Mcenareta Ekologia

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W alter Larcher. "Physiological Plant E cology Ecophysiology and Stress
Physiology o f Functional G ro u p s", Springer-Verlag Berlin Heidelberg
New Y ork, 1995, 494 p.
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d Q313" (10 -1 )
c (336^0 (lO-2 )
m 9 n0 n ( 10 - 3 )
9 nihm (10 * )
n 616m (10 - 9 )
1 J = 1 N • m = 1 i& ' 92 • 69-2 = 1 3^) • 69 = 0.239 i i c = TO7 3h6n

1 W • h = 3.6 in 0m3i^n • 60 = 3.6 yiQrrvsmxn = 0.86 in0m ii0mhn>
1 MJ = 0.278 yi0m3i^bs:>cnn
1 j>n0fnib0nhni = 1.163 3^ • ben
6633s
1 MPa = 106Pa = 10 bihn

1 6ihn = 1 0SN9'2 = 105Pa = 100 J ^ ' 1 = 106 3I1& b3' 3
1 6ihn = 0.9869 ^ 9
1 ^ 0 = 1.0132 6ih
Byonb h3gn0n
(p [MPa] = 0.462 Tabs In = -1.06 Tabs log 10 ( 100/RH)

K20 = 310.7 lo g 10 ^ [MPa]
6y0nb qjihgmbncnn aw = pw/p s = 10' 2 RH [%]
Pw = Byonb mhcnjonb 66331 [Pa]
Ps = Byonb mhovjonb 6633nb fcijjhQ&i [Pa]
RH = Si^hnb qjihgmbncnn j)36ni6m6i
1 osmol i i _1 = 0.00832 Tabs [MPa]
Byonb bSQ^nqjn^hn 9 ib i (p)
i6bmo3ib3hn bn9 v,3hn33 4 ®C -<b 3 = 0.9999 &h b9'3
i 6bm03^3hn bn9 i 3hn33 20^ 0 ^ 3 = 0.9982 &h b9"3
6yonb û6b3(nh^n
63360b 6iiipn = 9m0n 60"1; 45 boi'1 ; 936 9'1

6iispn = 9 mon 9 "2 69"1; 93 9"2 69 "1 ; 9 69"1
- 10 -
3 6 0 ) 0 ^ 0 6 0 <56 W qffl3d60k 3 6 d 6 n 6 3 6 0
63360b 6si)sgnb bnRjsh] = 3 ben"1
X 3jtihmcf\yib {>i0v^>hos6r>6s
1 s = 1 Q-1
1 s a -1 = 10 as ba-1
1 inmhos ba-1 = 1 3S ba-1
im6o36^hi(3RS
1 ppm = 10"6 3mpn ampo'1; 1|I 5I1 ftlr1

1 ppm = 10** 3mpn ampn'1; 1(J. 6s6m6h ^h'1
6sbcns U3(j»
1 j)h COg-fcsbcns b3ps = 0.73 &h V2-5sbcns 53ps (RQ[C0 2/0 2]= 1)
1 &h 0 2-6sbou> 53PS = 1.38 5(1 CO^sbOTs 53PS
gnqj3bonb y’130305036^0 DCq, = 0.64 DHzq
go<33bnob 2)™30305036^0 DH2q = 1.56 DCCh
0.03%vo1C 0 2 = 300 [il I"1 = 282 |J.bar = 28~Pa C 0 2Sshôspjhn 6633s
1 |lp p-1 = 1-963 |X j>h CO2 p-1 (1013 3p6sh 6633:^3 gs0®C-b3)
1 ppm C0 2 = 1.83 a& a"3 = 41.6 |X ampn S'3 = O.IOI Pa (20°C-bo P> 101.3
ipPa 3s3hnb 6633^3)
1 02, COg 6h_1 (39hspn 6m6s) ben'1 = 6.3 p. 3mpn COg 35'® ed-'
1 0& C0 2 g0-2 bon'1 = 0.028 65 C0 2 a-2 69-’ = 0.63 |JL ampn COg 0"e 60-1
1 06 C0 2 ar2 69-1 = 36 06 C 0 2 ea-2 bar1 = 22.7 \l 3mpo c o 2 a*2 ea-1
1 |i ampn c o 2 a-2 ea-1 = 0.044 aa c o 2 a-2 ea-1 = 1.58 a& c o 2 ga-2 ben-1
1 06 H20 pa-2 ben'1 = 1.54 H 3mpn H20 fr2 63-1
5saly>fins6m&s
1 ampn a-2 63-1 = 0.024 33 63-1 (20°C-b3 gs 101.3 ipPa 3s3hob 6633^ 3)
1 3mpn H20 S’2 63-1 = 0.446 To/Tabs P/P0 ba 63-1
T0 = 273 K; Tabs = s6bmp3(53hn ttjaaQtaajfe [K]
P0 = 1010.3 ipPa; P = s^0mbg3hxn 6633s [ipPa]
qjnjyn0sbs
1 &h 03hspn 3sbs 3"2 = 10"2 a 3s"1

1 5h mh6s63pn 93hspn 3sbs ~ 0.42-0.51 &h C ~1.5-1.75b C0 2
1 5h C * 2.-2.2 5b mh6s6xn 33hspn 3sbs = 3.1-3.4 5hC0 2
1 5h C0 2 - 0.59-0.66 &h mh6s63pn 33hspn 3sbs~ 0 .27-0.305b C
- 11 -
hignsyrii
i j a-2 w a -2 = i w a -2 = 1.43 • 10-3 iiC . ba-2 gor1

1 w b3-2 601-1 = 698 J 9-2 63"1
1 y.0- b3 '2 = 0.04186 MJ a-8
1 Ompn <3fT)(5m6n = 1.8 • 105 J (X = 650 6 d)-gb 6 2.7 • 105 J (A = 450 60 )-3 g3
y>eay3a6i ( { ^ , 3036* ) W 3 "2 («D3ih) jCC3*>n (400-700 63)
fi 9m(v» oj<n^m6n 9 "2 69"' |i 9npn <grt£in6n 3"2 63"'

ge>nb bn5ioi03 (S brufn qjvj) 4.6 18
BC>nb bn6io>Q3 (&a&6 ;)3Qn brfiaoicj) 4.2 19
JuxnjjCjhn Bicn^hi 4.6 14
cnjcnhn 4.6 12
3ah3*h]dnb 6xnph> 5.0 20
McCree ( 1981 ) anb^ggncn
O3033hivE|ha
i 6bmc3lj)3hn ^ a ^ o h i^ h i Tabs: Kelvin [K]

qjihgmdnonn Celsius [°C]
gpnb biioi36nb b ^ iQ m ^3033halj 3h36n:
TD = 0.5(Tmax + Tmin) + ( T ^ - T ^ / S ? (Milthorpe & Moorby, 1979)
TD = 0.29 Tmin + 0.71 Tmax (Gouderiaan & Laar, 1994)
paanb b ia ^ c ^ TN = 0.25 (Tmax + 3Tmin)
3hon33Q3&nbi Qi 6ip iy 3i 6nb qjaj^mhQ&nb 93bsb36 n5gmhab(jni nb. 336935

3hn6359n: Sestak et al. (1971), Slavik (1974), Incol et al. (1977), Savage (1979), Hall
et al. (1993), Salisbuiy (1991; 1996), Nobel (1999).
- 12 -
__________________________________ 80066(500)6 baUbTOriObOÔ &)rt08n
i ai536:>h3aD b^hoarn
8(536ih3 93gi9o6i<b3 ihb36m6b cnnoijdnb y3305i6, 8101 8mhob b(p336bi 91

3 npi 6yi0bi^33369o; b83 C3cn'b3 8(336^3 39166^13600 9 1 yo6 x f i 3i>6 bn3h|j336<b3i3
i n 53639361. 0mh33Q 53mpm5n3h 6ihb3Q0o b03P3cnnb 3^3390 8u36ih336n
1(^8^1^69636 bi8 yihni9 n, hm83cn(j 6ycob, Si^hnbi 91 8oinb ji636obj>i6 83953-
6^91. 8icnn bnhb36m 51113801 i vj)8mbqj3hm, Snghmbq^hm 91 0ocnmbq33hm 51691.
8015301636001 03ih30o 0j336ih3xo bicghob ^338 8n>|hmmh5i6o!b836obi 91
(3bm3X36ob 001616093016001 8i536ih33dobioi3ob 13150936390 b^bb^hiô - 60191-
50 6ih8mnj86i.
1.1 ai336^hnbW
l5bn3h363Cnb^hoan
1.1.1 ^ a n b o jD h n
5Cm6i03hn ^ ih ^ n b y s x ^ D b36bn(jQho 6160901 i^Soibqôi - 3i3hnb 016390
[mbar] 10'4 103 1 0 * 1 0 1 10° 101 102 103

100- ^— * - ----- j------- 1--------1—----- j--------
\
90 (
Mesopause \
80
■ ......V ........... \
■g-
|
70
60
50
Stratopause
\
................. 1 ...
\
\
\
\
..............\ ..............................
/ \
| 40 \
/
30 Temperature Ozone Atmospheric pressure
I / \
{ \
Tropopause / [mg 0 3 • g'1] Mountains . , V
5 10 / A , i Plants \
. “t v
■100 0 1001°CJ
6ib . 1.1 i^3mb<33hfnb (H ackel, 1990)
- 13 -
9 (3056600)6 &6 6 0 8 D
gphbn, hm03Q0n(3 5 sb 33X ni e 3ga9 n6 i (6sb. 1. 1). Q3gs3n6nb Ss^hnb j,shbn 01536s-
ho^bb s0shsj>36b 6 sb3 nhmhgi6 j)ncni gs gs6j)6sgncn gs 9cns5cnjs3b smhoij036nbsb
Bsh0 mj96 n0 n 6ypnb mhcrrjph. 6nmbqj3hm9n 9 cnQns6sg gs 2)3hdmg s^mbq^hmSn,
3nghmbqj3hm9n gs 0noimbq)3hm9n bnmOsbnb j,s6s6n036nb bmj>sgn 3s6m6bm0n3-
h3&s6n 0 mi5303pns 6sb. 1.2.
IsQhnb 6s636y33(t)30n 0mdhsm6s 9 nbn 930sgj>360m6nb 03p0n3m6sb 3 ^ 3 6 3 3 -
pymqjb. SsQhn 78 0 m(530m^ncnn % sbm^nb, 21 0 fT'l5X rn^ncnn % gs6j,6sgnb, 0.035
0 mi530m6 noin % 6sb9nhmhgs65nb, ^>{,1130133 050(13 hsmg36m6noi 6ypnb mhcnjQnbs
gs i3cnn09m6npn j)s'b36nb&s6 03g&36s. &shgs s0nbs, SsQhn 93nys3b ^sbnbOsj^sh
gs 0ysh g>s0s^3^yns63d30 0n6sh3336b, 33 nh3303b ym3pnbs f,mj,nhgnb mhgs6&b,
sjhnQsg sbm(t)ri3s6 6fi30ifi3h3636b, 3s0mj,3636b, 0^33hbs gs ^3sh(j0b.
0(536sh336n 3b3&ns6 s^Smbcr^hmb J33gs cg36sb - ^hpOmbq^hmb (^SQhnb 6sh-
bnb "s0n6gnb ‘bm6sb“)- oi3ncn 0mbsb(p3h3 hsnm6369n 5s6bb3s33&30n 03nd03&s
nymb s0n6gnb shs 0bm0mg bs60m$03 0^30363611 (3(5s63gn 63n0s, ^DJs-j^bngn,
jshnb gsjhmps), shs03g 303(1™ bs6j)hdpn3n 0g5m0sh3m6s (E>3n0nb s6 yn63nb 33hn-
mg36n), s&h3oi33 ignOs^nb bsbnscnn (bs03s0m 0g&m0sh3m6s gs s0s 013 n0 sg&n-
0sb s0n6gnb ^333036(1030 h3gn0n). sgj,npnb cns3nb363h3&nb, 0u36s(i3XfT'6nb 0 3y
h30m6nb, bn0s(ppnbs gs ^nSnb 93bs6s0nbsg 2,s6bb3s33&30n qgshcnm6nb ^hn^m-
hn3&b3 6s6bsj>3oih363Qn ipn3sl$3hn 3nhm636n ys0n6g36s. 3hoin gs n0s33 0sjjhm-
2)0n0s^nb ^hn^mhnnb Ggsh60369n 5s0mnymq)s 5s6bsi3cnh3&30n 0n],hfTij)0n0s^nb
0n6si33Oi36n; 3b 03nd036s nymb, 3s{js0ncnsg, hm030n03 q)3hgm6nb, 0i536sh330n
ou>6sbsbm5sgm36nb 0nj)hmj)0n0sljin, sj>h3ai33 3s3hnb bsbs'b[p3hm 9h33&nb 0nj)hm-
iCn0son 6nsgsj,nb s6 cgmcnQ36nb sbprnb. s0j)3shsg, 0i536shnb 6s6n036'b3, hm0-
BIOSPHEiRE
6sb. 1.2 bb3igibb3i jnSn^hn x o 9 o6>in^

(s^m9n/9ncnm5 J i 6i 5-
riQj&i bnmbqj^hmSn, i^mbgohmSn, 3nghmb-
gjjhnbi 9* QncnmbojQhmSn. rnhgnBs^nb ipjh-
dn 0fn53hnoi0xni. a s js c 001^ . H, O gi N
6sh9m9536npn> y s X ^ D 9090 3hm3mhi}nncn:
4.98 X 105 ym3X 9 ncnm6 3.
n. 3scnn 93qjsh93&s 93^95360 y 3 X 5 »iim9nb
9ssbpm36nOT 50% (Deevey, 1970)
- 14 -
boobn3(n35aô JW ftoan
Qjftnu 5i60i5o6xn> (pn^ t» 3^Dhm bnû^dn. 6i3C3bib ibg^Sb 0fbnb h*gru(jnnb,

(j>3933h*03hnbI ^36n*6m6nb* Qi jahnb gngn 15300036360; *0*6 9^36ih3b "bm633h
03nd03bi 060936300)36^9 * 36mb.
1 .1 .2 3 f ie h n b < 3 D h P l
Snghmbq^hmb 0032)3013636*: 001300 3'b*h9s'bihn 9bmqj0nm <^213163 (hm03cbau

Q39*9n6nb (b3Q*3nhnb 74 % 3^nhs3b), &*9gn6ih3 g* Qgg^hn 9 ng;> 6 y*obi(i)333^n
(0Qn6ih336n, ^6360) Qi, 6mcfTib, 5h36»jnb 6yc3&n. 6y*0n 0mn3m33&i io 3 fT,b<33 -
hm9 nu (*(1360360^ 6nb0nb* 9* Bycnb mhoijcnb b*bnoi. abmqjcnm m213*63 - Bycnb
3b fc n ^ ^ h n *hnb 363h6nnb* 9* 6n3ain3h3636nb 5* 63 b*'bc’3h x i e en-
gn h3t)3h33*hn g*, *0*33 ghmb, 53mqjn'bn>)3hn 9* 63mjn0n3hn 3hm(33b36nb 0*b^>
6non'bnh363cn qpjomhn.
pH
Bab. 13 6ib3nh6i e x ° bnb&j93&nb pH Ijpsnba (j6y3>jn bi'bn) ga 3,u6ah (D^Sj^nhn) 6yoo63n.
eaS^hnbxn ihj: pH-nb baSâcm 9n6i(jo93&n 'b(«3nb 5yac3n (Ott, 1988). mhgn6alyib pjWba
6aR336o6n> COi-nb pa CO3-" -nb 3hm()36|jn.
(jbhnco 1.1 6yac9n C02-nba ga C^-nb bb6apm£>a (|iM) ga 0mux,r* no,n i<ri6(J06(>hioni (%)
bb3agabb3> ^3933ha1>]h»'b> 3a]hnb 3ah(jnac3hn 6633a: 35 PaC 0 2, 21 kPa 0 2 ( estak et al., 1971).
6ab3nh6agn pnmjbmngo 5s656agn gnmjbngn 2IC0 2

(CO) (PM) (%) (|iM) (%)
0 26 0.059 458 1.02 17.6

10 18 0.041 356 0.79 19.8
15 15 0.035 318 0.72 21.2
20 13 0.030 291 0.66 22.4
25 11 0.026 263 0.59 23.9
30 9 0.023 245 0.55 27.2
- 15 -
9B3G6(f)00)6 <y>rt09P)
<b(j>3nbi g i 9 lj)j)6 ihn 5yc3&n jn9 n^hn 330i06360m&ncn 3hcn9i63ainbjp6 di0ni6

&:>6bb3i3936:> [3 ]: bp^nb 6 yi 00n 9 ihn036 nb bi 33 i 0 m 930i5330m6 i 35 £>h/0-<b3*;
nb Sgngihna Na+, Mg2+, Cl" gi SC>42' nm636ncn; bd 9bhn3 ihb^mbb igfcnombhnsn
5^6^3^33636(115. 0lj*,6bhn By036n, hn^mhii 63bn, gngn himg36m6noi 93n(5i336
Ca21 gs HCO3- [ 142 ].
6yi00n 5it36n b bb6igm6i g i S m i n g ^ x ^ ii3933hi^^h>bi g i 6 6 3 3 ^ 3 . ^35-
^Dhi^hnb giB33iborii6 g i 66330b bhgibcrii6 3hcnig bb6igm6i 9 i^ 30 m6 b g i ^03-
630m6b 9ihn036nb im6|536lj,hii5nnb bhgnb j,3iom6ib3 ((5bh. 1.1). oii3nb3<3i0n
6ib3nhmh3i6j,nb himg36m6i Qi9 ming36 x n i 6ygnb pH-t>3: nj,n 9i(pi0ni 63^33
360-
340- 18- O
ra
320 JD \ -'V ' 16-
300- \/ ,
O V F p(JU? 14-
280 O
12-
260 /
18 22 02 M AM J J S O
Time of day Month
Bib. 1.4a,b bb;peibb3i 6yc3&9n 3bb6sQn ih im h j^ x n 6ib9nh6igftb (D1C) pa ipB^baenb 4^61536-

(Ondok et al., 1984; Pokomy & Ondok. 1991; Williamson & Platt, 1991)
ih39n (6ib. 1 .3 ). b 3g i 3 nh30n 9 hnb By03&n 93n(5i336 Snghmbc^hmb mh5i630n

6 ib 9 nh6 ignb 0.2% . gi656ignb 3ih^nb03 hn 6633i ^giSnhgg 6 y03 &9 n bi03i0mg
n6°33i. hi(5 6yi0oii6 i^mbq^hmb 0rnbib(p3h3 q)36i3n (^jbh. 1 .1 ). 6 ib9 nh-
mhgi65nbi g i gi6j,6ignb 339u330n 6 i 6y0nb b 3g i ga y330sb3 3^301 5i6ioi3630
033636^ tbm6i) ni53036i gpnbi ga b3<bm6nb 6 i 63 i 30 Pi6 i 9 n. i 9
I530np36nb 0nb3bni 6y0nb grnjjr^^rn^hmqjxn mhj)i6n<b036nb ij^n3m6i (6ib.
1 .4 ). Snghrnbcgphmbi g i i^3mbq33hrrib 3mbi‘b(j'3h3 qj36363n gi6j,6ignbi g i 6ib-
3nhmh3i6j,nb ij^njhn (330i 9n9gn6ih3m6b.
6y0nb 9(1636360 0nb ( ^ ^ h i ^ h i b ioii6i6h36b. nj, bigx3 ib3<7>n gn63&36n
ihii, 6y0nb b 3 g i crj3636nb 3n3h 3bnb higni^nnb dgn3hn 3 cn;>6 orj3 nb 83935^9
6ih0nnj0636i dQS^hi^hnbi gi bn0^3hn3nb j,hign36^n. 3b j>n gng {^ 3 9 36 ^
ibg36b Bygnb mhj,i6n'b036nb ^ 3 3 6 ^ 3 , 3hmg3jljn30n 6ib i g i 6 ^ 1 5 3 0 3 6 ^ 3 .
1 .1 .3 cnonnb<33hn e:> 6n:ie:>dn
93gi3n6nb fohjn ihnb 38113^1 bii3^6im 0 hi 3i 0 n jn0 n3hn xD^D^^nbi, hm3 -

C3&f»l3 mh6i6n'bd36b bb33Qnb 6r»3om3h36ioii ibi53& ig 3bi^nhnvj6i (nb.
- 16 -
0(3066630)6 b6(3bD36050c?C) &6608D
Bib. 1.2). pnaimbqj3hm 6n3cnn3h3&3i>b D(33Cn^ SnghmbqQhmb, ifthQcr^ 5i0m3y-

mqjb jp’bodb gi bb3i 6n3cnn3h3&36b (0ift., 3XJ>i6nb i0mq}hj333nbib) i^0mbq)3-
hm3 n, 6i&hi0, 33nh3303b ym3pnbb, pncnmbqjQhm biBynbn 0ibi0ii 6nigi5nb 6ih-
3nbij063Qig.
6nigi&n ih ihnb 9bnpmg pncnnbqjphmb t>3g i 3n h x r* ftif^bsn^hQbxn <J336i: nb
3n6QhiC3hn gi mhfci6;)cn 6n3cnn3h3&3&nb 5 ihgiJ 36 nbi pi J^h^nb Shm g^ni.
6nigi5n ijnnjbipn mh6i6n'b03&nb 3m6i6np3m6nCTi 6ih3mnj3636i g i 3 nbn Ri3 myi-
pnbyii jph^Qm Snhm&^b'bji gi0mj)ng3&xn; Bnigi^n &i636y33ljpn3 n^3Q36i: n'bh-
g3&i, SmSBncgg^bi, S^^dpni gib^hg^b g i jjiBigjj^hg^b. qjn'bn^hn g i jnOrvjhn j>i-
Smqjn^nbib Scnnb g]giji6n gi6 ^636933^003 5i0moii3nb3q}0g3&i 0n63hi03hn
6n3cnn3h3636n, i0i3Q ghmb nbh663&i 0 (j36ih 3x n Qi ubm3303hn 6ihRQ636n gi
tjbhncn 1.2 Bnbgi^Sn nmBjbnb 9o9mt53Q3 (JelEy, 1987; Kuntze et al., 1988)
an9m8i53c x ° V)g:>3nhn6 633(500303110 ^30101*163606 (530°^

oj3hcnn (d2^ -1) 363hn ( i i ' 3*30330.)
Al 93 Fe 6y3c9n 25-40 0.03-0.05
53666000 mj}09360
jj3RQn6n^n 10-20 0 .03-0.15
n0o(jo 100-300 0.2-0.5
33h9ni3cn^n 600-800 1-2 (8)
9m6cn9mhncri6 non 700-1200 0.8-1.2 (10)
33836n6 6366036(503 800 1.5-5
Time (relative units) Half-life of detritus Id]

6 ib . 15a,b g^hn^nb g39cnb ghm. a bsqpBnb bb33g 36b 33 m hîB xn im 93m636o 36o6 93(533606
ghm (R3i39i6, 1976). b g30po6 6365^ 0031*163. A 9nihm 6cpc0(536ih 336n, B 6i ii 6ihn Bypob 8(5363-
113360, C ^ m 6nb 9(j35ih 336n, D 9i,i)hm6yic 9(j36ih 336n, E bf^jnb 6(5363113360, F 9^ ( 53( ^ 136360, G
nbc36n, H 036&h3(yi ^ynb 8(5363113360, I q}moiiTi09(5336o 633606 qjmonp36o, J 63^3606 bsqj36o, K
6063^1363606 6063360, L 931139863363 633606 C9Tio<c36n. 8330 6311,50036001 3060863003 b3333cm
9m6i(53936n (Enriquez et al., 1993)
- 17 -
aoaGcwfiooi^ a ^ a a n
g*boi(5o0o 6o*g*5ob oili5*6o'b336o. bh66ob 3(10193^360 (560133001* 3d^J)h393&-

03601*6 3(101*9 01*69*01*6016001 5*hg*o;j3636i 33606013*6 6*3hcn36*g, hm0p3&nu
5*amognjj3nb 0063(1*03(1 6030103(1363601*6 3hoi*g j 06 ob 2>o'9^03jb36b. 6 o*g*j,3 n
y*0o6g3&* 3hmq3O0O 3mhn‘bm6 (jO&nb*1 (1019300(5 S^S^bnb* g* @3^hnonb (g^hn^n
- 6n*g*{,nb *b3g*3oh*b3 9*0336000 J3h 5 * 3 ^ 6 6 3 0 0 3(J36*h330n g* (56013303^
6*11636360; 0cn*h&9630nb 3360336*) 8 (1 3 3 6 ^ g* 5*8oiqjno3ob 0063(1*03(1 3mhn-
<bni6l!)b Soihnb 33606013*6 6*3(10101* 5*6 ^ 33*336 30 0 830153300160b *boi636b 830-
(5*3b (ubh. 1.2 ; 6*b. 1.5 ).
6n*g*£,nb 00^360 i)eo0*ooon, 8(536^330(1 b*q}*hncn, 6o*g*5ob 01(15*60*6836001,
93Q*j*6ooi g * *g*8o*6ob *635*3036001** 5*630(101636300. 9330* 3*oi5*6nboi3nb
5*hj)3330n 3hoicgn0n* 9*8*6*^1*0136300. 6o*g*536ob 6*h3 mj36*, 3*oin *5363036*
g* i0*bnqjni*(jn* *hnb 60*g*58(5«ig630i&0b b*5*6o. *3 33ij6n3h36nb b*qg3d3036nb
Ufig6s *3(3(1036300* y33C*bo<3nb, 306(5 (5900016b &*3hi33b 8(536^301* 32)0100150-
ob b*2)Oonb368o.
6o*g*5ob 8y*ho 6*600*^360 02)3(136* 2)018^36*9, 110163001* 8oihnb 83*030360
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59590560, (10150111(5 501603110 5116365, 0153060 2>X 03 ^ X ft 65630cn5h36o6 6 y50oi6 ocn
01(1556300 6001(53601^)3606 3o6hoi 65'b(?3h3&0956 6i90i309i 9i 9o i x o 60016033(1-
016 330193(5006 96o93630oi3i6 qjijôihig oj(ji. 59590560, (101501(1(5 6363606 835-
6 3 6 x 0 95 9o‘b5666h5qjxn 5ih959j9630O, 5(15633303611030 3<33i03hoi6ooi 3h33i
6363606 3hoi(53632)0o 9 i j 96 o6 5659, 56oihoi3oi6363h 60010013366. 0039155 9 i 6 cni-
3060 656652)3Oih3630 O 9 g 50 i95h3m65 03 odo35i 6 oihoioiQi(5 5i8oioy36oi6: 5(163-
- 21 -
00066600)6 a6A39P)
6m6b 6503)10163, hm9 59500560 gs5h(j>333b5 g i 55565g53h36b 63360 3956360b

y3395 6 nm^m3 b, 013 9 obn 1501965 “6015590 3S38oh36nb 555365893 ih 9ooy356b gi
013 8nbn 636b 35636069536901606 5hd6oi6i6 ih 953895)1365.
i .i . 4 . i 6 nn 6 ornjnan3hn 3hni50bD6 n hfTbnb°33hnan
^3)1^901356 8(5365)13015 0)363015 5)1380 (hn'bnbqQhm) 0099065)130166 6030103)1365015

lj3pnb iiyijhn 3)101(53636(1, hm0300fnj 80165609301636, 3homb 0bhn3, 8(5365)13, 0nbn
0336306 bibncn pi, 03mh3b 0bhn3, 60595506 mhj)i6n'b036n: 69590(5365)13360, 601301360,
iiyi6m0oi33^36n, 65^33)10360, (5601339360. 015365)13015 5i9b05hn g i 03395)10 65609360,
(1018936015 6o5g55'b3 153035 (6503360), 6635956635 95889393606 65835936001 556015906
0n63h59O(b5(3O56. 958906 3)10193^360 558010901035 55^3606 b5booi (C 0 2, CH4, NH3,
N2). 3301606^38580 9o63h59o'b5(5nn65 95 mh55630O b36b^>6(5036o6 3hmp3j(500b 3hm-
65b. 1.6 CCb-nb g i 02-nb yn 6(j;)6ah:>(jnnb

jhign^n omb6ih 6(1^0^5811 (6nigi5nb bnph9o
10 b9 ; 12®C) (Ohga & Ikushima,
1970)
^360 3)106563010156 8^nghoig 5hob 3(10103)101953^380113&390. 013 36 3)101^360

93733^636959 9 5 66)150359 6009065)130166, 65806 8(5365)13360 35)1559 65)155936056
65^3360 6030103)13636001 95 05010 600185606 955)1013365(5 6mh0593h5g 8089065)130166.
01330 8(5365)13390 65033606 958906 3)10x536360 h5o93 0o(b3<bo6 550m 83033116363905
95 3383606 65h9mj95(5 6390 ^ S o o i 9o9go65h3m66, 50 836016333580 913365b3015 'bhgiu
8 3 6 3 9 ( 6 5 3 3 3 6 0 603010311363606 6o86ohn6 558m) 95 3301606^39580 60018565
9(3nh3 h5mg36m5ooi 5)10139365. 65033606 8063)159^65(5006 b n fij^ 959013^363905
60595506 803110103901)106 b5b3m6ho3 0385g5369m65,b3, 53h5(305'b3, li3833h5vj3)i5'b3, bo-
5fTW133't>3. pH-(b3 95 956589390 6366^56(5006 8305gj)369m65'b3. 8o63h59ob5(50o6 606-
j ^>3 05^5905 601^001 6nsg5{j8o 630,^1593)10 pH-ob 30)10163680 95, (101501)1(5 6360, gs6>
905 ^5116^36056 60595580, 55h30133 83533 60595580, hn0398o(5 359(303806 830(33390165
- 22 -
BiM66 ff)0 cna cWfnoan
Scorzonera villosa Trifolium trichocephalum

Allorhiflc roots
Polygonum bistorta Aegopodium podagraria

Homortifcfc roots
6*b. 1 .7 <33t>3au> bnb^^dnb dnhno^gn ^ 3 6 0 (Kutschera & Lichtenegger, 1992)
0Unh^. 6nbgb6nb <y>0m9hrn6:> qs 9nbn ^ ^ b ^ o b 4 -5°C-cbo ^333001 9^633^ 06333b

6niQi5nb fnh^bSn'baQbnb iiyismfenb 9 ^ 30 0 136 ^ q x d jo & x irb se, dc&$&c
3hrxj3b3bnb 9o'bcl3Q3ib. (jhrnioix 53n9ru>5 (jy33&0o ^'bihQi'bbhn 6nmchbnb 9^1*133-
6nb ^hoKjhom 06o93630m3s6o QnbgVi b6mhx 9o63hs0n°QslSonb bnfijhhî, h>tau b3pb
36ym6b 60^9^50^ ^303311^ ^ 06^ g i bnCSm^nb m^v^nQ^Q^hn 0^ 336360360.
60^9^3680, hm003&0(X5 qjmfvj&ob 8mux»T'&* Qljrtv^ 6i't>3£,nlj 0o0mux^ 2>*d6x^
- 23 -
8 0 0 66630 to <W(n3 an
Tap root system Heart root system Sinker root system
Abies alba Larix decidua Picea abies

6*b . 1.8 'bmSn^h 63^ 536* 11301* <33 b 30i* bnb^30nb ,503360. A bies alba-b * j 3b
5 * 63,5^1360010 <3 3 b 30 i5 bnb^,33i; Larix decidua-b - 5 x ° * J06hn; Picea abies-b - 6n ig ij,a n 33(1,505*03-
h*g fi* 93363^0 (K o stle r et al., 1968). ^ h m io ix i5y 3063n g i 3*65659001 (pshn6 60* 9* 536^3 63-3(5361-
h336o 03001*113636 mRmqj3 bob65535h, d(vi3h 5 * 6 ^ 1^,300-95^3000 91 biyh 936ob3553 *h <3363011 bnb-
,53836b (R ichards, 1979; Longman & Jenik, 1987)
630*15 (65b. 1.6). ohm3njj]{)9 n j)5h&5g 5580165(1 5059553680 05p 5on 6360*^30010 5 ^ 3 ^ -
5ob 550m, 35656590b 3301333(^165 81501135. dpo3h 9501)33800 56 £>(16036056 60595530,
55h30133 yo63Qnoi g50)5h x 60595580 55^015 3n3m(j305 0093659 83033116363005, hm0
5056 3300)0365 550mo63nmb 8(5365(13300 b5qphob i 3gm05U yi. 6059550b qjm(i363o
65b3ohmfvj565ob yn6^36oh5(5n5 (15393633(103 5(^3850365 353(180 3 ob yi6(j36oh5(yu>b.
60595580 C02-nb h5m936m55 950my^rj6 x a5 6059550b mh556ob0oi5 95 0^365(13015
6om0m5o3h 5jon3m&5b3, 6059550b b o li^ jo ^ ^ 9s 3°^ ° ^5(3006 01530b363(13b5b3,
335305(joob 33hnmpb3. 55b3&ob ^1615360(15(505 6059550b bb3595bb35 3mhobm6 o3o 556-
bb3533630O5.
6059550b ‘639530(10956 5o3mbqj3hm3n C 0 2-ob 553mymqjnb 3hm(53bb 6059550b
^3601^35 36009365. ob, 55(130133, 9095955 953mi09363Q0 65b3nhmh£)565ob 35(1150593(10
66330b 5h59036ob3 95 6059550b b3053ohb3 ^53(106 (3360b 0mdh5m65b3. 6059550b
b36aij35b3 dj^rjh b3553036sb 56936b 03933h503h5, bm0m 3hcnb5 95 08533 039^3h>
Ojhnb 3ohm6368o - 6059550b 036n56m65. bm3o3h b5hoy3C)3 o oynb 6059550b 6360*^306
- 24 -
annfi.^fnaoia b m n a fra fra ^ o &>ftaan
20 -
40 -
Trifolium repens
60 -
Urticaurens ' Calluna vulgaris
6*b. 1.9 6n:>gs&nb 6vjp36:> <33 b 3Cis j>i6 :>6np3&:>V): Trifolium repcns j3n96sh-oinb6ih 6 fiig ii,3 n (3 ih -
ljb36:> bjh^cnn) g i cnnb6ih3n (3ih53]6i bjhioin); Urtica urens cnnb6 ih 6(u g ^3n , bsgs(j i'bfn^noi
3gng>hn Vjpi 3h;)»; Calluna vulgaris mh^s6xn 6030103113636001 dgng^h 6ro@:>59n fb 3ga bjhôrm -
^hnp3n. J33g> b3hsom - 33KyiiiC3h ShmjJyijSn) (Kutschera, 1960; Kutschera & Lichtenegger, 1992)
o6£j66o3m6:> 0 .1-0.5 5b C 0 2 S^bor1 9319536b, bmpcn 8gxfib 6nî5r>bi - 9^1b-

00136001 1 fch C 0 2 S^bor1. ^hm3n sx n iMf y f o h 60^9^50b b^6o<J3nb
336369360 ( 1-15 5h CO2 9 ‘2bar’ ) i(pnh(\5b36i 6pn6 636016 33hnmp3&9n. 60191506
63601^306 ] 0ijnh]bn 3^336360360 (0.050.2 C 0 2 d^boi'1) gi 9ibibrcior)6x ni
j3o36 iho 91 ^36ghnb 60^9^53606^0*3^ [225 ].
6 ru(u&3 n 6030*1311361011 m e* 9|j36ih3b, 0ot(hoirnh6i6o'b336bs pi ihimh6i6x i f ® -
3m636^36b Smhrtb <0 3 0 ^ 3 06^36603(119 q)3b3oc> 1(1330, iphdmg 0 ^ 3 0 6 bjgbinhV)
0o0pn6ih3n6b. 0336380 6306013361 6tppo 91 9 sb9 o 5bb66oQO 6030103(136160, boipoi
ho(bm93h930o 35(139360916 5idmoynvgi 0)36306 5i9m6iyoiq)3&o gi 916906 Shmgjj^j-
60. 09363^6 qjihomb n6p3Jbn (RAI) 01651)109361 hm^mhy qj3b3ob qjiho¥n6o (9 ^) 60-
- 25 -
90366(5)3016 &6A 39P)
ARBUSCULAR ECTOMYCOHRHIZA
MYCORRHIZA
Gymnospermae
Vascular plants Angiospermae
and Bryophyta Basidiomyc&tes
Zygomycetes Ascomycetes
Zygomycetes
MYCORRHIZA
IN ORCHIDS
monotropoid type
Basidiomycetes Basidiomycetes
encoia type arbutoid type

Ascomycetes Basidiomycetes
MYCORRHIZA IN ERICALES
6sb. 1.10 Qn^mhrfbib bb 3ig ib b 3i qjmh8nb bj39i^3hn 5 i3 m b ib x 3 ^ i (Gianinazzi & Gianinnazi

Pearson, 1988)
Alnus incan a
C=) Betula pendula
[...I Quercus rubra
^ 3 Larix decidua
70 - 1 Pinus nigra
Pinus sylvestris
60 -
50 -
40 -
• 30 -
& 20-
%
5
6 ib . 1.11 bb3igjbb3i bnb o j^ n b un^bico
Snimhobnb (%) nfjg^b^nxn 9^33
moderate high extreme
Pollution load pa0»&n6d3lv)&CD&rib 5130361 (Kowalski, 1987)
SQStfk qjihonb j h o i g x ^ (S2)- ^3> 3sj)i(}nau)Q, "br^rrjh bih^jyxSn 3b 3^336 36 30 0

qjmcnc»ny>6n 63360601306 0.6-2.4 qphj^bQoi, bmpm 60631^136360601306 -
0.050 .1; Sjjjhx 6063013^6 (jy39o RAI Itkib (jmoni, bânshng^ <bm6ob 63^^6^(13690
- 26 -
3036 . ^ 30)6 a ^ sa ri
yi 2-4-ru. [ 104, 125]. 0£)b3nb 653^ X 3^5 g5 bnbSnh], 3nh3x yro3onb5, gbQfnyig^bj-
pni 50s or) n9 bib^robnb q£foyn> bob£)8nb Qmhognicmyi^h crttyib3&3h3&36‘b3 (nb.
qj-jojcrb (yiS^bn: 65b. 1.7 gi 1.8 ) gi jihnh^bb 60nb 5568530016590: n‘bhg365 gpWj)-
bx^D & 6308056 b3<bm69 n g i 8(5ohg365 3D63i!)S(jnnb ^Qhnmgnb 6m0mb. 6n5g 55nb
<71330636^36^ (pH, 8n63h503h 6030103}!365015 g i 6y0ob 338(3330m6i, b3hi(ini,
6050550b bnc+103, 6059551^ 3mhn'bn6li 36n gi bb3.) dpn^h 860936301*13560
6ni, (10180360(3 556b5<bc'3hi336 0^ 30 15 j^sh^xDîb (65b. 1.9 ).
bSnhig £^001356 QbDîhQOU) <33b336'b3 bibcgpbi 8 oynhrfb30n bmyn^&n g i ig&n-
pn ij3b bnSbnmb^h ^hcm^hcnm&ib. bmyn^bn ibj^h g i ioiibj^h lbhgni6 033630^ 936-
6013 <b3gi3nhb. 8|336ih3aii gngn 30hi303lxn6i 9ib3n6d3pni ib3cnn 0 ^ 30 15
6nbi. u6mi)noni 9n»)mbn05nn\i)hm3nnb 0h5350n qjmh9 i (6ib. 1.10). y a x ^ ^ qjihcnmg
553^30363005 8oj)mhn<bob 30tin])X3h-ih63biXDhn (VA) tyâ 050 63b3g3&i 9 hi3iQ
83(1^6056 gi 6505bm3i6 8(5365(1380, 55(130133 5303(1565 g i bi3b9n [208 ]. Snynhn'bnb
ihb36m6nbib 3dnd3 83050360b 93cn3nb36i n'bhg3&i gi 95b3o6d30o 8(5365113 33301
85(1550365, dijpcnoiig, (1306001, 0138(55 3b b3gb jBymbb ^ 3 0 1 3 3 o m jw ^ o 6030103-
h36360b (855., 35980380, ^y3ni) 83013^^36565(5. 03mh3b 8bhn3, Soynho'bx 60131^1366
83663301 36ihn 35h3gnb 3390^6 yn33p3^368n 055656 8dn93 9^ 9 ;)5bh30oi6
3(101535(10 958(53300 63cg3hnb 0336^5. qj3b3nbi g i 601301360b bo36nm'bn 63660^3(105
33533 5i3ri6iymq}36nb 9n3 ihcn. ib3, 3555000150, 60630136360 35(155336 ou>3n56oi 80301-
600160366 S 0 2-rtb 'b33niJ83g36nbib (65b. 1.11 ). 5800018 3nynhn‘b3pn UT1301360
gi6n6d3h3?)3Cn 605955360b 35(150 069035001(13605.
1 .1 .4 .2 6nTOji!)n3hn ai3 3 6 ^ h D X n 6n3(JinDhD6D6n
8(5365(13360 6ih3mj36ni6 8115350 6n3cnn3h36ib, (10180360(5 5530365b ibg3636

bb3i 315365(1336^3, 8ni)hmmh6i6n'b336'b3 g i ^(*133036^3. i8 bibnb 6o3cno3h3-
636b 3601936501 33016016360 i6 n6a)nijn0n35o;)£>n [45 ]. 3ynbnb,i38i8n nbnSn
860936300^6 6560565001, 5585^3330, g i 3|J330 9^ 9bi336nb 3n3ihcn n63n5n-
001(130 hmpb 5^303636. 6ooi5;)on3ho 015365(13300 60301031136560 65(18015953636
8301(1590 03l356oipn‘b8nb bibmgmn 3hoig3jo 3&b, 580^018 bb3i&3ihig 8501 8301(15-
gn 8(5365113300 6030103(136560 3601936501. ^01503(1010 ib3cnn 6n3cnn3h36nb 83056-
mpn'b8nb ^385 6ifi33636oi 6ib. 1 . 12 . i8 3pibnb 6o30i03h36ioii 8n8nibn03i 801-
(3383005 (3bh. 1.3 . 33mhig 8(336ih330 6n3CJin3h36ioii gngn 8hi3i0cg3hm363&i &i6-
3nhm&3630ni 33rn03(3nnb 3hm(33b3 n 3(336ih3cni ijo n3hn i i 39nh3&noi 9 nj,hn630 3i-
hibnoo&cnie g i 6i6bii3aih36ncn 2,m6b3036(irl3bfTi3OP36oii6. 3d 3303bn 958(33300 jn-
8n3hn 6n3cnn3h3636ni (6ib. 1. 13) ihib 33(3nq3nj,3hn Oj36n03hm3i 6nb 6i3hoi3&n
(h3‘bn6n, 0nj)6n6n; 6ib. 1.14 ), g36mpnb 6ihfi363&n (t,m6g 36bnh36x o 0^606360)
g i qj053m6mng36n.
n6cgrrijn8o3h 6 n3oin3h365cni h5mg36m65 5i 6 bb3i 33b x n i 3(536ihob bb3igibb3i
mhj,i6 n 9 n, Jbm3n0bi g i cn3ncn 33^13990(5 j,n (b3 j)h30 fnhxn 303836^360, o^nB-
3ho ngnnbpiboobn). 8icnn himg36m65 (3350365gm6 b m6 oPi6063'bob 856do0 'b3 g 5
6 0 3 6 0 b 5 5 6 8 5 3 0 ^ 16 5 9 0 . 8 6 0 3 3 6 3 0 ( ^ 3 5 6 0 6 56 b b 3533 b 3 b o 5 (p8 m fi 36 o 0 o 5 3honn g(j>nb 5
05 b 55o io b 856 d o 0 <b 3 (3. 59 6 n 3 0 in 3 h 3650 i i 8 h 5350 q j 3 hm 3 6365 b 5 g 5 h 5R g 3 6 m 65 (b 3
3 6 0 3 3 6 3 0 ^ 1 3 6 5 0 8 fn J 8 3 g 3 6 b 9 n 6 3 h 503 hn 3 3 3 6 5 g 5 b o h 3 b - q j 5j(j<T'h 3 5 n b b 3 0 rnJ 0 3 g 3 -
65 ((3bh. 1.4 ). 8i550ooi5g, boh3b9o 0ymcgn 9i5365h33&n g5 5'bmoocn i5hj,5g 0 n 3 >
- 27 -
913066630)6 &6 f?»38 rJ
hi63& X n 9 l536ih 33&n 350113303^9 bgo6ni6 963 h]6nb 939 n ^ 33nb m&n3j^36n.
ga9a8ti33hfto636Q36nb dribnfg*; ‘bmmjmhni: 61536^301^ y3i3no36nbi gi 6iy-
mqj3&nb 93<33hnpm6i g i ihmQi^n ihnb 6 3 ^ 9 oajotohn biSîCD^^ 9lJ36ih3aii
9n3h 963h36nb, cghn633036nbi g i bb3i (jbm33036nb 9mt>ng3nbi gi9(i)33h3nb gi
6iyncgnb &i3h[53036nb 9 nb5 nai ('bmmjmhni). y3i3nQnb <33h5036nbi g i 6iymqjnb
cg3hb J,i 6 i 3nhm636 b 6i6n0m6hn3 6 nm9396 hi636 nb ag3hign 6i3hoi36nb 6*301360
(i>ihm^n6mn936n) g i 33h3gnb i 39309 n ihb3& xn 3 0 ^ 6 ^ 3 6 3 6 0 g i 9 x i 6 n6 n,
cn39|3i 39cni3h3bni 3jh3gnb 633680 3bb6ign 60nyn'bn036nb 939 u330 n cg3hign
6i3hoi36n. 'bmj^hain 9 tf36ih3x n 3nj,936 ^nb (qj0i3fT'6mng36n, 6 3 ^ 0 ^ 6 3 6 ^ î-
hm^n6mng36n) b ^h ^j^hi 9 mi539x ni 6ib. 1 .15 . 9i536ih 3cni 3nj,936 vi)36 nb <33113-
&nb ^higiuni p i 'bmj^hoin gi9i9^33hni63630n 9mu393oni (jbh. 1 .5 . y3i3n0nb
bb3igibb3i q^hb &i6bb3i33630n b33j^hnb g,i9 mbbn33&i ij3b, hibiy bb3igibb3i
963hn i(j>nj3i9b. q)hn633036n iihj>ig i(j>nj3i036 9£>b3 q)3h36b gi, i5h30i33
30i!)hinnbq33hn b33^hnb 5i9mbbn336ib. ihm9 i , j x n 6030^131136360^ 963hcni,
O3hn6330cni g i ybm330aii 9m'bng3nb 0 3 3 6 ^ :^ ibhx3&36. i 9 6n3cnn3h3636nb
b36n j,i6bij)3aih36nai 9dicghp3&:> n9 33hnmg9n, hmg3bi(j y3i3n0n 9 b ig ii j,i6i-
ymcr)n3h36nboi3nb g i hmg3biu 6iymq3n g i 96 nq}3630ni. ihm9 i ^ x n 6n3cnn3h3636nb
Carbohydrates
Nudeotides
Amino add add

metabolism )dism
PhosphoenoJpyruvaie
Acetyl-CoA
Shikknicacid
,COOH
Malonyi-CoA
Acotaiepolyketide
pathway
Isoperctenyipyrophosphate
fob. 1.12 0\S36ihnb 9omhig 8> ji 6ri(vi'bd9 n fcJicnxn 00601313^(1 aVj&nb ‘bmjppn (Schlee,
1992)
- 28 -
ati0 G6 fn00)6 tWôan
UbhnQO 13 6nmiiyi]hn 9>nhign 9u36ih33cn Shrnp^jftnb 9Db30ghnpoi6i gi 3qolyi (Rundel, 1978;

Harbome, 1988; Crawford, 1989; Luning, 1990; Henssen & Jahns, 1991; Schlee, 1992; Bennett &
Wallsgrove, 1994)
jCsbn 33b33ehnpm6 i 3° » iS "

^3h336mngo6n
9m5m^3h5] 636n b9nh3g. dnhnoi3g 1330136911 ga9 btobnicn3b3cn bj&i @0 539m
i 55nmb33h0369n, 536l» 430ih36noi 863(13 g i ^ m jb n ^ o

b3b i 3n^3h3] 636n
AsteraceaeSn, XtyrrySdn g i t^txfidn
gn^3h335mng36n bSnhsg, h3bn63n g i Qi^jjbSn ljmJboi,3hn
5^6bi43oih36noi: Solaneceae g i (jmjbo43hn, 36ô9o4hro63co
b i 3m6n63&n
Scrophulariacea&dn 3°Diy>
j>i6bii3oii 361101: Apocynaceae, iynjbo43hn. 663(13
4>hg35mcng36n
Asclepiadaceae
36m(15360 qjihome 6i 3*H}3C3&3Cni 5 >3i<33hsg363co
3n(yV)]h33636n C^,ynjnq}3h369n 93943336300 9mj33g 36i
i<yn{)36ribrh3bxn ,bc>3nlb 6tpc9u36ih 3369n (4f»i#*i43hn
qQ6mcnfa)bn
9ih^n3n (336^360 b9 ohig qjncnpjbbi g i s6ô9n^»»i63c a ôyyiiscrrrfrin,
ir*y,3jj0n ^h 30133 (^363630 93943336300
3iQmj,36n'bnh3i)3cn <336^0360 "b^nb 6y if 9l536ih 33?)3 n 863(13
qjQi3fn6mng3bn QihovTig 5383033(13936300
^>606360, g3oJbng36n 3^h3&jn9 nb 3jch3g 369n, 963(13, >6ô9 o4hin63oo
ngnm6Q3blil3&9 n, col) 63&9n
gn&3&bmqj3hi6n 0^)363680 0^3636811 i 6 i4n9o4hm63co, ^mjb043h0
s'bme83Bu33C36n
gshwmg 6
iOJPCmneo^ ^rf^qr^Jty^jojtiSri, 5^6^430^)36001 ^mjbojohn, 96sh3
qjjb33&9a cgmaipQ&bi £3 6symqj3&3n
ophonmg 6^3^^00800^
3906360 gi 333^093611 qyJijqijjOPTj6369n, bSnhsg 5363936^ 9393631x1301363011 (3360
!ff>3nC02® i
9bb9n>6 6ipo9lS36ih 3363n lynjbn43hn. i6ô6nm^n4n
bniCD*ln 3mon333lyig 36n (5i6bii3cnh36(101: Cyanobacteria,
Dinophyceae, Chrysopty'ceae)
[jnprib ih930J963pn b9nhig 53639363 3shimb6363 n ^003)1 (Jffljbn^n
i 9 n60193^336(1
dytaiyyyytm 5>3hU303&3 (631^10336113 lyn#j043hn
ljn>6m5363hn 5cn4mbng36n
gi qjpwc^**1)
{,0 3^ ^ 6 ^ 0 ^ 36 (1 939oib333nottg Bh3Uxnj3(j333-9 n 83333, 863(13
bb3» 93mhjgn 6^3^101360
b9 nhig (Fabaceae, Rubiaceae, 3C3CfTîOiir)hn1 i 6^n9o4hm63on
4383(111636(1
Poaceae)
3mo(is(j3l5n0353^ 39ot>3hjbig Apiaceae g i Asteraceae 53)14333060039 iynj*i43hn
- 29 -
8 (336^ 00)6 <Wfn39 n
6ib . 1.13 ShnQnvjnx ijs:jbm63&9n Sjm hign 93ty>dmcn'b9nb inm ij^n^hn Shnfrjj^^&nb j>s6:>6nc3&5.
03mc3unnb ^ b p ri& itg S n y ^ n ^ b j'bib (S jjig ajqohnon) 365(330065 933icm 6^ -i( 53o>i iinlj 6^*
(("flip Sjqj^hnon). 1 _ pnj>6n6n; 2 - j>m6go6bnh;){>3on ii>&n6;)&n; 3 - n'bmj3n6m0n6nb ^ ix y 'f'P O '
6n; 4 _ &3(!)ipin636n; 5 _ 6 - n6gmQnb iC iX vr'riP3^>n P* nhnpfnnp36n; 7 - b^jhmnQ^-
6n; 8 - bobj3n^3h5363&n; 9 - imcniuo^cD^n (Harbome, 1988; Frohne & Jensen, 1992)
b]6nb bn0d5O)h3 (53503650016b ppnb 5568530016580 95 3nbn gpn^hn hn^9n 9ob-

5955363005 (5601330015 5^03016560156 (65b. 1. 16). 556653301(136001 6506^3(136015
mhjng^^bnb y353npm6i. bb35 3(10103660560 815365(133606 9bj>53b5g, mhjng^bnb
y3i 3nOn b50633hrt56n5. y353nQb5cg5hnb 55(13 8(13 3395365 b50n 350nb 09010101-
C5inb 5556, hm0Q36ni5 j 36o56 350b. 8og5 3h3 - 530(153060, 8395365 b 53n 530(1-
53n6nb 03311(50065556. 8501556 oiho 655553b 3(1018563016, bmpm 33b533 b5b383(53-
OnCf'5 95 8 5 b ^360 361^9365. ^finb qjmh05, b3h6x n g 5 530183^(105 30608363-
00135636 hmpb 56(130366 95850^33(105636300 063hnb 801^093580. 63j(j5ho, (101830-
0(5 00101536363005 6563^11380 563 93^80 ^3606 33565 3b5h3b, 65h3ni59536b oih-
jo933&ob 0O3h 063hob 9ncbng3ob 9585^36001 658350365b. 55b5m(55ho5 oihjog33-
5ob Ophrys-ob 535hnb 6ih3oi85g53603&ob y353opnb 553630365, hrn030oi5 363hnb
9mbng3ob 80^6001 5bg36b 0g3gho 363hob cgmhSnb, 033(106, ^ jb ^ h n b i g5, (15(5
001535(105, b3h630nb (033(10180163606 0b553bo 801033303(10 b^h^j^hob 3jm63 63(1-
6300b 5568530(101636300 Jo3o3hn 6531101360b b5335036ooi) 6 h x 080^5(5056.
50X^3501(15 315365(13015 8oihob 3(10103(10101606 nb30io 0301(1855, (1019365(5 obo6n
3h0i35630i‘b 3 80^0303636 0501 003(1 550oiyoiq3o0o Jo0o3ho 6o3om3h3636ob 65835-
036001 [ 154]. 50 6030103113636b, hoi030oi5i5 50010016360 36010365 ((5bo3oi356n
3953360, ^301360, q336oi03ho 653(101360, 503500109360, b,33(10100360 g5 3305ho6nb
65801360), 0(5365(13360 5580^010336 (101501(1(5 60(501(56006 5560530016580, ob3 6033-
gnoob 330g 355U - 55bh66ob b(j>5gn58o 0yoiqjo o)h656oi36og56. 50018016360 5580m-
ymgj5 353(180 0301010360956 (b8nh5g 13h56b 3oh5(5nob ghoib Bygob oihov)ooib6
3hoi5g), 56 60595530 03363360056. 63080b ghoib 3b 6030103(136360 gogo (150193-
6016001 658010(13(56365 p 3hoig56 g 5 0301010360056 60505580. ‘601503(1010 8(5365^
5030oi35oi3h 653hoi36b 550oiyoiqjb 6y503o(5 [53], 65b. 1. 17-*t>3 65R3363605 50
- 30 -
3036663016 U6libn3A 063^0 &6639 P)
Ubhnon 1.4 3n6jhiC 3hn i33&nbb g i Eiycnb g^cgnufxyib {>*3036* ^1^0^1346 9u36ih;)oii a^mhsg
So^s&mpn'ba'bj (Gershenzon, 1984 ) (+ fci'bhp:.; - e s j 03&*; X h jiju n i ih ihnb 5 i 9m b iijx n)
,53h3 ;)6 mn@;)&n x +

6^0^636(1 + x
b;)3&n
CQ36m0nb 6:>hfi;)6;)&n + + + + x
iCyjCmneDbn + - x + +
6 C3 i^ l 3n5 niC»D&n + - +
Ur>i6m536]hn 6cniFi<bnQ3&n - +
OCH>
H2COH , .
| H jCO X 0 °^
HjC % ---------° w
H j C o '^ f ^ o c H ! H9 — 0 H? 0H och. ----------------- 0

ocu
Hip— pH
HOChfe-CH— CO
H fo'0*
con
HsCOH OCH, O
OCHs ocH>
C^
°Ob f 9 I 1 10
ChjOH
I 9—^ i i S - 0 — 9H H3C O 'T > rcHi
9O------CH
HOCht-CH-CHO CO
HjCO
HOC^-CH-CHO
H3C0 T oc**
OH
R,=R2=H: pcoumaryl alcohol

R,-OCH3;H2-H: conlleryl alcohol
R^R j -OCHj : sinapyl alcohol
65b. 1.14 6nq£nb pn£,6n6 nb b ^ v jJ a jh x 0 StoWO0 6^ qjD6np3hfn3i6(ib (|jn6i3n6nb b3nh,yi) b,jh]J-

liD^XO oji^hdxS' CNimz, 1974; Lin & Dence, 1992)
- 31 -
3(J365fn00to adfnOdD
6ihnb bib^mbnb, mh5i6m3&nb, bbbnumybQm 3gj>m0ih3m6nb, 5i3hu3P3&nb Snhmbj-

6 nbi g i bb 3i ogij^mhQbnb 3ob3Q3ncn.
hmftmhu 3 3 3 3 ^(^>36036301, 8 3 ^ ^ 6 ^ 0 0 ^ 3 6 0 8 u 3 6 ih o g i6 R i 3 mnh 3 u b 3 & i 6306001,
gidmoymgji og3b336ngi6 gi bg3&i 6 nigig)9 o 3 icno bidding gngn himg^mbncn
pij)hm336i, hnb &i3 nnj q j^ o b j,ih938m 2»fT)6u36(i)hnhg3&ni6 3nj>hmnh5i6n'b336n.
i 3hn&ig, nj863&i hn‘bmbqj3hm, hmScnb mh{)i6o<b8;)6o &i6bbj)30ih3&3C qj36jun3h
36303b - 3ihnbmj!)hmq336b 6i ^3i3cn3636ni6. 33i(pp3bo 8u36ih33&nboi3nb 0icno
hmpo 3iph3big gngni, 3n6inpi6 nbn6n J36 oi 6 cju>30b363h qjnp^hb, hma^pSnij
5i03Qnb oj3b3nb £>i9m6iymG9n gi, ^(130133, y330i nb 6n3cnn3h36i, hijj q93b33i>nb-
336 8n38ihcn36i. 3]>hnbml!)hma336n ibg3636 qj3b336nQi6 h3<bf»ibq)3hm0n 338i3i0o
g i j,ih38rr>gi6 8m 6m p3&3pn 6 n 3 ain 3 h 3 b 3 &nb 9 3^ rn jb o ^ ijn ib .
n6orrfctfn 6 n b A s 6 p iU 3 i g n vy ry y n jb o B 3 6 n b i g i gn & fn iC 3jb n63& nb b i 9 3 i 03 &ncn.
5u36ih336b 3 3 b 6 33a i 36ihn b b 3 ig ib 3 i 03ji6n<b8ob bi33ip3&noi p inu 3i6 0030
n6<33jbr*36nbj)i6 , 3icn Jbm3O0369n cgn,i)m3icnm636;]ho 6 ijlj3hn36nbi g i bmyn^&nb
33^hobj,i6. 3b 03ji6n ‘b836ni: 3n33hb36bnj)3hn h3ij(50i 330ml!)33nbig5n, hmg^bi^
p i‘b n i6 3 6 3 0 n 3 3 h 3 g 3 & n b B h io p p n p 3 3 3 6 n i 6 ; 9 3 0 m<}hnpn mh5 i 6 n'b 3 3 b n b 3 m0 nb>
6sb. 1.17 ^C3OfT’3 icn0 hn b ] 6bvi i 6(jn 36n. 3 oi3M>3en3 &nb 3 3 9 li3 X fr|^>i Eucalyptus cilriodora -b
{^Bjnaohj&nb (jnjjpnb dnboe3nan (d itx jb ^ i bjhsow ); urjbQob {>i63fKnjh36nb ip^pmSicrrjhn n63n6n-
(jn i bb3*g ib b 3 i 81536^330 ( 3^ 3336 1 bjhicnn) (Rabotnov, 1995)
jihognb îq jb xD ^ri 3mju33b g i 6ioin ndmbngrfbiijni; i6 li,n3nh3b3pn, i6^n6ij-

(i3hn30n g i i6^nbrTit,m3i6n 6n3om3h3636nb ^6013^0. i8nb 6icn3pn 6i5ipnoi36ni:
i CflC X n b3Qgng36n (B ra s s ic a g i A lliu m - nb 53^3630), 0iJ^m636n (^0^3630 g i
R anu nculus- nb bib3m63&3n), bi3m6n636o (b3hm3&3n g i j3h3n), j3o6m636n
( 3i 30 nb b 33 n), bi0nyo0ob 8 ^ 336 0 [ 192 ], qjQS3m6mng36n, vi)i6n636n gi vbDh^3_
6mog36n (6ib. 1 .18 ). 9b6b3D0n 6i3hcn3&ni qjn^mi03^jn636n. gc«3b giib0R36ncn
20 0 b b 3 i p i b b 3 i q jn ^ m ^ jb n B n b jn 9 n3 hn b ^ h ^ 3 h i i u6rn6npo, hm 3 Q36 o|j b b 3 i-
gibb3i 63ihnbcn3nbii gi3ibibniori36xn: n‘bmqj0S3n 6mng36n (Fabaceae), b3bj3n-
^ 3 3 6 3 6 0 (S o la n a c e a e ), 3mpnii53^nQ36n (A ste ra c e a e ), mhjn6mcn ( O rch id a cea e )
(6ib. 1 .19 ).
- 34 -
9 0 0 6 6 6 3 0 1 6 l^ U b ro tn O & O ^ O &6 6 0 9 H
PH
K*
H H2 ,8-Glucosyl COOH
HjC=C-C-C*
N-OSO3 CgH,
Mustard oil glucoside Lichen substances

{Sinigrin) {Ordnol depside)
XXX
C H ^ '^ '^ O C H j
Phenanthren
(Orchinol)
HO iHCOOH
HO
HOOCHO OH
Hydrolizable Condensed
tannin tannin
(Gallotannin) (Catechin)
64b. 1.18 s6^n0nihR&xn 0l33iS r*J 9(j36:>h3xn 6iDhoi]6nb ‘bm^njhonn 9sj>i(y>cno (Rundel,
1978; Harbomc, 1988)
Stiibene Coumarin Anlhragallol
OH
R = H: Pinosylvin R = CHa: Scoparone R = CH3; R, = OH; R j - H

R = OH: Resveratrol R = B-D-glucosyl: Scopolin R = H; R ,= OCH3; R2 = OCH3
61b. 1.19 eb9^330n 6 bQhcn36n - <gn^miC3Jbn636n. 900)360030100360 (b,5nc&n6n) 6ih 9m 0^6636:1

8h i3ip OmfbhjSn g i i j 3b bmynb bb6o6b:>(n9g3f>m 3033^ ; y W o h3<b33hivjhmQO 6bh8m nj9636i
ebmpng 3 jb 338935, hsy dsbV ) njhnab dnn^>6b 'bmftnjhonn bm|)R, ifch 30133 x^ M in b o jah n {pSmbb-
n336nb ghmb. bjn3ihn6b gb byi3m (vi6b (^38^ 06360) i j 30 > 3 mb>y i6q33j|y i3hn 8608363pm di. i>jh>
5 bpmQ36o, hm0c 36no b^fc>jn6pi6nb 6:>hfi363&n;>, {>i8mnhR330h6 ^301503^ 0*1 bmynb 9o3ihcn (jmjbn-
i3hm&oai (Gottstein & Gross, 1992)
jnciirjhn 6 ^ (5 36 0 36 0 13 5^ 6 . 9 (536ih 336 b d^Q^don 01^39^3^ 9 \JHS3 ^C 3 ^nl>

j>i6 g i i0nbcn3nb n y 3 6 3 6 3 6 nb3cn b i 9 3 i>C3 6 3 6 b , hm ^m hnuii 330 36 0 , 9 3 6 3 b 3 b,
331139360b 0nj)6n6n‘biuniI 33h3gnb 33gco69n bnonyn^nb 9i(j>i0n 939(533CfT|&>-
2,i 5 b i 330 ih36 n0 i i(Woi6 n9 5 i 3 n i jn0n3hn 6*3hoi3&nco 01*39^(53^, hm0C36n(3 dyyd-
- 35 -
90066630)6 &6 (f)09 n
R - fJ-Glucoeyl: Prunasin Benzaldehyde cyanohydrin Benzakjehyde

R a p-Gentiobiosyl: Amygdelin ♦ cyanic add
H3C s / ° ~ ^ a[ucosa R . CH3 : Linamarin

R - CjH5 : Lotaustralin
6ib. 1.20 uni6 m6063hn ^Qn^mVieoinb fi30X ^ n3n ai6s6nc3&i: amigdalini Rosaceae-Si; prunazini
Myrtaceae, Caprifoliaceae, Rosaceae g i 53n9h369n; 0 n6i 9ihn6n g i omî3b^hion6n Fabaceae,
Euphorbiaceae, Linaceae g i Asteraceae-9 n. 013 ushjgnb 9iVii6;)&nbib 3i i 3mc3d3 n ihbj6x °
5pnj)m'bng3&n 93300® yn 5^ iij 9n &3^>-5C3>)pi'bnp36cni61 Bih9mnJ9636i cr^nb^qpen 3nphm(5ni6nb
9gi3i (nb. bjhiomb 'b jg i 6i6nQn)
P3&i nymb 636013^611, ihibibtii0m36m b^Gnbi gi 53601b 9J0163, ^mjbn^hn (6ib.

1.20). gi0U3Oc 6n3cnn3h36iOU> 30h:>3Q3boi6i gi3ibibnicn3630ni i0 i 013 n3 bib^m-
6nbcn3nb. 6113013113&iaii 3hcnn jjpibn îSbi^^cnhQbnai 0<33jiJ3hni d3d306oi3ihoii
0n0ihcn, 03Oih3 - 0nj3636nb 9 n0 ihon, 03bi03 - 063h3&nb pi i.0. (6ib. 1.21;
1 .2 2 ) . b id m 3 ih 0 9 3 ^ 0 1^ 3 n0 b ib 3 0 i6 ic n i qjoioiQ360n, hoi0c3&n(5 n 6 ^ 36 b n 3h ig
Solanaceae
Convolvulaceae
Solanaceae
Piperaceae
Apiaceae
Campanulaceae
Fabaceae
Lytfiraceae
Magnoliaceae
Ranunculaceae
Papaveraceae
Rutaceae
Rubiaceae
Convolvulaceae
Apocynaceae
Loganiaceae
Rubiaceae
Rubiaceae
Aquifoliaceae
Theaceae, Sapindaceae
Sterculiaceae
Asleraceae
Phenanthrene Papaver
6ib. 1.21 iCiiOoing^fen p i n9 m jib 3&nb
9i&ignov)&n, hm9oo&9n(j nbn5n ih n i6
6i3huxD*>Xn
- 36 -
,o
COOH
nv
HOHaC *n i
Saponin (Hederagenin) Cardenolide (Calotropagenln)
r 9"
OH
due
HO
GkioGal-O'
HO
H
Steroid alkaloid (Demissin) Sterol (Ecdysone)
6>b. 1.22 »>hn^3h33636n, hm9 x o ^ 0 9^ 536 036 00 (dnhnotogê 963h3bnb) b^6o6>^(^6Q>

&n pi0(j33cm6noin jndnjhn 3 * 3 3 ^ (Crawford, 1989; Schlee, 1992 )
n d m 3 3 b i, 6 i i h d s p n 8g30m b 05033 b 5b 3m 6 3 6 0 15 6 3 3 9 ^ 3 6 0 0 ), 830866335 bncngn-

38 11b 35^365. i 5bfn 330 Oi5 8 0 3(1 8 ( 5 3 6 5 (13 0 15 95^056365 pb 0 6 ^ 3 6 6 0 3 (10 dm 336i
815365(13360 0 06333b 3 m c ,o q j 3 6 m p 3 6 o b , ( > ,5 6 0 6 3 6 0 6 5 g i ^ 3 (13 3 6 3 6 0 6 3 1116 (5 3 6 ^ (15 -
(503606 8 5 ^ 3 6 5 6 [ 2 7 ] . 0 5 0 3 3 5 8 6 3 (13 6 0 0 15 g i 6 m p 3 b i3 6 o o i 9 5 ^ 0 56 36 0 b 8 3 8 0 16 3 3 -
3*30 (5.
1.2 awib hienibni ei icna>sn

9 3 9 5 8 0 6 ^ 3 bnumi5bQ3b ©“b o b 603b 5 5 8 m bbo 3 3 & x n Q* 6nm bg3hfn8o 03b30O
363h5oob 653590 3’bh363x y * ,io)b- 6on9ibnb 8365;)36305g gb y330* mh5i6o(b9ob
(5bfn3308mj9xD^n^;>cn3r^) bi^bhoboi 8bob (i5gO5t500b ob 8 x^ 3 6 0 0 1 8(50(13 hbm-
g36m6i, hm330bi (5 8(5365(13360 q|mli)mbn6oi3,bnb 3hm(5360o 630013063636. bbo33ho
363h500b 3b 656000 qjojbohgjdb jo0o3ho 66360b bibooi. (15905(5006 3ghm 96o-
83630^13560 (1501936^65 3(536 55hg5oj06365 booi6mg, hm63cni5 656o0m6ho3 6ypob
5mhaij03&:>b3 0^(13365; 9565(16360 6001601 6ipQ> b633b 9x^80606 *bx^nhnb
(!>3833hi^3hib. 58535(159, 8bob (15905(505 boo»63ho 65056b0G5 gs 6y0ob dn9n>
I530ob, (*1(1556300 6030103(1363606 65h8mj86ob5 95 55(195j66o6 Byihm. 58533
ghmb, 6(5365(1301501306 (1*90*15056 5j3b ih5 8bm0mg O}mj,m363h53^ni30n g i qjm-
0>mio6 3 h63 j , o i X f ' 3 <3 DAi3 &n> d i^ D Q (150 ^ ^ 3 3 3 0 6 0 0 5 9 , 6 ^ 3 6 x 1 1 o p f o m h o ^ i g 5
5556605 qjm^mg3b(i)h3Ji5n30n 3033^0(5 (ybh. 1.6).
37
8 0 0 6 6 6 0 0 )6 cW ffjo an
(jbhnpo 1.6 9%nb hsgru>unnb &»303f» Q yaB ihj'b] (Ross, 1981) ( * g:>dmyip3&3Qo;> (Jbfib iJbnb
imbnbm'b] p i (^hp6cj6nb bo^nghrr^Vj)
atwib ajownpob. 0.3*1630"
hign>iyinb ^CC'nb bbfl33hn ov)b^« Smhojpv
b3j)^hn bn5hdo (59) 363h&oob &3&Di5nJ 3^n
Shmê^n
39609363- 6606363-
3C\jhsnnbo93hn 290-360 0-4 9|jnh3 38608363pm
Cm 0013560
gm^bnbon^'b^hig
6608363- 6606363-
ijôjho hipnîyib 380-710 21-46 s 9(}nh3 661183630013160
0013560 0013160
(P3flC>)
39609363- 6608363- 36608363-
nBghiSnorxyi 750-4000 50 -79 > 86093630013560
0Oi 0013560 cm
396o936 x 39609363- 38608363-
6hd3WPCC'n^ ' 1 4000-100000 86083630013^60
pi Om
i.2 .i h s e r u * s n :>
i . 2 .i .i t o Q r m D c n 6 ih D a n
6nmbq}3hm 6t>nb h5g05ijoob Qiibcm^bncn 2 9 0 68956 3 0 0 0 66-693 bn&hdnb

jjiQP^bb np^bb. 3qjhm 3mi|03O50crin56n h5Qn5un5 9oi5fi6oij9365 m‘bm6nb 6hnb
0n]h, bmpm 'b(p3ihn jjhd^pn o^C^ôb 365680 95601^0936x05 3531180 Bypnb
mhcnjQnbi 9s 5 bb9 nh6 ignb rihg565 ob 939 |j330m65'b3. 559 mbbo336 x n 6bnb
3 6 3 h jjn n b 4 0 - 4 5 % - 8 g 3 380-g56 710 68-893 3 6 5 6 ^ 3 9fTi9ob. b33johob 59 65609b
iCP3nj3i8cn hmj,mh(j bnp3Q bn6icnQ3b. j0mhm305booi5 3058360360 80156015336
5i6mbbn336ib b33^hnb 59 36^660, 59001*18 85b 36019365 "qjR^mbnba^'b^h.ig
bjon3hn h5g05i505n 563 qjih-n. b9oh5g qph-ob bi(bp3h36i9 600^6335 4 0 0 -7 0 0 6 9 -
nb 36560. cgbh-b 303(101 801^93 oi CC’D^)n'J 9 bhng56 3b5bc>3h365 39 oh5nnbqj3 hn
(159051305 (U V ), hm630ni3, 01530b 8 b h o 3 , nym<35 mh 6560059: U V -A : 3 1 5 -3 8 0 6 9
gb UV-B: 2 8 0 - 3 1 5 69 , bmgm 303(101 5hd3QO oi C^3^,n^) 9bhng56 - o6qjh56001300
(15905^05 ( I R ; 750-4000 6 6 ) 95 013(160^159051505 ( IR ; 4 0 0 0 - 1 0 5 6 8 ) (ob. ubh. 1 .6 ) .
93956060b 5o9mbq)3hmb 55h30i5 b5t>c>35hb3 8bob h5g05(500b 06036^30165
5hob 1360 W S'8 ( 8 5 b 3bnb 8398035 36mg365). 93958060b b3953nh53g3 650130
9C>3b (o53cnbb83&5 9^66369090 560690 95 bbn33&ob 3311336902)395(1x0 8 o65 h-
013936001 55‘bm835) h5905|j0nb b59350oig 66019019 47 % 5p633b. 6bob bbo336nb
65b335h<b3 6300 8580633 55066335 b50 y5hmb bn3h^39o 5o3 mbq33hmb ^395 093-
63680 55h9503bob5 95 553hxD&ob 95, 330113303b yoi39ob5, (*^36936001 5h3$ 30 nb
9393559, 55h3OT33 55066335 95 8015060^8365 ph36o3&nb, 6o3hob5 95 6y0nb
63301360b b583503&ooi. h5g05U05, I101839015 6059555693 56 8 u365h3x b50)5h58Q3
50633b 86019019 656o0n6ho3 8395365 3ohg53oho bbo33&ob5556 (3ohg53oho h59*15-
<505); 9565(16360 656090 y i - ^53(165 95 (3*3693680 3^33 55<356 o x n:> (6i<3 i,6 -
0 X n & 55663390 h59051505). 3oh953oho 95 55663x0 h5905(joob 3hoim60noi65b
- 38 -
980b tfU>«SOaOlK> 3^09560
6bb. 1.23 epn^hn jombic^hn hsenium (GJ Jjfr2) (Schultz, 1988)
& pri 6 i 0 3 hn h ig o i^ o i ( i 6 h ig o i^ o i) 36 0 19 3 6 ^ b i S ^ m 5^63936^3 b ^ n b

gm63b3 50m6i0ohn higniynob 8ijbo8i03hn 8^^33636300^ 1 kW S'2, bmpm qjih-
nbi - giibpm36noi 1800 |i 3m0O ci}m^m6n 3 -269 -l. 8bnb higniynnoi 3 ^ 3 6 3 30 9 -
mcgnb 9bhn3 jtfRfchiqjnxn 6^Den^' 'be'3nb gm6ogi6 bn9 ippnb, ig&opob
h3cn3qjnbi gi ("h36pnb6m6ob bihnbbob 606393001 gngn h3j,om6sQ3hn g i Qfnj)>
C3hn 5^ 63^ 336 36 0 ^ (6ib. 1 .2 3 ). ib3, 9i5iQooiig, 3i(r»i0n iâmbqfthxn 663-
30CTU) gi 0uoh3(T>h36oni6ri6noi 5i0mhR33po ^hm3n i x r‘ hinm6o6o di0Oi6 633b
3bob bbo3b 0^3636: i j 3i3hnb J>i3^ 3nh3b03 fcihbo 6nbgij,ob <b3gb3nhb3g3 iî-
h]6b ihi 47 %, bhbdog bi33ipnig higni^nnb 70 % . gng bn3i(j>0O36b3, gi60m-
636006 33gih36noi, n3 ob j>i8m, hm3 bbo33&b 3qjhm 801303 j>bi hfooi j,ibi3p30O
gi 3i 3 hnu 3qjhm 5 i 8 ^ 3 nh 3 i 0 3 i , hign^jonb j i 9 3 hn o 6^ 36 b o3m 6i 83^0^. 83^0(5,
g39>9n6nb b3gi3nhi8g3 3mp6330n hignijjoob bngng3*b3 8mJ83g36b 0mip03hn
^nim&hiqjoopo oii3nb3&3h36i6n g i g(pobb g i 60nb Sohomgo.
6yip 3o dbnb higniôb 5^0036001 oqjhm dgnoh g i b6hiO]ig 3(jnhg36i, 309(13
3i3h3n (6ib. 1.24 ). 6nigi53o 30 higoiôob 938(jnh36i j>i63bibc3h30ig gngni.
ib3, 3ij)i0noTig, <b3gi3nhi3g3 3mp63x n higniynnb 3bmpmg 1 % 0*3 i£>633b
6nigi5nb bo(j>h333n 2-5 88-^3 [231 ]. higniijnnb 3ou>6oiJ8nb 33g 36ig 6y0nb y3o-
0 ib 3 *b3gi 3h336n 01636^ O)6n0 6yipb 3qjhm 6(30(13 ^833(1033 ij3b, 3ngh3 ljn3b,
g i jihnot 5i3m63330n go63&3&o bn3i>3hn33b 8bmpmg 6i hi33X boph333p3
i6m6ib6mh3636. i8nb 5i3m o6^36bo3ho higni^nnb b3<bm6nb gi6y36nbib 3g5hig
6yi0bitj33363n 6ih8mnJ8636i ^3833^3(1001 j>i63nhfn636x n 906*6 3g 6hign
6ygob bo6>)3ho3nb b^hi^nqjoîyni. ,563680 6ih3moJ8636i 3h0idi630inb5i6 &i8n-
3 6 x 0 6y0nb 3ib36n, ôhdmg d0noh j)i6 io i3 6 x r>. 0* s9nb jpBm 3qjhm oi6opo g i
3b3&3jo b 3g i 3oh30n 9h3 (33non86nm6n) g i 8nb ^338 9g36ih3 3cghm 1^130 g i
81,3^30 6ypnb 8 ib i (3n3mpo36om6n). &igib30i 8icn Smhob biâimg 3i33o4ini,
i8n(jm8 bibib(p3hm bm6ib 6ib^rn0ob 3h3b (o»3h9mj)cn6b) 36(1193636. 838^9-
5m3i b 3 b 3g i 3nh30o 6yi0n iyi3g36i, b^hi^ngniiuni jho&i g i
^6i 9n 6yi0n 8cn0ni6ig inh33i. iS ^ ih ig , ^6ob iKjS^hi^hoon h3gn9nbioi3nb
- 39 -
9(306660016 &6 ff>09P)
gi0ibibnicn3630Oi b3‘bm6jhn 153000363611 33n0 n06 nm69 n gi 3(101-

6inhn gibign ^3033(1^3(1360 3n3mQO06 nm69 n. b(p3i 9 n p>h0 ig 33031^3(10 bmenb
J330 03g0n3n Oi3h0mj)(vi6ni; gi0ij)36ncnn 3h3 l53033hi(j3hnb 0^3301(10 15300036001
Bih0mnJ0636i Sgnb aihi 3330 ghmb ©bmgmg bs93s0m j)i63g36'b3. i j 03bidQmi
nymb ^(130133 ^ 3 0 3 3 (1^ 3 !^ b3'bm63hn ©^hyombobn y330i‘b3 ©xpi0 15-40 03^hn-
i6 3 h30 n. 32)3i^mh3Q gi Sm gihx ^1*163680 By0nb ^ 3 0 3 3 (1^ 3 ^ pni b c ^ iS n
0oi30n Bgnb J,i 60 i 3om&i9 n ©3g0n3ni. mj)3i63369n Bygnb ^ 36 33(1^ 3(1^ 3 5^303-
6ib ibg3636 gn63&36n.
a b
Percent of surface irradlance Wavelength [nm]
6ib. I.fi4a,b 6<£>e8n cJbnb hipni(jnnb (350-700 69 bn&hdnb (jiCP3^n) 3D9(yih3t>i (6ycnb Jerlov-nb
bb3igibb3i ^n39n). a 6y(vibj333 9bnb hsgniynnb %-n 9]9i5nh]&i <b3p}3nhou>6 drobx hjenj(jnsboii6
33gih]6ncn: JI 6yipn 04316301} (jhcOnix 6i 6np3n; J 1 b3q)Ou> 6yipn bi6i 3nhm bm(<rii6; 39
bi6i 3nhm 6yc3&n 663i63^>c(pm3i6n higmiyinb dijbncfecjhn &s3lH53p]&<Kn. & bs6>3nhm 6y03&3 n
9bnb higniynnb bSjJ^hnb 5i6}6nc3&i pc>nb biioiQ&Sn (Jerlov-nb ,jn3n 5) (Jerlov, 1976; Luning &
Dring, cit. Luning, 1990)
1 .2 . 1 .2 9 l5D62)hD3 C b ^ h 0 H h ^ G fl^ n flb 6 ^ 6 flCD6 ^
0hi3iQfiih3bni6n 9(536ih330n biqjihn 0np36x higni^nib bh30ig ny3636b.

6^63(130 030^^(5361^^3600 2^.6031*1(1360010 ih3»(30fibi gi cni60 n0g 33hxn ckrii6-
oij0nb 03935ig 6oigi5nb lb3gi3ohi3g3 nbj ymî bn6ioiQ3 i(p633b, hm0 bn6iov
gnon qjihgmbooio gii0iymqjO036i ij b x hi3g36n03 3hm[536ni (6ib. 1.25). 6(53-
6ihnb îb o^ô £>i6ioi36nb Snhm&pbnb gibibiboioi360ig 3n'b6oh0i (Wiesner)
03-20 bi3»,36ob gibi6ynb0o 030mnpm (5636i bn6ioi0noi qjihgmbnato gi^9 iy-
m<gno36i- bn6iongncn qjihgm6ooin gi^diytngnoobi 2>i0mbiî3b yn6j)h3&3c cgm-
oimQ<b3 (i 5 0(536ih3Xn biqjihnb 36i69n) fciGiaiQbjcmbnb Bfipb
- 40 -
880U JC ? 08i &0
bib. 1.25 bb3igsbb3s

(jnSnb 3(536:>h3;)(>m6:>3 n
cfbnb higmtyinb %-n
S^S^nh^bi: imhjip^hn
ihy6ih-6id36ihn
93h3X n i!>ya CbDe^
3ihlib36i b^hicnn)
(Kairiukshtis, 1967),
cgn^bihn (bop* 3ah-
33361 bjhicnn)
(Cemusca, 1977), Sbj-
b39bnhnb 5ihj)i336n
(j33e i 61*1^361
bîicnn) (Hiroi &
Monsi, 1966),
bn9 n6gnb 6^5533611
(j33pi 9ihs 336i
b3hMJin) (Allen et al.,
1964). R U36m'bnb
cb 3g i 3nhnpi5 shîpncn
hspnusm.
(ânbibôb %-6n) bnSiOxyib n3 h5m936m6n956, hmS^nu 50 815365113300

bicghnb ^395 'bcîh'b} 5(pn6n3636i. 3:>5x^0159, 013 ^ynb bi6]h33£>^ ^33001 qjih-
nb n6(|)36bn3m6b ihnb 1500 ^ Smgn cgmon6n 8"263 ' 1, bmpm j33oynb 11111830063
fo^fijnb <31*1010360156 200 (J. 3ri0n ojm^PiBn 8’268‘1 , 85606 i 3 636jnb q)moi036ob
bn6ion0ncn crphpmcbncnn 95i8iymq}O036i nj63&i: (200 / 1500 ) X 100 = 13 .3 %.
j 330 ynb Rhpn0 nb>3 oi)6 n 8 (5365h336o 3353363060 5hru>6 659056 b^b^n higru>yn-
nb5g8n 9s 33(1 $0933 0 ^ 9 36 0 56 5-20 % bn65Oi0ncn cgihgrnbncnn gij)3iynq]n036nb
3nhm&368n. 50 0ifi336360nb 1-2% p;)hmqjmcnon:>6 6t5365h3oiio»3ob bnumijbcnb
j33g5 lb(j'3hi9 00139365. 389^60368^ 0153651133665 63060365 05hb36m6 (53959
5^6^013630 595n0369 n (0 .5 % ), bmpm 3yig3h3b 3380*5333560 0 . 1% (b8303oiob
Byi08 i5365h336n) 8 n6 n6503 hn bo65oi0ooi gii0iymcgn036nb 3nhm6363 n.
h59f»5(5nnb 5h59036^0 cgnljm(536n,b3n 956m^09363005, 33nh3303b ym30 nb5,
33 qjnoi03nb bnb9nh3b5 95 qjmo>03&nb 556955365^3 (65b. 1.26 ). 63qjmai03nb
bnb6nh3 h5mg36m6ho359 63069365 5i3mnb5bmb gmoi0nb t^giSnhnb o6g3jbnb
(LAI) 338331*16001. 050 5633636b y33Ci gmoionb b53hom qj5hoim6b 6059550b
556b5lbp3h30n (b39i3nhnb 8n85hoi [271 ]:
LA I y330ib3 b3 nhsg 5>0mnb}b3<bi qjmcncQ&nb qj>hcnm6nb 8 ®-9 n 6n5955nb
■b3953nhnb 1 62-<b3; 013 L A I = 4 -b, 6059550b b3953nhnb 311013390, 563 1 6*, 95-
y33Qi ogncnQnb 5563110 095(1011*160

LAI --------------------------------------- ----- --------- O-2 )
6059550 b <b 3g5 3n h n b qjihcnm&n
- 41 -
8 0 0 6 d(n 901 t) (W ft 3 8 n
qjih3on D^gi oymb crjmcnQO&nb 4-53(1 Q ^n b3gi3nhnoi; 5ibi53&ni, hm3 031^010360

516015363001 bb3igibb3i ^(^6336133. ^ynb (536m'b363o q)moi036ob fcihgi bn-
6iaipnb bbn3369> 369b jpriihn6 15(130133 P3hm36nbi g i ,3m,3360b bi2>3li330O(5.
i 9 339o*>333i9n( 01651(109361 9u36ih33(vn6nb 1b3gi3ohnb n6Q3jbn (PAI), bigi(5
gmcnp36ou>6 3(10119 5ioi3i0ob6o636xfu> ^n&^nb cgihaimbnu [3 2 ].
0 20 40 $0 SO 100
Canopy cover [%] Canopy cover [%]
g < , % § |l- 2 % H 2-5% [ ~ | 5-20% |~~| 20-50% n 50-80% |~~| BO-100%
Relalive Iransmitlance
6ib. 1X6 qjih-nb 3op633bem6i 6n63m3i6-q)fncnpm3i6 S^hjx 0SO9n (3ih|jb36i bjhiom) g i

C+io6pni6 (0ih j3] 6i b^hicnn) i9n6g3n (Tselniker, 1968).
higniym 9(336^330 biqj>h3 n q3moi036ob c»u>69 n9Q33h x 911336b 5in30ob.

i 9 ibon 6 , 3 nbn 06,336^30161 gi<3ihxoi6nb bihnbbnb fci'bhgnb 33190161^13 3Jb3m-
636150103IH9 9(3fihg36b 0i963h,!r63hnb 3^0631501^ 3i6m6ob oii6ib6ig. ojmov
03611b 9 ibob 93^-61303610 3(101531(11*13160 i53&3036nb>b higmôob 5^9036^(1
83099361 J,i 9 mcn30 n0 0^636 3Jb^n6j>i5nnb 5b6^riC36nb 3 m6 bobi g i bi33ob [ 155 ]
9ri3h bib303(j3onon qjmhSncn:
Iz = I0 *e-k LAI ( 1 .3 )
bigiy, Iz ihnb 8(5361113x0 biqjihob 13391 gm639g3 3 oic 63 x n higniijonb

n6^36bn3m6i; l0 - (11901(501 p o i 195001b; k - 3Jb^n6iijnnb ^30305036^,0
9 fH53930n 9(5361(13x0 cni6ibi‘bni5igm3&nbioi3nb.
3jbj,o6j)(5nob 3013030(5036,30 biboioig36i 516101360b 33(1^03193(10 5(1190366001
qjo^m(536 mi0nb 9 o9 ihoi 6 i 60 i 536 x n:> gogn 3;joibnoi (qgrri0i036ob 3 /4 -
*b3 93^0 450-^3 gngn 330*001), 3^,306 31500b 31^30)0(5036,30 8319536b O.S-V)
613936b, g i 6i0ib6ihob bi 9 ;)icm 90163^3 51610136300161 8150(^361 ihi 393,33b
mli33h p o i 19500360116 33911136001 (16 6ioib6ihob ^ 3 9 1 bi'b[?3ihoni6 8391-
h36ooi) (61b. 1.27 ; 3jb^n63(500b 9ihi530m363&nb ô3o). 3 ohojooi, 9 (536ih3x n
- 42 -
980 U (3 ^ 0 9 6 6 0
6lb. 1 SCI qjmcnpnb 'bjpiSnhnb (1693jbnb

3}fi336] 6pncn 5^0^633^0 bn6Knpnb
n6|j36bn3m6nb 3jb 3m636uni03hn 40361
bb3ipsbb3i ^n3nb 3u36ih 3x biqjihSn.
D qj>hcnmq)mciO(Ti3i 5n mho366m6n
80361(1336™ G 81^30036360; Tb33&n
(Monsi & Saeki, 1953; Kira et al., 1969).
6yonb 3(536}h336nbai3nb 161002,0311(1
3m6nj333&n nb. Pokomy & Ondok ( 1991)
cni6ibi‘bn5igm363(!)nbicn3nb, 659513 <35(10101 0301010360 556015363005 3mhn'bm6-

(biBy^hnb 693001, Sb^b^^bnhnb 6 5 0 ^ 3 6 0 , 8 5 C'506505 b 330 oi6 5 ), D jb ^ n 6 y
(500b 2)01303015036^0 OJ-tQ 63^05; 0^33, 32,33 &505b65hob 6 o35poob 6ib33ihb3
3cnin6CTij036i 95(536300 606501006 2/3-qi6 3/4-693 (65b. 1.27 ; 3jbl|)o62)i5nob oih-
0366016361^ ^nSn). bn65oi0ob 83(363306 958012)093630365 830301010306 bob-
8 0 (13^ 3 ^ 336 8 0 (5 5hb 3 6 oi6 b. 3 (j»h56 o ^ 3 3^ 33 ^ 3 9 5 05(13680 bo 6 icn 03 b (1893659
3cni6cnji3b, hoi8 6059550b ‘639530(15693 d50O56 1501^ (15905(505 5F6336 (656.
1.27 ; 656. 1.28 ). 85680 55650136300160b 5(159036^0 nb3cnn33 56 2)nP33 303(1m
82)3301(105, 309(13 01(103660563606 6i0ib6ihnb ^338. ^ 3 3 6 8 0 ^>n, (101603665(5 8395-
(1361101 03535(135(130560 b33&n (ihyn, 030^30, 3 3V 0 nî5n) J36056, 15(130133 83663(1
2,01(10183680, 55650136300160b 51159036,50 063010335, (101501(10(5 85(1153001363606
Oii6 ibb'bm 5 i 0 m36 i 9 n .
556501363001606 3(101-3(1010 36 (13363001356(1 856560501363005 83030101030b 60b-
3oh3. 050 06555(108365 LAI-956 05 615365(13300 65035(106 301(5300160956 95 55-
301065^365 1101501(1(5 qjmon0nb qj5hcnnb (S2) 609093 8 (5365(i3X n 65035(106 801(53-
00165^3 (83). 5b3, 8555000159, b5gb303o bo8n60ob y5658o, hoi30ob L A I 39(106
4 .2 -b, 83q3oioi03nb bnb8oh3 ^01005 2.6 8283 [ 103 ], 636^(1013021300 035(10101030101-
0013560 651159863563 ^yobi (LAI = 955600136001 7 -b) - 2 .5 -3.5 3e3 5 [279 ], boi0oi
63(^36065 - 1.5-3.2 8283 [ 126 ]. 83030101030b bob8ohob 85633636300 55665^301-
(136001 060301(135^03005 (5502)3300 63 815365(1065 95 (jyob I536oi‘b36o6oi3o6.
83030101030b 805Oi85h3Oi65 3(5365h330m6nb 5563noi5(i365lb35 95801210936300,
53oôi3 bo65oi0ob 5565600365 60ob 5563530016580 0(530365. 3b 3b365 hoifcmhu
goioioi03u336 OiyOD&b, 063 6505601356 oi565b5lb0155901363665(5, 6595(5 830301010306
35jbo338o 95 (159051300b 9585656050136300 5(159036^0 3bm0oig y0oih(j36ob bhgob
958015311360b 838935 95069365. 030101001356 ^ 3 3 6 8 0 '65801(106 3030101001 8950185-
h3oi6ob5b 6059555893 5(316336 bo65oi0nb 50-70 %, q3oioi036ob 5580ob5b - 20 -
- 43 -
8036 ^ ^ 00 )^ to tfo an
Radiation and Cliinaie 39
(n b . 1.E8 cfbnb hipnuyinb 5165600361

qjmoipm3i6 Ciyodn p i ^ynb V ) 9moi:
aaqgmoifgnb ^ 34,130361 & i6 ia 9 &x<’* i'b 3.
gid^hnbxn ^ 3 - 3bnb 3nh@i3nhn hignujni;
'b^pi bi'bn - 5 om6io ]h n hipnujnnb 013(131*1
3a9n; g iS ^ n b x n ihnb J33p i bi'bn - gnqo'b-
n]hn hipnujni; h^rnQ jiniSn bi'bn - hignityii
^ynb q p 6jh ]69n i 3 j6 ijn h n - h ig n n jm bnb
3ih j 36nb ijjd ; 6ib l'b n b t j g i a b ih jb : gni&m-
6i c 3hie QiSijhnbxfi bi'bn - S^jgrnonpisn
3]hnmgn; 3 i 3n b iV i - b h x n Sjogmoionb
Sjhnmgn; 6;}h,jn(vn3i 6n bi'bn -
qjmoim(X5336nb Sohnmgn. (Walter & Brecklc,
1986)
40%, bRQfn bhx<i 9oq3rnoip3nbib 035h3on 3ihj36nb ^338 - 10% -b3 6150360
(65b. 1.28). y330i'b3 nih^bnb 015065(13360 bn650ionb h39080b b3bm63hn
(530003636^59005 33603&Xn- nih^bnb 05365113360, hm3036o(j gmaimp-
3 153360 ^y33&nb J 330 y 33n ighQ^’b ig b x '^ D y3i3fu>6, 01530b bibni5m(5b(vn 150506
dicm6 0 mjj03 33hnmQ3n 559056 (cnm3pnb g6 m6 ng56 b h x 330^0130^693). ib^-
01360:1, 3555900159, igh35i‘bsq3bxnb 5301030,5360: G a la n th u s-n b , Sc illa -b ,
A n em o n e-bjîhnb 61(13013595360360. 5b5065bhg5 bo 0tf36:>h33&b, 05550110159,
03bnbi 95 600300b i(n0m6i(536b, <bm633h qjmoioo&n 51(50036001 53056 6(53035,
3ngh3 ibi]>m3i6 b33&b; 50ob 6y50m6nai nbo6n oy363&o6 556501360b 353013b
3nhm636b, 309(13 3b 333mg6m35b35. oynb J33g5 nih^bSo dmbbhgn ^^601^063^63
3(5365(13360 (H e p a tic a -b , P a c liy sa n d ra -b bib3m636o), 01350 b53oihob
6n 50nb b533503&5b ^03336, b i3cnih3n(5 s^n3hsg oy363&36 bo6 ioi03 b ibo3 n-
0ai5finbs g 5 bhgnb5ai3ob. 0 shag053 i 63 63015 3ih536^338 bo6ioi0noi 9558^0103003-
6s 0cn30o 60nb 5 i 60 i 30m6 i 3 n g5650ni; ^ 5 3 3 ( 1 i j J3395 O5hob3o 5(16360165
0bn0mg b3mhni356 0(536bh33&b 013 333d0fiioi.
i .2.1.3 nseiBnQibeDhn ai3D6^hnb hien>sracn 6^hoan
63636(103 3nhm6369n 3j536ihob 0 o 6 o b b x * 65600360 9bnb (159051505b 0(^3636

bb35Qibb35 qjmhSooi: 3nhgi3nhb, 0h53505bnb 1(1350006 (0501 3 mhnb 6n5g56nb
b 3 g i 3 nhogb 6 ) r gnqj3bo3hb. 015365110b biboumijbprn cgmhQi gb qjm oioob bri3hi5noin
m hnôôb ihob nhrt 801531110 qjij^mhn, (10100360(5 5i6b5b(Ti3h5335 505 013 n 0
0l5365hnb 5ryi65 h3l5X (15905(5030 55h30mb. d50O56 1501^5 qjmoim0o5 bo3h(533n nb^
mho36lio h 3 6 x r', hm3 056 yoi330oi3ob 0nnpmb 0bob 3ohg53nhn bbri336n.
cgmoi036ob 5563bm005g gogn 656000 65b33h5g g5fihgo030 0g5m05h3m653n5 pi.
nbn6n 0(^3636 5h350O0 g5 goqj3bo3h h5gn5(5n5b. ob 03^1010360, hm0C36n(^
yn330Oi3nb 56 d50056 bSnh^g 0^3636 3nh5g53oh h5g 05^05b b3 nh5g 5hn56 5595b-
- 44 -
9 oaB^(f,om^ w i 3b n 3 6 a&ae?o iwrtoan
5sb. 1X 9 qphgmbncm
5p 5ioi36x ,T'^ bnb SJnghm gs
Sjpih^bnm cgi3ih 3ib j 3 n
3“bni6n 8m6936gn<yi 360690b N
3nhm&360n. bbliDibbli ___ ___ ___
9^ n b 3 no. 6* 33636*11 □ ' 0 0 5o % E E ]so .2S% H » ik
5 s 6 i 0 1 3 6 x < ,<iinb 3 3 8 ( ^ 3 6 1 dysphotic zone ladling toliage
3 h 3 & x 8 g50 i8 ili30 i6i8 n .

O'ohmqjrncnQnifjn 6(5361(13360 030011(13636 biibn6o0H5oo‘i bnb)538ib, (1018300(5
311018163010b 5191803111130 9 1 gi8R(ign0i3O qjoioi036ob mh^bobnbîS 8395361.
bnb 3ih33n l5X?iOXn <3010^360 bn6ioi0nb bbn33&nb 3 n8 ihoi 3 icnn 516015360b
91 3ihjnb 8056001 89361(130161^ 3 n b x 3no> hignujnnoi 516 ^ )31336 x ^ 6 8ihi5g3-
6ni6. 10 3bhn3, 33oh33p3b yoi30obi, 36 n836 x m&i iJ3b 3ihjnb qjfnh8ibi g i
q3mcnQ3&nb bnb3nh3b. 3ih3nb 3b cri3ob3636o 516130^10163636 higm^nnb gi8ib i-
bnioi3b3o 5(1191136^ 3ih;snb >>ih5ig 51610136300 33(10033(100916 3ihjnb 80560011
gifihgnpx 61600361893 (61b. 1.29 ). b8oh 31(133680, 8151000119, 3ohi8 o gxn
53o3ihoibob 31(1380 bo6ioi0ob 6150360161 80560011 6i6o08n 83od03&i ob3 9090
nynb, hoi3 ij bnbo8o0i(5ooi 01(151601360 ih Riobib36i, boipoi igh3 51630011-
y0mhij36n 0^33361. bo6ioi0ooi gi])3iyrtiqjn036i 83030101030b bib(j*3ih‘b3
3ih3ob 80560011 6160080 (8 n6 n3 i 03hn bo6ioipooi gliBiyol 0300361) b18310361b
nd033i 513153010101 gib536i 8011536300 bib30i6obioi3ob bo6ioi03'b3 b33i5oqjo.50ho
3oioiboi36o036ob 33bib3&: (liy 3Qjhoi 6150360 1^301 3b 3moiboi36o036i Rhgo0ob
030101036b, 3 ooi 303(101 335h x n 83nd03&i oyoib 31(13360. 180^018 5 16 ^ ^ 3 3 36
bn6 ioi0 ob3oiy3i h x 8l536ih3on 3ih336b 8 o6 o3 i 03 hn bo6ioi0ooi 9156^01030036001:
10-20% (ihyo, 0ihojbo, 030^30) 91 8(19000^8(516 815361(13011 3ih336b 80608103(10
bo6icn0ooi gi38iyoiqjo036noi: 1-3 % (boiô, 6id30, 6003300) [2 ],
1.2.1.4 wefwnnb 9 oo 6(na> <3 ncnent> aroh
030101010^3 91(538300 higonjonb 6i6ogn 10(132)0361. 6i6o0n 8oiin6oij836i,

9161(1^360 616000 y i 030101010b 510300b. ojoioignb 8n3h (119011500b ^3.5301,
9 oii6 oij0 i gi 51,51(1361 83(^330010 3hoi(53b36oi, Q.n. nbn6n (5i0(jvib bn5hd3'b3i
- 45 -
8036660016 &6 6 0 9 D
6ib. 130 33hb3nb 3^h59n bn6ioiQnb

(gihgm6noin ^3430^, gi
9cni6arj9i (jicpvib bn^hdnb Snb^ggncn
(Xvhhinnbgohn, qjih-n, n6gjhb6ncri30n)
W avelength [pm]
(Gates, 1965)
Qi0 royig:)bxn (6ib. 1.30 ).

^ 0i 3 0 - b33j(^hnb n6ojh:>6nav)e 365680 qjfnotmpn 5oh33053b 33h336gn 2,305(13-
02>q gî530x n h5go5(joob 70 %-b, qjbh-nb 365680 0bnpmg 6 -10 %-b [7 4 ). 'bmQnQhn
bih^ycnb ^ 3 3 6 8 0 Q5 (jhr(i3o»,x 6308056 ^336811 55650136300 qjma»036ng56
in h 3 j>0 3 b i 1 2 - 1 5 % boQ30 n bo 65 o i 0 o b i; 3 b 5h 3 j>0 O(yi 9 5 5 >6 5 3 X n b o 65 oi 0 3 ibQcnn
^yjQbnb 9 5 6 3 1 1 x 0 J33Q i nbh^b^bnb 5 5 6 5 0 13 6 ^ 0 6 o8 3 6 3 0 fn3 5 6 o B y ih m i [ 1 1 8 ].
863*63 bn6ioi03 >nh3](036i 3qjhm d0O3h (10-20%), 65ho63obq)3ho 95 6001390 -
y330i'b3 b^bîg (3 -10%). 6001300 bn6icn0ob 51131,035 8 3 ^ 9 860 836x^3*60 5
8(5365113015 9 mhob j,m836nj)i(jnb3n. t,3hdmp, ih3i0(i0 bo6ia»038o ,j50(nob bn^hdnb
ai565qj5hQm65: 6ooi30n/&hd30o 6001300 rVbhQ3&i 8(5365(13015 8nhnb gnb^B^nnb
bhgiboub 3(10159; 5b30in 5(132,01100 bo6ioi0ob 8n0(?3&o 0(5365(13 qjo^mJhrnSob bob-
(|)30ob 83933(116001 “s h ^ b " 8i6do0b 8 3 ^ 6 3 0 3(5365h30g3 [2 3 2 ]. x^snnbqg^hn
65b. 1.31 a,b Camellia

japonica-b q)navn09n
9bnb higniunnb
939(5nh36i. a pm j^noTSxn
930(yih36i d^bhnbj&^h
3ih 36Jn3i 3 n (PP) g i
(ph;)&cnbo&h 3ih 35Jn9 i 9 n
(SP) bn6ioiQnb b3]ijhnb
863563 (550 63), 5nan c
(680 63 ) gi
6hd3(Xj5C(rm3:>&* w 30 n
(780 63 ) 3 m65 i)330i36 nb.
b 33<bfngncnb 3hnb 0n3h
higm^nnb 3ou 6o<j3 &
Thickness ol layer [mm] MesoptiyHlayer (Terashima & Saeki, 1985 )
- 46 -
880b
hign^nnb 336n 336 xo ii (qjmcngnb 803(1 ih i 333153b 3 %-nbi). 5i3m 6>

iC n b n i 'bmjjn^hcnn y 3 i3 n p n , hfTi3 0 3 6 nu 30 li,hinnba 33 h j>i 3 mbbn 3 36 ib dpn3h i n h 3 y
0i3O6; ibjcn j>i3mbbn336ib 363h3&n iih&ig i(r>nj3i336 pi 09363636 mhn36^nhig.
‘bmj)iQ}Q, ih3i3onb 36ihn Qi3 mine36x n i b 3Qi3 nhnb 013063636^3. ib3, 3 ij>i-
pncnip, qjmcnpnb ;)3finb36hn 3363b3i (3 ib bb3i 3>)m0mj)n3hn q336j^ni|j ifeb) 2 -3 -
33h bhgnb ih3j>30nb n6^36bn3m6ib gi 33bi6i3nbig, 3j3nhp36i iboain qjfncnp36nb
2>iQib 3 h35 nb bi<ghoib3.
a b
Wavelength [nm] Wavelength [nm]

Bib. 1.32a,b bnBicnQnb idbmh6(yinb bSjJ^hn Jnb^Q m jnjhig 5i 6 bb3:>33& x qjmcnpjbSn (a) g i
jQmhmqjncnb ^ jb ^ ijd n (b) (Kubin, 1985). Sh ijxyi bibjm&nb qjmoico6nb mi^nyjhn SihiSs^hjdn
nb. Gausman & Allen ( 1973), Gausman ( 1985)
3 oii6 o*j3i. gmcnmpSn 33(j'633cn higniyni n6^36bn3hig 9cnin5cnj336i (6ib.

1.31 ). 30^>hinnbc33hn hipniijni dnhnoiipip 3533 33np3h3nbnb 3n3h 3cnin6oij336i;
ib3, hm3 qjmcnpnb 3cghm ph3 i 3h3363n 3 n bn 3bmpmp 2 -5 % ip633b. i3j>3ihip,
3 3 ng 3 h3 nbn biibnSriQ iunm 3 ih 36 jn 3 n b io i 3 nb 3<33jii3hn Xibhinnbcg3hn gjn c^ h n i.
qjih-b jprnhm^Qib^nb 305336^36(1 3cni6oiji336. i 3 n^m3 gmcnpnb 3 cni6 oij3 nb
b^j^hSn 3nj>36n jomhmqjnpnbi gi 2phmlin6mnp36nb 3n3h bn6ioi0nb 3 ou>6 cnj3 nb
3 ijbn 33036b 333bi6 i 336 i. 33bmqjn03 n 03b3Qn qgih-nb 60 -80 % jpmhmUQibj^nb
3 n3h 3 cnin6cnj336i (6ib. 1.32 ). ^503(1010 b3i3036li)3hn 3 u36ihnb ngnn6pib^3-
6n pi bjj03hmq3fip3hn 3^36ih336nbi gi 3ic33&nb gmai036nb 6m^^m3i6n 6 ih 3 m6-
ij3636n 630b 3691*1636 bn6icn0nb 5 iljih36ib qjmai0nb jbrr^npaii bnph333n [261 ].
pi63h30n (|)hm5ni30 n 63n3ni6n j>y336nb dpn3h piRhpnox d336i> nih3b3n 3ubm3h3-
6n ‘bm^n^hain 6i0ibm3i6n 3u36ihnb gmoi0nb jbm3n0363n ihb36m6b 0n6bnb3ij>-
3ihn 33h3g36n, hm3036nu 5113636 pi 6iP:>:d3cn bn6iai03 jbm3ri0cni bn(ph339n (nb.
6ib. 2 .31 ). bmj^hcnn qjmai0nb 3^ 3118 30 0 3jh3pn pnpn himg36m6ncn 33n(5i3b
i6vyrii5ni6n6b, hm3xni3 33bmojn0ngi6 3533 fci3 m b x n bn6icn0nb h3i6bmh6|5nib
ib g 3 6 b .
qjmonmpSn j>i30nbib hipni(jni b30 3qjhn pi 3qghm b3bli)p36i [110 ]; i 3 nb
- 47 -
8 0 0 6 6 6 3 0 )6 566080
6sb. 1.33 dîpfjxibib 3bnb 3nhgi3nhn

higni(jnnb & i6i6ne3i>i 9^nghm
9(536ih3xn biqjihnb gifihgnQx 6i6np3n
(Smith, 1981). bb3i 3m6iu 363611 6163 m-
336 gi l4hm3n i x (MOD^'bO nt>- Alexeev
W avelength [nm] ( 1975), Johnson & Atwood ( 1970)
25
20
£ 15
■o
c
(0
W
o
6ab. 134 a-d bn6 icnQnbig9n I*jh3j£)h3cn ig:>3iy>un3&n 33b6ih-h(3bnp6ih (jy33n. 9h}gn ip63hb 8 3 }
epnb biiJîer'i &i6iOi36}b n3Qob3n 3m6d36pnc i 6o5 g3n. a qjmonp3i>nb n6icn6i(5n;>; b qjmcnc36nb
gbdihjrgnb bnb3nh3 (b3hicinb 3ihub36i 6i6non) g i bsfrjaa &i6(v>j,3&nb bnb9nh3 (bjhiomb 8 ^ 5 33 -
6 i 6i6npn); c 93bhnb3&hn 3ih36jn0nb 35h3P3&nb ^nSr^n q)mh9i; d JcmhmqjnQnb 336(3330016}
3jh 3g3 n 639c 6016^3 (b3hicnnb 9 sh(jb36s 6i6nen) gi qphonrnfc^ (b^hiomb 3 ih j 335i 6i6n0n)
(Tselniker, 1978; Goryshina, 1980, 1989; i 6 scm6n3*ln 9rn6iU393&n 6n63m3a6n (iynbcr^nb g i bi03hm-
<gno3hn 63fiJ5ihnbio>3nb nb. 93bi&>9nbig Aussenac, 1973; da Gratani, 1993)
- 48 -
880b J ^ 0 8 ^&D
5i3m cfbnb 363hg,nnb him936m6i, hm93pbiu qjmcnQnb 3hcn9i630inb 9m6g336m

8(13360 0(^3636, 3jb3m636unip3hig 8150(1936^. n6qjhi6ncno0n 6603360 2000 59 -
9q3 o6 i 69 n 0 oiin6 oij9 o{>i 636^^9, bmpm j,hdo0t!PC(?ni6n 0163(10 higni^nnb
36i69n (7000 69-^3 93^0) - cnnoij9 nb 8cn0ni6ig (9 7 %); 81536^3 bncn6;)hn
higni^nnb 9 o9 ihoi nb3 rr)U33:>, hm^mhu 8530 bb3xn .
bbn3nb febobhota. qjmoiQnb bo6 ioi035 i 9 ^m0 igm 6 i 9 nb i& 36xo6ibi 95 bnb-
J3'b3i gi 9 n 1)0936300. cnbo0O 83<bm8mhqj30n qjmcnm0O iîh36b 9 bob bbn33&nb
10-20 %-b, diQni6 cnbxn qjmcnmQn - 40 %, bmpm bj30n gi 638390 qjmcnmQn -
3%-‘b3 6ii036b, i6 8308036^ hignijjnib bi3hoimgiu ih ^ ^ 3 6 9 3 6 [11 2 ]. y330>
t>3 g n g o {>i9^m0igm&i b33johnb 08 6i6op363ni, hm9036bi(5 di0ni6 gngn i8-
h3i0i3O 36ihnu &iiR6ni; 3bo6ni 963^63 gi &i6bi2>:30)h3i>nai ib0m n6qjhi6non3pn
366360. ^800^9 cgmoi036nb 9 ibi 9 n 6 i 3Cn0 n hignitfni 830^9 9 gngihni
0iibpm36ncn 500 69 -nbi g i 800 69-^3 9300 bnj,hdnb îoc^ncn. qjmoi036nb
6iO)ihj339 5i6iom63630Oi 6ncn3Q-963i63, bmpm ^yob 6y3goig9n ^»n 9bmQmg
6ncn30n gi o6ojhi6non3pn Rhgn0f» (6ib. 1 .3 3 ) [ 1 88 ].
1 . 2 . 1 .5 a i 3D6 ^hDCrD C n iX 'D h f l h ^ f W d D C f l i c n a ^ n b ana^cn
higni^n^Qn jj0n8i^n6 8 o9ihcn 9^36^3360 i3Q3636 igiSî^nnb 6b35gi6b3i bib3b:

9 rn030ii5n3hb, gmgnqjoiiyr^hb gi 33m03i5n3hb. igiSîtfnib R336 5i63nbn0i3<7i qphoim
ibhocn; nj,n 8mn(ji3b; bi69 m x 3 h3ijun36b, iion9 i lyVbiuni6 gi 536(^^033606 33(^03-
UOib.
9 mgx*un3hn igi3ly>(5ni. ib3cnn ^nSnb igiS^^ni ihnb 66hiqjn gi 93J|J33ign. 9 i&i-
0ncn36ni; 33Qom^hm3n‘b8n (hmg3bii5 9oi0ni6ig 9u36ih3, i6 qjmoimgn 015300b bn3h-
(jncn mhn36î\3nib gi cfbnb bbn33&o6 9nbi(T'36ip nj>i3366 m5^n9iQ3h Smtn^nib), y3i-
3003606 Qib3h3i-5,i(?36i (hiy 5i63nhm6363pni hmj,mhi5 gp3-e>i93hn ho^9 noi, i6333
^m6 j)h3^ x n i 9 o6gnb 3ohm63&ncn), q)rril!)mgn63'bn i63 jgmh('i30iblj36nb h33m<bn^ni
(hiu 5 i 63nhm6363Qni bo6icn0nb n6^36bn3m6nb <3C3Jii3ibn36ncn - gi6i0 ^ 6 ^ 3 6 ^ 3
J0 mhm30ib^36o6 mhn36(î(jni 66033606 6n8ihcn 33h336gnix^>hxn;>- bmom d0O3h
5i6icn36i'b3 obo6o 5igio6iU303636 3jh3gnb iXC06obi])36 gi un(!)m30i'b93hn qjnoi-
836li|36nb gibSihjbnoi 0^33636 533hgncn Smbn^nib).
dmgngnîijnohn ig i 3oiuni. 5i9mn6i^36i 9u36ihnb 9mhqjci5363'b9n gi 9nbigij>3-
&30Oi bi93i0m higoiijox 3oh(n636cni6. hm5mh^ 63bo, nb 933jl533^goi. 8i5i0noi36ni:
j6m3n03&nbi gi mh5i6m36nb gnqj3h36i5ni(jni; dgn3h ^ 6 ^ 3 6 ^ 3 8m'bihgn 8(336^33-
6o O3noiih3&36 9i3hn3n p3hdnb bnbj)39ib; 9 icn q3moi036b 5iifi6nioi 93'bmgn0nb hi8-
936083 8h3; 33h3g3&n 9 gngihni j0nhn30ibli)36ncn (6ib. 1 .3 4 ); i6ih9m3636 3qjhm 83^
88hi0 8ibib, hi3 n(5 363hj,nnb 939i333pfn6i 93^ni; y3i3n0m6i 3qjhm 9d0i 3hni; n3ncni-
h3636 93^ 6iymgb; b3b^n 5i6ioi36n6 3ohm6369o 9 m‘bihg 9y36ih336b ifooi &hdxo
93b0a)8mhn636n, oib30n, 9i5hi9 gngn qjihcnmbnb 9Jm63 03(^1010360.
33mCDl}n!3hn i@i3oi(5ni. ib3cnn ^n3ob igi3^x5n36n b309nbi63gm9n 9 bnb higni^n-
nbig9 n ihnb 536m^n3nb 6i6n0n gi 5i6bi'b("3hi3b n9 ib, 013 hm030 3i 6 nî^b 9 n>
6o^36b 33nhi^3bm6ib 9(j36ihnb obi 013 nb bib3m6i gi qjm^m3],m^n3n. 9u36ih30ii
jjOibngnî^ni Rhgngnb gi 6n6gnb 9(j36ih336ig (bi5nmqjn^36n), bn6icn0nb9rriy3ih3-
036ig (330nmqjn^36n) gi dgn3hn bn6icn0nb i 9î 6 9u36ih336ig (9|j36ih336n, hm9-
036015 o'bhg36ni6 9ifpig bn9i(pp036<b3 pni ig5np369n, 3gi'66m363n g5 b ^ n b 6i3nh3-
6^3) ihnb 3im0m5n3hn gnq33h36i5ni(jnnb 5i9mbilj)3036i, hiy 5i63nhm63630Oi b3p3j-
- 49 -
8 (3066630)6 &6 6 3 9 P)
Ijnnoii gi 935306(101. ^rri^n^bnb 6363i!)niDhn lJ^(33d33Cn i^hjpg 93(^36936}, 01350

0(536ih3Oii b b 3 ig i b b 3 i 3m33piun 3&b 3 3 3 nb6 i 3 Qncn j ,i 6 :>ai3i>30 fn&nb 3hcnbi g i 06^33
3nhm63&9n [255 ]. 01536^301^ bi3ib3bm 6 mh0 i 6363iifii3hiQ Q3(J3h0n6nh363cni. bb3,
3i 5ipnoiig, bn6ioipnb0my3ih3 01536^3 03nd03&i 93353mb fihgnpob 3nhm636b, 9i 5hi9
33(1 0ni(T>633b 0353360b nb3oi bihnbbb, hm^mhnij 3b ihnb bunmqjn^36nbiOi3nb p:>9>
bibnicn36300 g i 3nhnjncn. 33001^130^36(1 O0i3ghni30ig igi3(jnh363o6n ihoi6 9 i(j>:>p
^3033hivi)3h36oii6 , 5 i 0 m93hi0 3i 3hoii6 gi iihîg 0^636 Bypnb 6iiC36m6ib.
9l536ih33&o 353360^6 bn6ioipob ihi 9bmpmg n6^36bo3m6ib, ihi03g 0ob b33j(|)h3p
iPi03m<bouoib. jhroOiôcn (<33higo) i g i S , ^ ^ ^ ^shftig ihob 5^9ribi^3cn Q3hj gi
Bnoix 6yic0b36ih3369n. 9ionn 3058360)^^0^ 0^3036} Byonb bo(ph9 nb (03bi6i9nbig,
b b 3 ig ib b 3 i bSoj^higohn 030ip5360m 6ob) 0ob3g3ocn. ib3cno i g i 3 (ji( 3036 n 5i0mbi(53-
0Oi i5h30i33 b03C3Oinb 0|j36ih3360n. 0i5ipooiig, Jcmhm3Qiblt)36ob 30503615360b yi0-
im 'bnyni o u 3 0 3 6 i 5i6icn36nb b 33;jlihnb 0ob3g3ncn.
1.2 .2 ic n a ^ n
1.2.2.1 icna^Dhn h06nn6D6n
j)Cn0 i ^ n 0 u 3 6 ih n b o i3 o b 3 0 6 0 8 3 6 3 0 0 ^ 6 3 ^ q jij^ m h o i. |jbh. 1 .7 -3 n 0f*n 530 3po i

5 0 m6 i 0 3 h 0 i b 9 j5i 6 9 n 9 u 36 ih 3 30 m6 ob b n iftig o 5 i6 i6 n p 3 6 i ipn O i^ oh n n 3 3 6 n b 0 o b 3 g 3 n o i, bm pm 6 i b . 1 .3 5 - ‘b 3 6 iR 3 o 6 o 6 n i 9 (5 3 6 i h 3x n oi>-
6 i b i 'b m 5 i g m 3 6 3 6 n b i g i 0 ( 3 3 6 ^ 3 0 0 b ib 3 m 6 3 6 n b 5 i6 i6 n p Q 6 i ^ p n O i^ n b pm jiip^hn
3ohm6 3 6 nb 0 n b 3 P 3 n o i. 0 ^ 3 6 ^ 3 3 6 ( 1 0 ^ 3 3 6 3 6 nbocn 3i 6 nl5i l5 3 6 b , h m 0 p 3 6 o |j ij) 0 iy -
m0 5 0 0 3 6 3 6 0 i o i d n h n o iig bib n ijm ^ b g n n 0m on bni36np3636b [2 6 6 ],
g 3 g i0 n 6 i'b 3 Ibm 6 i 03 hn $pn0i,5nb 0 h i3 ig q )3 h m 3 6 3 6 i 5 i 6 3 nhm&3 6 3 p n i b b 3 ip ib b -
3i 53R5hiqjn3cn 5i63gnbioi3ob gi0ibibnioi3&3po 363h5nnb 6i0i6bncn. 0i5igo-
oiig, bi03igri g i 0ip i0 5i63g3&'b3 .jpnOi^b ibiboioi36b bo‘bm63hm6i: 0m6ii53p3-
m6i Sm'bn^njhn g i 635ii5d3hn higni^nopn 6i0i6bobi (‘biqjbogrv'biaoiihn). oi&npn
b3'bm6 nboi3nb g i 0 ibibnioi3&30ni 5 hd30 o g(p33&o, bmpm uo3nbioi3nb - 9 (^503.
353i(5mh3g lbm6 i 9 n, bigi(j gpnb 0bfii6o biiOi36nb himg36m6i 6pob 5i60i3pm-
6 i 9 o 3ihnh36b 0O(5ig 306o0363gmg, 5 i6 il5Pi6363QOi 3. 6. o^ham-gp^-piOohn om-
3nb 5 0 0 0 ^ 0 (6ib. 1.36 ). gi6ip 5i63g36'b3 gi (13501*163600, bigi^ b03poii03i-
(b(p33oiob ig5O0n ij3b 63n9di6n gi 93hi0O bo<bm636nb 9 m6 i( 5303m6 ib
(nb. 6ib. 6 .57 ). ^pnOi^nb h35on6xn 3ihoi^n3&n (0i5-, 5m6i5o636l53hn g i b (31303-
hn) 5i63nhm63630Oi i5h3oi33 myjiSohn gn63&o6noi (oi6ogo gn6o63&o - &mp<3-
b(5hn0o, j)3hm8om; (5030 gn63636n - pi6higmho, ^©bmggô, 636533gi), jih36ooi
(3ibil536ncn) g i h35om6nb mhnr^hicgnopo oii3ob3&3h3636001.
Ib m 6 ip 3 h g i h 35n n i6 o p 0 ijjh m j)0 o 0 i l5 36 0 n < 3 n h 0 n h g 3 6 i 0 i o i 5 i 6 g i 0 m32,n go 6 o-
pn g i b b 3 i g i b b 3 i 6 i3 h U 30 3 6 n b 0 jm 6 3 5 p o 0 i(5 36 n (© o b m jjp n O i^ n ). 0 3 PI)hn m hn fthbqjr^pn o ii3 n b 3 & 3 h 3 6 i6 o , hm0 p 3 6 n(j
l5 3 p o i6 j i h o b i g i h ig o i(jn n b (b 3 0 n j 0 3 g 3 6 i b . b io 6 l5 3 h 3 b m i, hrri0 g n g n g ib ib p o 6 o -
6n(j, b ii3 c n h o 3 0 i o i l53hnl5 rih n i3<33h3p <b m 6 i0 n , J 0 6 o i 6 3 . 6 .
j i p i j n b 5 p n 0 i(5 b (0n b o i3n b g i 0 i b i b n i o i 3 6 3 p o i 0 3 g ih 3 6 o o i 0 [jn h 3 h i p n i ^ n i , 3 i3 h n b
- 50 -
880b <3^ 0 8 ^ 6 0
Ubhnoo 1.7 jc^^Dhn 'bm&3&n, Brogij/ib ^n3]6n gi gm3n6i 6e>n 9j536ih330(Ti6i (Walter & Breckle,
1991)
6nî^n <3(53bih 3X fr*>

jiSi^hnic^hn 3 >>3i,jmnni03hn yi3nb<33h(i 8 ih ig 9(l>3i (33 ^hmanijjcn
jjpna^n, hn&nhu crmbbihn, qj^hipn^hn 6 3n9 ni6fi ^ 3 , hm93o9nu
B^bn, 3^83635^x^9 SoSng^hn 6(iigi{,36(i (q33hi6digi536(i) b3'bm63hn (330(103636(1 bh>
0dig ih i(nn6n9636i
ljhiTi3nj)Xn 6(10130(1 cmibjdn i6 600130(1 l5hm3d i x n
bicgbxnb 63(1636001 g i &h(i0n 9(16360, qîgn^^hn b i3i6nb qjmoi0ri3i6n ^ 3 ,
99 hi0 n b3tiri6ncn (S^dng^n- 6nigi536n b i3 i6 i
gi6 ihngx^QD)
b]6^hm >iixn Xi66m b 6nigi536(i (ih^nv. ^ 3^Cphm3ni)30n
ihn gxn Dgib5nib n3h9 m- g i Jb3hm6nigi536n), Xi66m b
iC n S i^ . (163(1101(1 63(1836(101 ij,h3oi33 gi 9pi 0363cn 9(}36 ih 330n 6i; gni9 n6nh36b
6nigij>36n (bm0n63ua J3(ii6n 0i 6 g9 iO3)536n
bmcm6fiiyi)
b0 x ® ' i 9 D»'bp3,’l>5f*1X n b93 QOii93i‘bp 3nb- bXDhciojnODhn b3-9(336^33-
"bidcnhnb 63(1936(101 g i "biojb- inh^Qn yi3nbq)3hn 6n, hm903b(H5 bi65hd0(i3(i
X nb b(i99 hignoi (b39dih(igx- 9(16360 yn6336nbig9n b36bnj)3hm6noi
ngi6 b39n339ne3hi9g3) 5,i 9 mnhfi33ni 6
aibnpn 'btnSn^hn 6(10130(1 i6 y3(ioi3cn <bm9fi3hn bih^ygnb 9ih ig 96-
bSnhig 63n036nb "biqjbx0^ (jynb 6(iigi636n 3163 (jy336n
9 ijbn 339 ncn
5 hnco ‘bnanjhn (jynb y i3nbo)3hn gi a)ihoinqjnoi0ri3ibn
9 m x 3 I5n3n 'bidoihnoi 93h i 6nigi536n q)mai(Ti09(j336n ^336(1
J)rA y l5o5vi3h(i b(533(ib BdigiM&n (9i 39n6i, b^333hngi6 3g 365(nb
ihng3(%-'bm8n3hn 15(13(1 6i6pi, 9^hi) 9 (j3 6 ih 3X 'Ti6i 9g 3
'b i 9oihncn
6mh3so3hn 3393^(13630(1 6mh3iC3hn
15113(1 bmBnjhn 5hd{yi 6digi536n 6n63fn3i6n ^3360
‘b ig b x f'O li gi J)hd3cn
”b i 9 oihnoi
3 (npihxn lj36gh(ib 3] 93bfioi 9gngihn 9398(13 Sfchigmfci^
dQfiQh 9 m x 3 lb iO)bxn<:n 6nigi536n 5i 9 mbii|)Xn 5>i63 noiih36x n 30y3m i53&-
bm^ncQnî^nncn ghnb 9(536ih 330m6i
^cghm 0 i("ipn ^ ^ ^ h i ^ h i , 3i]hnb gi&ipn ^36ni6m6i g i jihnb 03gihoi>noi

gi&ipn bnfijsho).
0cru>0n, igni&i^hn âSQ hi^jhxn fchign^^nb cni6ib0ig, ShmfchQbx^g 0(jnh-
93&i i^mbqjQh^pn 6633b g i ^Q S^hi^hi Ol3Q0 i bÔîpmg 6.5 K yjH-'b],
bn0 ippnb 0i(j36ibou>6 ;)hcnig, 0m6036gnp i 0 n6g 0 n 3 bnb higniijni ai^^pmbb,
jih n b bnRjih] nbhp^bi g i nb b x jghm b3 nhn bgj&i, Siphnb |j36ni6ni&i iCD&D-
pm&b, 33s3mhiijni rfbhgQdi gi, hm&mhu S^bn, 6ip3;fc)&nb himg]6 m6 i g i cnm3pnb
biqjihnb bngngo g i bi6j)hdpn3m6i S i ^ x ^ b (6ib. 1 .37 ). ^hminiX 3 i("ip-
9 cnni63 cn0 n (ptr^bp^&n 6ih0mnj063&i ^qjhm 0ipip bnaiCppOD^D- bigijj Sijhnb
v!)36ni5m6i, fi33X D ^n3< oojhm 0i(j>ipni. 0ipip0cni, bi66mj)P3 ^ 3 3 5 3 ( ^ 0 0 1 3ohn-
mgnb, 0mip3 gnbî6ijni'b3 ipn0i^nb 0j>3ooihn L53pnpQ6nbi gi mbnQj^hig
3jbj)h30ip3hn 5^3601 SnhmcbQ&nb j,i0m, 03ndp3&i 5i63nbnpmcn hm^mh^ 3hoi53ihn
- 51 -
90066 (^00)6 &6 (n09 n
Wind
1 1 1 1 1 1 1 1
A i B i C ' D i E ' F i G i H i
1 1 1 1 1 1 1 1
Wind-swept side 1
Shady side 1
'S n o w *
'm a x . 1
.c
Q.
~<o
D
5 1 S o il
o
c
1 te m p e ra tu re
W
— 1.....,J~t; 1
X Psychrophilic s '/ 7 \ Winter \O p portu nistic
fungi / xdesiccation /H e a t a n d \ fungi
o radiation '
Duration of / \ Snow
snow cover \ pressure
6sb. 1.35 (jj^hiCDhn 103360b b]6i 033h bih^yxSn ,jm3mj,hiojo30o cru>3ob363h3636nbi 9b i0n3i-
li^hn qjs^jmh^bnb 6i 3C36i 3(5o6ih3X m?>n*J ^n3] 6nb {,*6*600365^3 95 bnQriii^^h j>0P33&‘b3 Sriigi^nb
q jm h B o f^ i^ .A a^hRênpn cnm^n, Soldanella. bi^^dn; B Rhododendron ferrugineum. bi3b3&o; C
Rhododendron ferrugineum g i Vaccinium uliginosum; D 533* &3fiJ6ihn Vaccinium uIiginosum-ck>
9m9n6i6^m6noi; E Loiseleuria procumbens; F 0^36360 3hmgnh36x 9* 6i(?iho636x 601915^3: G
hm'b3vj x n 9 ^ 6i0n 3i 61536^3360, Juncus trifidus; H Arctostaphylus uva ursi 9* Vaccinium vitis idaea;
K I j^>e>:>hn&36xn 6ni9i{,ob 0^ 3^ ; J Juniperus nana, Calluna vulgaris, Vaccinium vitis idaea: K
Rhododendron ferrugineum 95 Juniperus nana. bi6j,hd 0^130 onr»i3oni6o 33hnmgn 66033630^13619 i 3 -
ynhj&b 33630>ilinnb Sjhnmgb g i 913(53300 315361*13360 bg36ni6 qjbnjhriq|nc3ho bmynj&nb cni3pibb-
3nb mbn3j^. 'biScmhSn 6mh(j3nb jihm6 633*1^3 g i 9 bni6 qj3hgm63&'b3 8(5361*13360 bSnhig *163-
6ni6 cnm3onai 91315330611 91 5160159016 gi6icn l$3333hi£)h36ob ^38^1^339361^ 1^ 30 133 ig&neo
ifob "bi 9 oihob 5i3m3hm&nb" 3053)^. bi9bh3cnnb qj3hgm6o 91(53001 Jihob6i6 g i i8o^,ro8 biO)b3o9n
i j 3ic>ioo j,3033hi^)3hnb 518m b8ohoi 3(5361*13011 519163*1361 (Aulitzky, 1963)
^3C3i5n3hn cgnQ^hn gi iionSionbiurohn fchigr^fSon 3 ibcm&30 n h^nmS^&nb qjQm-

hnbicr^nb. i j di0ni6 QmjjQQ SiBdnj^&lû y i gngni SnjjhmiQnSiOOhn bb3im&3&n.
9 cnnb bQmbQbSn g i bb3iQibb3i ^jbSm'bnunnb qj^hgrnbÊi'b] 33 j 96 nQni 3j>330ihig
5i6bb3i33b3Qn bibn^nybpm 3nhm&3&n. 93bi6i9nbig, 0 (j36ih 3x n biqjihn^ 5 i 5 -
bb3i 33b30 ni- 3 i&ienaiig, fihgnQmQcnnb 3jb3m‘bn(3nnb q^hgm&Qij'b], bi03iQmg,
5i6iOi36nb bi6j,hdpn3m&i 60 %-ncn 6ijjQ3i>ni bi0bh3cnnb 3;jb3m'bnunnb qj3hgmd36-
oni6 839^361101. 'bmQn^hn bih^ycnb 3i(pip0cnni63cn9n biSbhjoinb qjQhgm&^bn
- 52 -
980b 66^06(306 *56 <3^09660
6>b. 1 3 6 Ji^hnb b i3]b(xn hm^mhu ppvibi (mhpnbvy'b (n^hdn) g i 6(yib (i&btyVbnb

P3hdn) phmrib qj36juru.. ^ 3 3 3 . 6 ^ 336 36 11} g (n j^ i9 nb 6 ic n x n ((T0* <3Qhnb p s S ^ n b p ) p i 6630*1
(9jJn pi91jhnb3i) SghnmpQftnb bs6fthdon3m6y 339^3630^^ RhgnomjcrSn &s6o i 63& Xn nhiMbyib
(3sh(Jb36i b3hscm) pi 3i 3^yT*'f'X 9iC>i09omi63oi3n &*6ci63i>Xn ynynb (6ihj336i bjluoin)
3m6i(j393&n.
fciljnQQ&noi oqjhm cnbnQfii, 3ngfvj b>3ii32*Xn ig&nQQ&n 6* Rhgncmooinb qohgm&j-

6n. cnb3036nbcn3nb gidibibniovj&QQni jihn. i8n^m0 i j 33>3mhi(jni 5^150036001
9 i(j> iQ n i, 3 0 9 ( 13 jih n b jp S g :» u x b iQ ib n jh m g , jih n :> 6 *9 5 0 0 3 6 3 11
Precipitation
[mm]
5ib . 1.37 io n S ^ n ^ l a c n c o ^ bn 3 i(ronb bhpibortiS 3hou>p p i 3u36ih33(vn&nb {p 3 ^ 3 0 3 6 1 ^ b 3 p i

<b(j>3ihn iQ33b3n. bih^yc0&n pi 3u36ili33cmdnb 6 ^ 0 X 0 ^ omi3cnb biophnb bi6ahd(yi3m4>i
(pi3l!)hnb3on ^ 3 ) pifihgnQx fihpn0m3ainb gnhgrnfc^ pi ffbni6 biSbhsoinb ^ i^ o ii^ ; 3063i!>i|yinb
33hnmpnb bi55hd|Vi3m6i (ftj,3Qnbb93&i 9^336nb himg36nidi 1 0®C-’b3 9i(nspn li 3933hi(j 3hno\);
gii3nh33636n 6ih3m3dgi 1063-1878 66 (F. v Kemer, 1888) (3ih^b36i b3hxnn). 3^+1360^1 i 3 n6g 3 n
5pmi>ip3hn hbgn^jnnb fehpi; Bpn^hn 6*03^360 (99) (536ljhip3h pi fihgnpmjomb ipij&Sn; n3pnbnb
3i3hnb lj3333hi^3hnb pi633i; 1 9 bne'h3;),b3 6nigi&nb biSoiCni 6pfr)hn ( ^ ^ i t f j h i (3*113336}
b3hioin) (Stelnhauser et al., 1960; Turner, 1970; Franz, 1979)
- 53 -
30366^0016 <VMf>39n
0101300 b3 nhig 5 ig io b 33l536i g i bd ghmb l53033hi£)hig 3j>33cnhig 3U30i. fihgn-

0013010b q}3hgm63&'b3 g i (130030)06 033h3bo3&3n oioi30nb biqjihn ih i Qih^m
3qjhm 0d0i 3hoi, ih i3og bi^Qimg bi6j)hdpn3rnj. ^hm3ni)X Q* bo6j(hm3n i x Q^C^C'
9oini63ai9n 63016360 dpn3h l536oi6n g i jihnb{>i6 g iU X ni- iQo^mQ i j 63^^360 g i
b3 8 5 3 6 ^ 3 3 6 0 6301636b biÔimg 3 x ? i 0 bn0ipp33di0g3 0n3y33&i.
1.2.2.2 b n n icna^n Gi qjn^nbôhn
6nmj)0o9i«yi ihnb 0u35ih3xn biqjihob 0oihmicn0i^n. 6nbn bi'bc>3h36n

3h(j3cg36i 9 (336ih 3x n biqjihnb <b3gi3nhngi6 6oigi50n 0 ^ 3 0 0 5i3hi53036nb ^33-
gi b p 3hi0 g 3 . 050 g i 0 ibiboio)3630 n ico 0 i^ni, hm03 pny 5 i 63 nhm636 x n i 0 (J3-
6ih33Cn biqjihnb 33g636npfn6ncro g i 1^13^53*1001. 6omj)0n0i(5'b3 0mJ03g3&b 50n-
0i^nb 9330 i (join. Oijppncnig, (bm6iQ3ho j)pn0il5n 5i6i3nhm&36b 0"bnb higoiyoib
(£,i0m0gn6ih3 53m&hiqjo30n J)i63gngi6), 0 i 3 n6 , hmî 6iC3j36n gi0miop363poi
(135001630 j)On0il5<b3. bocn6mb bihnbbb, 33i3mhiyoib g i 6i03j3&nb 5i6i6np36ib
2)0 gng6n0ig j)i6bi'b(Ti3hi3b 0Oi2(iQ3hn ^m3m5 hicgn30 n ou>3ob363h36i6n.
n6gn30g3iC3hn diyjGihnb 6nmi£n<hiy). 6om2)po0il5nb 3b ,50^0 ihnb o6go3og3-
i03hn 0u36ihnb 5ih030m 30n0i^nb dipoi6 oib30n <bm6i, hm030nu 30003(13^9
5i63nhm6363Qni qjmon03&nb bnohynoin mhooBîunooi g i jihnb oqjoj^ncn. 0536^3-
36n 3mnj)n0mcn3h030n mhj)i6o<b036oi, 3. 0. 0ioio bb33gob ôO^hi^jhi ih ihnb
'b^bjjig nb30in, hmfcmhnu 5ih30m0u330n 3i3hnbi. hmg3biis 01536^3 np36b 0fbnb
0iC?i0 higni^nib, 050 3b303i ^ 3 0 0 Jb3hnb bii>3h3300n. jihn 9tj36ihnb qjmoi0nb
lb3gi3nhi3g3 hi0g36n03 0n0n03j)hnb bo0ic>03‘b3 ih(?333b i0 600163*1 bibjh^pb
gi 06333b bncn&mb 2>ihj>3ib (6ib. 1.3 8 ). hitf 3ghm 3ip i0 n i jihnb bnfijih3, 0 ncn
3qjhm 3<33ji53hig bg3&i bnoi6mb i)ihf,3i. hm(b3v530O g i 6ipn0i bibnymybcm qjmh-
0nb ©1536^3360, hm03pcni bb33po ib0mi 6oigi&nb (b3gi3nhcni6, ^qjhm 3033^3-
hig o6ihfi363636 bnoi&mb, 3ngh3 3h3;Jl5 x n bibnijmub0m cgmh0nb bib3m636o (6ib.
1.39 ). 0 i336ih 3Oii ^3033(0,53(0 hi0 g 36 o03 ^higjbocn i(r>30 i^ 36 i J,ih30 n 0ij330 n
3i3hnb ,53033(0,53(1^ [73 ]. ,53033hij)3hob i 0 bb3im6ib 36mg3&i 6i03j>n ,53033*1-
63b. 1.36 33hb3nb qmon(yib pmhbxDh

0 10 15 b3£i3nhoii6 jihnb bnRjihnb {,hipn36^n
Wind velocity (m • s ') (Grace, 1977)
- 54 -
980b (^061306 3CJ09660
b^hb ("Overtemperature"; "Ubertemperatur"). q jm o i^ n gb p^hm-

0&n, bmh(jm3i 5n 6hymcg36n g i bnb ^m^&n gb n6bhfi363&36 bnan&mb
gb 9boin hmfcmhu 6]bn, 9bc>bcnb. 33h3mhiijnhb j>bbfi6nb &b9bj>hnp3-
&3pn o c g ^ n (70 W &s 1 &h H20 gO^bcrr1- ^ , 3. 3. 0.1 90 H20 bon'1). &b&-
nb9 n 3hn ^hb5 b 3 nhbunnb q jrib n m p m j^ h n yn 6oh m p n b bb 9 3 bQ36 noi 0i536bh33&b
bj3cn j,bh30mboib6 363hj>nnb bj^n^hn 0n0mu3Qnb 36bhn. 3 3 b3 mhbi5nnb bb^jbOD^ncn
5bj,hnc36b 5i 6bbi3cnh3&nai 30j3di!)3hru> 3iQhnb gbbhpn 036nb6m6nbb gb jbhnb 3n-
hn6369n. ^hb5b3nhiunnb 9b(pbpn n6^36bn3m6nb 93bb6bhR;p63&cbg 0^36bh3b 3&gb
933 dpm b bbi0bhnbn B ycn b 0np3&:>i5.
bnb 3bh3nb &bh030m gb 9ng,6noi, 63fij3&9n gb jjmhgm3b6 9bh(33CfT'36369n Jbhnb
bnfijbh3 9unhg3&b gb ibh&bg fcbOmnbb^&b 0nbn J>hbgn36^n (6bb. 1.40).
a b
Air temperature [°C]
6*b. 1.39 qjmcncnb &3933hb£3 h i bb 3*g*bb 3* bibfnjm(jbcm qjmh9nb 0*(Tiio3oinb 81536^380 dm5 -
3369(10 *9n6p3n. 6 n*g*j)nb <b3e * 3 nhai*6 *bpnib Sm'bbhp 01536^3360 ib ib n io ij& a i 3qjhm 3*(nipn iio3-
( 3 i 3n 309113 839^361101 3*ei*(yi ^ *6 ob 0(j36ih 33&b (01301110 63h^n(v)6n)
(Salisbury & Spomer, 1964) ( 0 *h(jb 36* b^hion). ^ O ^ itfjh n b 5 * 6 * 6(10365 8036*11300
(Novosieversia glacialis) 03enbnb 9b n *6 * 8o6g8n (Tikhomirov, 1963) ( 0* 113336* b3 h*<no)
0 l536 bh3x n oib6bbb‘bfn&bgm3&nb fcnmionOb^n. qjnofTtu 3 6 m,b 9 n 0bngb6

9np3630n hbgnb^nnb ogngrjbn 6b6nQn 5bhgbnj963&b bnon63h 363h&nbg. 363h&nnb
ij^n^hn j,bhgbj06nb 9h3 n6y36b 9u36bh3xn biojbhnb <b3gb3nhngb633. b6mh3g b9
- 55 -
8006663015 <V)tn38 n
Wind direction
y a/
1 / ✓
1 fob. 1.40 jihnb 5i6i6no3&nb Shmqjnen

/ bi3i6nb 9(}36ih3369n. Jihnb bnfijjhj (3
1 r
1 20 40 60 80 100%
59"1 ) 3i 3hn34Flhgn>6 0ih(j30ri3i 6 biQibm-
3i6 9y35ih]3n pbogi b^hbcnn) gi bj-
0Uj6ih;)3n (^3395 b^hboin) (Vareschi.
Relative wind velocity 1960)
Rn U f S C
fob. 1.41 0u36bh3x n biqjihnb bb3igibb3i briO ic^t] I3hm9n6i 9n ix n Shmqjnoj&n g(nnbi gi cpi6nb
hi60i 3poi6i 3n. S hipniiinnb &iPi6bn; u Hihnb bnRjihT; T jiTSSahi^Thi; e Bypnb mhoHpnb 6633*; c
C0 2-nb yn6u]6j)hioni (Monteith, 1973)
- 56 -
980b <3^09660
ij^n^hn 3hnbicri3nb ihnb g i 3 ib>bnioi3&3pn tU S^ h i^h n b gngn cgpjjîijnQ&n. hijj

^ghrn a^bpn g i gi&^hxni 9 uo6 ih ;jx n biqjihn, 3 nori oqjhm j^hfcigii j>i0mbi^-
X n 3 ib 3 n Jihnb fchign^B^n. i;jhmgn6i 9 n ix n 0n0mu3pnb ihm^bo&n (6ypnb
mhcrrjpnbi gi BibSnhmhgiBjjnb ftibnb) 06n9363pni36ig 5i5bb3i3go6i gpnbi gi
r i 3 nb 33hnmQ363n (6ib. 1.41). 3 oj)hx 0uo6ih33o biqjihSn, nj6;)di Qb
&}fij6ihn o r) 6ipib6ihn, bnmjjcnSi^n fciunpj&ncn cn&npn, 6]b^ni6n gi, i9 i33
ghmb, bî&npjhn gi 5 i 6 n 6 ib 6 mh;)&xni< 3ngh;j i 3 9 u j 6 ih 3x n biqjihnb jph^BmS-
15330 3i 0h3 n gi (jvii ig^np^bSn (6ib. 1.42). ^ynb bi&;ph3x J 3:)9 ibipfcibhgi bo
0(336ih3]6n gi i("0m6:H5o636n i3^3ihig g i ^ x ni gii>iCn ^D^^hi^^hobnb g i 9 >
!bni63&0Cn b ^ am ja^ go b n b ifciB . ^hm 3 n j , x n e * bob^hm Snôpn 3 ih i g 3 6 3 i 6 3 6 3 n9 n-
i6n ^yoo&nb bi&oh33pj3o9 ôctoohb^hnb gpn^hn S^hyombi 2-5 fchig^bb ih
bynpgobi, bmpm Bpn^hn Q^hyom&i - 10 ^hig^b'bo 6i]>p3&ni.
25
20
JL 15
cO)
'<&
I 10
£
PhAR [jimolTn'^s'1] Tfiif [5C]

25
20
IS
O 10
RH [%] Wind [m -s1] C 0 2 [Pal
6>b. 1.4CA-F <b*qjb;)03n 6nm2j0n0^ 3hn S ih id ^ ^ n b SOhyij^Q^hn 3hmqjn(v)6n 33hm3nb

^D^DX ^rnoxxnpB ^y^n. A cgncnQnb gjihomb 33h^.»p03^n B qjih-n; C 3v)hnb ^^9-
^3h^3hb; D 3^3hnb cg^hQm6ncnn 5^6^013630^6^; E jihnb b n fij^; F CC^-nb 2>m6u36l|>h*i$n:» (Elias,
1989). 8m6^j303&n ^hm3nj>3Qn ^yQQdnbcrqnb nb. Aoki et al. (1975)
- 57 -
ai3066ff>30)5 &5 ff>aan
^ynb 9*63(130 9l3D6ih g x b*qj*hb 5*6^3301(136001 gngn 5 * 3 0 3 6 * *j3b J33»iynb

g* 6*0*bm3*6n 9(536*h336nb 6y0ocn a m S i h i j ^ * ^ . 6*03j3&ob*b 6y0nb 9bm0mg
6*6n0n *p 6 33b 6o*g*5nb ^39*30(1*893. 9*63(1300 3*113360 0 3*336 36 6nb0b, 9*6*0
(j>h36036b* g* 0li)3hnb 6*6n0*336b. 3 3 * 6 ^ 3 6 3 0 8 380 16 333*30 nbn6n 3(10153*1™
qjn0o,hnb hm0b *bh30D&36.
hn'bm bqjahm b 6 n m i0 n 9 *ly i. 6n*g*5nb ^3833(1*0)311* (53*036 *gm6b 9ynh3 9* 6 -
dn036b3i5 30 g* 3b 153*036*9(^6* g*9ming36xn* hmftRhy 5*h3g*6 9 n(j>3&xn
bncn6mb mg36fn&*‘b3, *b333 81536*113300 b*qj*hnb 0 ,0 ^ 3, 013001 6o*g*5nb 839-
536n0m6ib* g* b^jJtEjha'bj. 013 8(536*h330o b*qj*hn qj*3 *hn* g* 3 n3330 n 9n6 nb
a)*hcnm6n gngn*, 9 * 3 n6 6n*g*5nb j>3033h*ii,3hnboi3nb 96n33630m6* *j3b 9 nb
cg3hb, 6y0nb* g* 3*3hnb 939i5330m6 *b, b^hjjôhib. cg*3 *hn 6n*g*&n, b*g*(5 *3(1*-
I5n* *j(jn3hn*, <71636* 8bm0mg b3g*3nhb3 9* 3 nb 9*bgm630 0336368^ 9 *3 n6 hm(5*
9*^3336090 0* b330n 60*9*50 bncn6mb *(3*h36b (?h3 * 033636*093. 131*93 6n*g*-
50, 5*6^3301(136001 9 nbn b39*3nhn, (503936*. *8(105*9, 60*9*580 bncnbmb
3 o0mi530* *^03(10*. b3'bm63hn ô Siô b 8jm63 (1350^163630 6n*g*5ob ^383311*0,3-
h* 60ob bb3*g*bb3* 33hnmp3n 6 3 3 3 ^ * 9 5*6bb 3*336xn * (6*b. 1 .4 3 ).
9*63h30 cgoli)mi536mb363n 60*9*50 9*15300* hn^mhu ^*h6n bnoi63hn 6*3*gnb-
*5*6, *b333 bocnbmb ^3933^0 i*h53nb5*6*(5- 6o*g*5ob h3gn83 n
96n33630m3*6n* oim30ob b*63h330O, hm3xn(5 b30b 330nb 3 nbn ^ 39 * <336361^
5*yo63*b g* o6*hfi3636b 33^-6*3036*9 bo,*6o03h li)3933h*l33h36b. *3 n^m3
011*130^338 lj,3333h*(j3h* 0830*01*9 3(538* 0 °C-ob j333moi.
Soil temperature [C°] Months
6ib. 1.43 a,b Bnsg^nb tf35eodspm&:>6n. (a) J ^ i h n & ^ x n 6*C*t>m3i6n biojhnb j 3 o0

6nsg*5nb (jO^DhÔhnb 3hmojn0n bigjboenb ,yOn;)h ge>3b neo6mnbnb (Rosenberg, 1974 ).
(b) 6nigi5nb o>3a33hi^3hnb 6on3hn <3 CDi!)3i lJnD&n Rhgner*^ 33hm3*3n (Geiger, 1961). 6nigi5nb
(B ^ h ^ o h n b 3hmqjnQ36n ^hmSn^bnbiaisnb nb. Schulz ( 1960) g i Vareschi & Huber ( 1971)
- 58 -
380l> 3^09660
1 .2 .2 .3 iCnaîMlb l33X 3 6 êm 6 i
X?n3^ n gb b0o6gn 9^9803^ 0153036b. 03O3mhmQm5n3hn qjbjo^^ôb (3bnb

hbgobijob, 3i3hnb 3b3hob (£)6nb6m6:>> jbho gb 6bp3j3&n)
3hcnn3hcnJ03036b 06333b b9n6gnb bb60m])03 (539003636^ hm0036o(j 93od03&b
j)b5hd3C03b hb0036o03 0("3 b6 jôhb (63n9nb ^hom gn, yn63nb 33homgn). bb30>
(539003636b bh &bbR6obcn (135x ^ 30 0 bbbnbom. 0bcnfcb6 &b6bb3b33630fib j>pn3b-
ôb j,ppi6bC3hn gb bb6&hd0O3n 15300036360 [273], hfn0p3&ou 03odp3&b 6o6g-
o6bh3m6g3b bOibb69330O6nb 0b5do0‘b3u i)0 (6bb. 1.44). g30b0o6ob 0rndhbm6b
P3hdob 5bh330m 5b5b3nhm63&b 3bncn ^b6boi3& x n 9* 66300 33hnmg36nb
3hcnn3hoi0(,n6b(J303m6bb (qgmli(m33hor>i0n(b3o). 6bort3po cgb'bb bhob 5bon6m6ob,
bmpm 66300 - j)b(i0336ob ^homg^&o^. b3oofo0 q)mli)fT'33hnmgo'b0b 03g0n3bg o»b6
bb0b3b «j*3h0n33hotngn(b0o. hn&mhu B3bo, cn3h0m33hnmgn ghmSo Rb3mhfi36b
gmom^homgb: 3b3hob 0hjbn030b btf>633b bhb 9b9o6, hm^b 0b3
'b36o(j0ob (03bb6b0nbb0, qjm^r^homgob 0bjbn030nb), bhb030 m06b3 53fib6; $ j d -
^3hbO3h30O 0o6o030o jjo 9536b 0br»0m0 pb0nb 6memb.
0b6b0 53m5hbgnx <^3936^)3 ajnvyn^hnrngn 690b 5 b60 b30m6 b3 o 0130(139
&T=o.5 K
6sb. 1.44a-d ig h ^ x oSmfoWn

50fT'6 xO hn 'aCDiiJiiy'D^nb hjimeb^h^Junb.
(a) dmSiiijdj&n, (b) nblyn-
hnôn Ri6j63h36n, (c) ’ibQo6 mQr^trr)n<\ gb
jQiljnnicmjnjhn 9 m6 i( 539;)&n ^hmSnbb gb
fihg. i93hny>lxn3nb, (d) (ph9i t>c>3;)dnb
g>6i 03j jb63&9n 3i6j,6ignb (Vbfnam3o&nb
5 i’b md3i. gc3teDb(}nnb i 93co-
^ 39^ 9 fTH5o9x ni v>C3n6nb 3 is(W i
(Lauscher, 1981; Webb, 1986; Roeckner, 1992;
Webb & Bartlein, 1992)
- 59 -
90066600)6 S 6 f609D
ni53CQ6i- i 9 n^m3 ,i)hm3nj)3&9n ih ihnb îhgjig fciSmbiôn Spnb ghm; ij ^cnQi-

^hn hn^9 nb 9ou>3ihn 5i3mbi^3Q3dii gpnbi gi Q>i9 nb q ji^d n b ^hcnnQhcnQm-
6il53CDfr|6 i (ppo-c-iaDhn ^pngi^n). 9 ou>9 n g i 9 i(ni0 30i vi)m36'b3 bncn&mb dpn^hn
5i9mbbn336nbib 3b hn^0n 93ndc]6i j^mnbi^mb ^3933hi^hnb îphgbig gng
I33i 0 3 6 ig p i 6 i 9 n. ^ h m S n ix 9 oio 6 9 n 6 0 130 0 Bcnb J>i6 9 i 3 0 m6 i 3 n 9 jj 3 6 ih 3o ii ^ 3 6 -
33hi^ 3h3Cn h3gn9 n ibôirti: 0 i 9 noi ig^npn ij3b b id in g dgnoh yn63ib (- 10° C -
9g3), bmQm g(j»nb biioi366n Sxjhnb 6nigi2>nb3nhi 9 hnb ^3933hi^3hi 30 °C -b
id>3 9 ij) 3 6 i ( " 9 m6 i( 33C3 (Ti6 noin yn 6 336 n b j>pn9 i l5n").
53rij)hiq3n3on g>i63g36nb bhgiboii6 ohoiig n(53CD6i gpnbi g i p i 9 nb
oii6iqjihgm6i Bpnb j)i 60 i 3CfTi6 i 3 n (b3bm63hn qjm^mSohnmgn^n) g i hiy oqjhm
9 ipiOni 5i63gn, 9 noi 35301 ihnb nb &i9 m bi^xn (Bib. 1.45 ). 3m0 i h x n 6hnb
^ Sn o i &36nm6nb ghmb gpnbi g i p i 9 nb 9 m6 iy 303m6 i 5 ygo6 i ("SmQihjon p i -
9336n gi g c l3 3 6 n " ) . b i 93 i c m ^63936^3 ’b iq g b o c S n gpnb b i6 5 h d 0 n 3 m 6 i g iib -
0m36ncn 1 6 biioini, bmpm <bi 9 oiih9 n - 9 b m 0 P ig fl biioin. q jm ^ m S Q h n m g n b b^-
“bm63h L53iCD6igm6ib oii6 3hoi3nb li3933hi(53hnb b3<bm63hn l33CnC3 ^);>b (bo'bm-
63hn cn3 h 9 m3 3 hnmgn(b 9 n). J>i5 b b 3 i 3 3 6 i 363hj>nnb 6 i 0 i 6 b9 n g i6 io 5 i6 3 g 3 & b i
(bigiu n6bmpi(jni 9 ipipni) g i 8i(T>iQ £^639366 (bigiy higni^nnb n6^36bn3m6i
gi6i(yii) 3 mhnb n6333b kpm&iQ^h ijnhj)XM3ni^ i|j8mbq)3hmbi g i (*1533633630 Qb,
i8j,3ihig, &i3C35ib ibg36b 6i0oj36nb j,i6i6no36i(b3. 93go&ig i 9 nbi n j9 5 36 i
99 hion g i 6iQ3jni6n 33hnmg3&nb 9 m6 i(5303m6 i - 3nghm33hnmgn'b9n.
fob. 1.45 gm^mijhnmgn'bSn b b ^ ib b ^ iiS a e ta .

6i6oenb 5130363 l^’bmBah Cbop3 b^hicnn) pi Bon^h
(J33g3 b^hioin) gn^mSohnmg'bj (Downs & Hellmers,
1975; Salisbury & Ross, 1992)
- 60 -
____________________________________________________________________________________________________ s a b a o f n & a m 0 3 ^ 6
2. 6ib0(lh6iGflb anbaihObi Qi aahXfi

6n301(lDhD6flb 6ihanj36i
2.1 6ibanh6ienb b30i Dxhoean

2 .1.1 o a w i b n K T O 'b n
i g h ^ x 63 mcm 5 nm]h SohnmgSn, hm g^biu 3 nh 33 c 3 hmiphnmli)3 6 9 n (ihj3&:>j-

O flh f^ n , ^m^nhgnb b ijo t f h r ^ n , U jn ^m & yj^hn^dn) yion 6Q 36m gi 0 3^ 10)^ 06013-
Protocellijiar
organisrris
Phototrophic
organisms Eukaryotes
| ph5o ? o p h e bsyis | Algae H| her P|an,s
Atmosphere C 0 2 NH3 H2 N2 --------------- 2222S--------------- ».

Respiration anaerobic _ 0 2 N2 aerobic
______ Eukaryotic algae

Relative
contribution 1 \ / \ , \{ .
to total Photosynthetic \/C y a n o - \ /

photosynthesis bacteria * bacteria ' \ /ip la n ts
1 ' / \ / v
Relative
concentrations
of 0 2 and C 0 2
and solar \x r 21 kPa 0 2
ratiation
arriving at the
earth's surface
Currently 35 Pa C 0 2
t------ 1 1-------1------- 1-------1------ 1-------1-------r
4.0 3.5 3.0 2.5 2.0 1.5 1.0 0.5 0
Age ot earth (109 years)
6ib . 2.1 qjm^mbnEkn^'bnb 33m03(jnnb 3 obid 0tn 6*603360 (L aw lo r, 1993)
- 61 -
GabBOfn&Ôli 8{ % 9M a 56 896^0 603Cn00fo0?»0b B^ff>8nd866
b^hig iJonDhn 0306hi536n, iâmLxggQhm bhxî© ih 330(5i3gi 3^656^06. 311(133-

QymqjnQn ^^©mbGg^hm ipg536ooio nym, g i Snghmbq^hmu d ip t ] (jmoi g ^ b ig b
3 Dnl5i 3gi- e 3©i3 n6 i b 3 bnumubonb 363(1530033011 Q* 0 iO3hniCDhn 6 ib i gm^m-
bn5cn3'bnb 36ihob 9 jm6 ] gm^r^^rnohrncrôai mh5i6ob0360i (6ib. 2.1.) 83-
J063b. q3mombn5cn3<bnb mho bi6mppim ^hmg^j^n - 5i3myricgn0o gi65&igo gi ibrv
0000(136300 6ib8oh6ign 3hoi6iohig 36n33630m3i6oi y 3 X i l5fTHjb:>pn Pih5i6nb-
3 nbicn3nb. gi6&6igo 5ibgi 6 n6 i 3ohm6 i b35oiJ3nb - 6om0rri5o3ho gi653nb y330i-
^ 3 3<333JiiDhn qjmh0obi, hm030O(5 363(15111101 3<bh36330yrTiq3b 6o30i03h3&ioii (5300-
bi gi 33(13930° b^h j^hn b 036ihfi3636ob Shm ^b^b. 6ib8oh6y036o ojî b36oij-
3fib 36033(1^03(1 b^b^hiOiQ gi 9hi3i0 bb3igibb3i 6nmbn6oi3<bnb bibîh^m
6 i3 h o ii0 . 3 3 fn0 3 ijn i b0303oinb 3m(i0m cgoo36ob (q j3b 3o b i g i p>3hmb 0 jm 6 3 0u36i-
(133611) 0i(nipn gnq)3h36uniunnb 0n0ihoi3P36noi bmh|5n3cg36mgi - 0(536^300
3hmg 3 J^ d 3 0 rrt6 i b30 3cghm g i 3cghm o b h g 36 m g i. q3iri^rnbn6oi3<b3hn i j (i)o3m6ob
0J1T163 03 g i 3 n6 ob 0(336i(i330m6i 5 3 3 30 0 6 3^ 3lbih0i<bih, i0iboii6i33, j>i636y33^-
0fi3 0bihg 6nm0ibig, hm0303nu ijj30 X nh3^D0ni 363h5nnb 0ihoi3nb gi0ibibn>-
01363011 bib3 - 6nm363h5fii.
q3m^mbn6oi3'bnb 3hmu3b3n bg3&i bbn33hn 363h5nnb 3oo 6cnj0i g i 0nbn &ih-
giJ 06i 6i3hoioii jo0o3ho 60360b 363h5nig. 6ib 3nh6igob iom03ho 6m6nb ym gx
5hi0b 3 0nn(Ti36i 479 kJ 3m^36(jn3hn 363h5fii. cgm^mbnBo^bo © m o^b bn6iov-
0noi 5i63nhm&3630 cgm^mjnanjh 3hmi53b36b, <33(1036^30 3hrnu3b3&b, hm0p36nu
ih bi^nhm3636 bn6iaip3b (3.6. bn6630nb (13^(503611) g i poqoboob 3hm(53b36b,
hm0036ni3 3bh3633pymcg36 6ib 3oh6igo bi Qi gi65&igob (530ib j0mhm30ibo36bi
Qi 0ioi 5ih30n0U330 3i3hb Smhob. y330i 3b 3hmu3bn g i0 m y ig36 xn i 8o6i5i6
gi 5i(i35i6 qjiJofT'h36'b3 g i i0g36ig, oinoim3x 3ioi5i 6b 833dpoi gm^bnB-
013'bnb bi3hoim 5i0mbi30nb po0nonh36i. 830935 0033680 g n li(rTibn6oi3,bnb 6nm3-
63(1530030^ g i 6nmjn3nnb biiooib3&o 5i6bn030n njojbs 0bm0fng 0ioin 3yn0m-
503(10 06n8363pm6nb 5ioi3i0ob6o63&ocn.
2 . l . i . i . q 3nc>njnan 3 hn 3 hmi33b 06 ri: Dsohannb i33e:>
qjrri(j)(Tibn6oi3bnb 5i6bmhunxD^nb 330 (13x3^0 3ohm6i i j0mhm30ibo36nb 803(1

0bnb 5i0mbbn33&ob 3oo6oij0i. i 3 5i0mbbo33&nb (13(533^(13610 bo6icn0ob b33-
j^hnb 600130 g i 03(13 3663680 8oo6oij0nb 0yjbn030ob 0jm63 j0mhmqjn036n (6ib.
1.32) g i Oii60b03&n 30ibôg3hn 305036^360 (^hm^n6o, jbi6ofTi<3O0o)1 (10180360(5
8 0 0 6 0 1^336 0bob higoi^nib b3ijohnb 03(13 g i 30ohioobq)3h 3663680. (50i6m6ij-
(i3hn36o, 6001300 6yi00y3 6 ^ 3 3 6 0 g i iho3^m0m6ig36o gi8i036ooi 8 3 0 ^ 3 3 6
6o0o3hmli)3O036b (qj0inuni606b g i qj03R3hooihn6b); qjmorômohmqoon 6001300
6 ^ 3 (10 3 6 0 30 - 6ij,ji3hnmj0mhma|opb, 3o o 6oij3nb dnhooiigo bmoooi b33J(jhob
8rrih330 600130 36i68o.
5i3mbbo33&ob 8oii6oiJ8i g o g ig ii gi0mj)O936xn q}morribo6oi3b3hig ^ 0 3 ( 1
3n5 0 3 6 ^,oii ^ 6 ( 5 3 6 0 ( 0 15 0 ^ 3 . o6o36b o3hn 5 i6 io i3 6 o b ib i 0 3 o5 0 6 o o i i 1 ^ 6 ( 5 3 6 ^ -
I50i 03odp36i 5ib g 3b q]m^mjo0o3ho 3hn(j3b36ob 0i0o0oooh3&30o qjij^mhn (6ib.
2 .2.). jomhmqjooob 6i^036m6i, 3my3OQni0ri i6 0130*0 83cg3ho0m6ob 0(536^33680
(„j0f,ihmbo“ ), ym330oi3ob 36o 33630m 36ig ij3300i36b q3fn(!)(Tibo6oi3bnb bnfijih3b.
ib3oio hi3 03o3R 633i 6mhRo qjmoioob 5i3pobib g i i5h 3ai33 qgmoim0i5336nbib.
qjmoim0o j0m hm bxn bg3&i jomhmqjocnb gi30nbib(5, hi^ 83odp36i 5i63nhm63-
- 62 -
6ib . 2.2 g*9ning36xO ^>;> imÊStyi^h
qjrymbn5oi3'bb, jpmhpicgnenb 3;)du33c>m&ib;>
gs bn6*oiQnb n6#)6bn3ni&:>b Smhnb
Theobroma cacao-dn. jpfomc^j&n
C02-ncn 5^53(13&nb Snhm&jbSn (2400 jj.1 1-1
3i3h3n) (Baker & Hardwick, 1976)
&Xn nymb d50O56 35p50o 56, Sohojnoi, d50O56 gi6:>0O 55650136001, 0n6]hbQ3hn
65056bob 95^333001, 85363 55'b3&ob 801^83936001 56 30(^300 n6q}3;Ji5nnoi. g i
6mQmb, jpmhmcgnQnb 93030(50010 33od03&5 63630>ni3hig nymb 930)3(18060(136300,
(101501(115 }0ib 595000 i ^ b am'bin^hn 56 y3ooi30o qj3hnb qjmcn036nb 3J0163
815365(133680.
q3mlt,mbn6oi3'bn bmhL5O30g36i jpmhrn3pibo)360n. j0frihm305boi36o mh9 bj,n 8386-
(156001 338mbi'b(3'3h30 n 33^3QDCn m(i556m36o5, (1018036015 015305601 l>o>hm353 o
830(55336 ojncî^ng^h 9386(1565015 boboo85b (<33010,0^06013^3(159 5 ^ 3 (1 0 ^05836-
0)360013(101), hn6rnbm036b, 305bo>o93h g68-b (j0mhm305bo>36ob £,3601811) p i bb35
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5 ih9 inj963&i 0)nCi i rnn63 ban.
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mhn 3058360)3110 bobo>385 0156808933(1x^9 3 m656 o03 m6 b bo65oi0 ooi 55630(10163-
630 h3iji5n363n. <71001013x0 85015560 9535380(1363005 bo6;>ai0ob 83831136 305-
8360,-150001356 3oi3303jboi56 (LHPC, 65b. 2 .3 ). 560)363(10 3058360,360 (jomhoiqjo-
0360, <33010300,060, 03030160006360, 2phoi0,060109360) 801560^5336 9 m b x bo6ioi-
0ob ;)3560136b, (1018036015 338932, 5 59 531538 5 b ^yju o m (5360,11b. <30101mbobo,395 I
(PS I) <300015363^3(159 303(101 d330Oi g i 93oî3b 5i6b^33Oih3630 bofajoohx
55(13801(53580 3ymq3 3058360^; sd 3m8303jbob dohooiigo 30183016360,05 jpoihm-
03000 a. PS I-ob bih^jtfooi I536^hb 65h8oi5g536b j0mhmq3o0-(jo0ob 3m9303;|bo
9 oii6 ov)9 ob 9 ijbo 9 i 03 ho 303001 700 63 -ob 365680 - ,,3053360,0 700 “ . j0mhoiqjo0-
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00b, 30gh3 Rhgo0ob 03010103630. <3mo>mboboo,385 II (PS II) 8301553b jooihoiqjoo-
yo0ob 3m8303jbb 8 ijbo 9 i 03 hn 3 oi>6oij9 ooi 660 63 -ob 365680 („P 680 “ ), hoi930 -
0(5 biboicng36i Jomhoiqjnc b-b 95 jbi6o>m<3o036ob 3qjhm 95(j>50o 838(5330016001,
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ob 30,000^5(50580 (jbi6O)mqjo03ho 150300, ob. 01530 6.2. 1.1).
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305836010b 801033300956 303jO)hm636ob 558010^01(1(565 9oibg33b. P 700-g56 558016-
oii3ob3qj036xn 30 3^ 0 16 36 0 b8ihg365 NADP+-ob 5pg5365b, boi0m j0oihmgjnQ-
- 63 -
G6b90 rt66 ^ 0 l> 9 n b 96 (f)0 W 9 3 6 6 ^ 0 603 m00 (^3 &0b P 6 rt9 nd 966
fob. 2.3 ^n0ij)mne3&nb 9]96hs63&3n jCDii^roBi&nb dhifjbSmh^nb &>9Ay’3D&Xn bj]9i. WSE ESypnb
eâacxn ^ “brian; PSII q)m,jmbnb,K)fo II, P680 e> 5oib^r>r)n6m6o6nl) itoîjnhD&n Qa gi QB; PQ
3C^Ws<T'Jn6m6n; (jn^rijhnian 66/03 i|m3 !3Q3jbn djny^b gjhjgmjbnO) (FD), (jnlynjhm0n b6(Cytb),
&mj>nhgnb:> g i hyiBnb (536^1360 (FeS) g i (jn,yijhn3n f (Cytf); PCY 3oi,JiJml5ni6n6n; PSI
bnb(S30i I, P700 g i FeS; 6ntnxn goh3emJbn6n-NADP<- h^g^jî’bi; CF ATP bn6cn3î<bib ynqjij-
limhn (Hess, 1991)
nb ^(pbi9j,36i9 bbôhm 303^(11*16366 ndoo3h 6yonb qjm^mpn'bn b6 3p3j^hm636nb

bb3h hpi63pn03 gm6mho - 0h5-, H2S qgfn^mi36m^hmqg30 6ij^3hn3&9n. qjm^mQn-
V ib 3hrii53b9 n 01^30630309365 5 ^ 3 0 13 3 3hm^m63&n 95 5b0Finyfnqjb gb65&bgn.
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636b ^npiimngnb 0306hb6ob 0 ogb g i 6bh3 <b3gb3nh3&b Bmhnb 3hm^m6x n J>h>-
9fi35 ^nb 03j 06 ib. 3b pH 5(1^9036^(1 3<bh3633oymogb 363(150001 ATP bn6oi3<bb.
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qji^b (ATP) gb 3 5 6 5 6 ^ :
8 hv
2H2CH-2NADP-H-2ADP+2P j------------- = 2NADPH2+ 2ATP+0 2 ( 2 . 1)
P 700+ P680
bn6ioiQnb 0 ou.6 oiJ9noi 5b63 nhro&3&30n g i P 700-nb 55^*63606 83935^9 dnb-

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- 64 -
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n(59116 i>5h6oiJbnpnh36ib 033(1936^ RuBP-jj5h6mjbopi‘bob 9m656np30i6ooi. <30101-
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(3op3&ob 50%-b 5(^633^. i)5h6oijbnpnh36ob 3hmg3;)oo - 3^3665680(16590560 9oip3-
33p5 65h9oiJ86obaii6533 o6p3&5 3-q3m6qjoi5po(53ho6o6 63530b (PGA) oiho 8oip3-
i3pob 65h9mj96ooi. oiocnm3po 59 80^32)305015556 9301353b 65b3oh65gob b56
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- 65 -
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5365 500(53(15093309-3-03016035^)5690 ( G A P ) ATP-6 g i N A D P .H 2-0 6 801656003-

m6ncn 8069065113 (153936093 6503363(106 5530001. 6ib 9 nh6 ignb 5pg536o6 653(10101
556,5010365b 563010 bib3 5j3b:
nC0 2 +nAcc+ 2nNADPH 2+2nATP>(CH2 0 )n+2nNADP+2nADP+2nPi+nAcc+nH2 0 ( 2 -3 )
^hoofb 0103016035,30 G A P 559 59 0 6 65b 3oh65gn b b b 3 ig 5 b b 3 5 6nj,hdo6 ( c3 - c 7)

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0l5365h336nb 33(139 36 0 0 03363360956 C 0 2-ob 90033'boob b 50 35036n oi, 50 5 5 6 0 6
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bo 8095 900336006 8393559 33od036i 4%-3g3 0o5[p6nmb [25 ],
6 nnrjn0 n3hn 3016(536^(15(50300 83^56068360 dohnoi5059 3033^3(105 958685(13
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- 66 -
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in;
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bxsnnb 3nh330ign S h m g ^ n ihnb ih i bi36ibdnh6ig n i6 n 3 mo3i 30i. ih i9 3g 85^36-
d0ih0j)i3s, 6ib9 nh&ignb 4 i^mSnb 0jm63 pnj>ih6ri6nb 0 g i3i, hm33pn(5 6ih3mnj8-
636^ ojmbq)m36m03nh33i v!)nb (PEP) P-îhbmjbnent^bncn. 0138155 6ib0 nh5yp36i 0g3
i(P9536i ym330cn3nb bmhun30936i ^B^mbmqjmbqjioohn fc’bnoi.
C02-nb 9i^ 33n h 3& i PEP-oii6 qjihoimg 6i 3hy3Q36x n 6nmjn3n3hn h 3ijunii
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6nb, nb3 30i(j>c3bn mhj)i6nlb036nb 03op6mpn'b3nbicn3nb. i0 6363ly\*)3h;>9 5i 63o>-
i nbD&DCn h3iji5nnb i3m bi3i0n B3h(ijn0n9i6 j)i63noiihgi gntphbrnBob 3gi336ob
jjbnb (hm030bii5 8gi36d 8ih 8gi3ib 0m03530i 3 n 6bb9 nh6ipnb 4 i^mSnb ihb3&ri-
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1. mhbiq33b3hni6n Shm^bnbi, hn03Q3nij C 0 2-nb qgnjbi^ni (C^Shm^bn)
9i 6 ib 9 nh6 yc36nb 6 ih 3 mj3 6 i (C^-ihm^bn) bmh(jn3pp3&i mh, bo3hu3Qig 3hoi0i-
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2. mh33hnmgni6n 3hm(53bnbi, hm0pob 3 3 8 3 3 0 ^ 0 1 801300 pi0nb J>i60i3-
0fn6i9n bg3&i C 0 2-nb cgnjbi^ni gniihbmBnb 0gi33&nb 6ih8mj06oon g i 9ioin gi-
0ihi53& i 3ii3mQ369n, ppnbnoi 30 bmh|5O30936i i 9 8gi33&nb 932)ih6mjbnpnh36i
gi 6ih0mj05n0n C 0 2-nb fiihai3i i0 i33 3jh3936ob j0mhm30ib(t)3b3n 0m 039i33
^b^m ’bmcgmbqjioohn fc'bnb h3ij|jn363n (mhj)i6 3 p 3g i3io ii 03l>)i6mpocb8o
C rassulaceae (0b3ji6ib3&haii) mjibnb 8 1336 ^336 30 , C AM -03liji6fn0n‘b0n). 2.1
Ijbhn09n 6ih0mgj)36n0ni b i0 n 5i 5bb3i 33&30n 83^61^03110 jjbnb (C3, C 4, C A M )
8ibibniai36036n. 3i536ih336n, hm00369ni5 C 0 2-nb qjnjbitfni gniih6m6nb 83^33-
6ob fj'bnan bmh|jO30936i, mhn 3hoHi3hoibi6n6iip0932>m b^hiô&nngiB ny363&36
3hai-3haib: 3ijbn0ip3hn ibn0npi(jnnb 0O(*>633i C4 6i0ibm 3i6 0(536ih3369n g i
6ib3 nh6ignb ibnOngi^nnb 3hm^3bnb 0mj6n0m6i, 9 nbn Scib^n^hm bi C A M 0^3-
6 ih3369n.
6 ib 9 nh6 ignb ibn3n0i|jnnb bi3hoim 3hm^3b9n C4 3hm(j3bnb 8m6i6o03mbob
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93qjihg36i &50mg53&i. 01536^3390 0ibignbioi3nb 80^3630 13C-/l2C 03qgihp36ib
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hiu 3q3hm 3ihymq)naini 5 13C, 0ncn 3oghm n6^36un3hoi 5 13C gi5hrn336i.

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(-23-gi6 -36-093, bi33i0m g -27%o), 3ngh3 C 4 0U36ih336b (-10-gi6 -18-893,
b i93i 0Pi0 - 13%o) [204], C A M 3i536ih336nbiOi3nb 3b 0ifi33636036n 03hy3m6b
- 67 -
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[ 114]. &*h9* * 9 nbs, 8 13C-nb bnpng^ 9*9013093630(1* *6069133, 5*6^3301(13600)
IjbhnQn 2.1 CC^-ob qjnjbiynnb bb35g5bb3* 4030b 8)0163 815365(13015 pibibnbovj&i (Black, 1973;
Kluge & Ting, 1978; Osmond, 1978; Sestak, 1985)
3ib>bnicr>363c n C3 Ca M
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po 03'bmojn(vi, 558- 905 55845(10 im636nb 83lbmqj09n. bsSihifcm
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(jmhn (b36b(j(u>yi: CO2) 1)06639380: PEP
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q3moi03&nb 5i9^.hn j,oi6o6nb 0(153003 5560553630° 9Q9m9oj36n 3jh3g36nb 55(1-
939m 9bb3d0n j 0 fnh36jn9 x n 33(139 360b 63nh53n6nb 5hb36m6i (3.6. 3(15615-565-
^m9 ni; nb. 65b. 2 .6 ; „3hi6ij“-53nh53n6n). 50 g i 03‘bmqjn0nb bb3i 33(1393690 C 0 2
33^390(1936} 3nh330ig ij^Sôihb PEP-b 0 gio69 g i 3ib 6 ih9 mj96 ncn (nb. 6ib. 2.12
dshybjBs dbihj), hoi030o(j NADPH 2-g50011)0936x0 qj3h036^nb 0505^93309(10153-
65(b ib bi 93i 036fioi i(pg636 i 9 i 05îg, 0139^5 ‘br^nohoi bib 3m6i 9 n b il|>hi6b 3mhl5oi
qjrnh3ib 65h9oi5g536b ib S ih ^ ^ n . 03'bmq)n0ob 3jh3g369n 3b 6i3hoi36n 9mnb9ih3-
6ni6 - nbn6n 55gi9mdhi3g36ni6 9390100336 33(1393690, hm930oii j0oihm305bl336-
9 n b33i5ngni)3hn qj3h936^nb 9 m6 i 6 n03oi6 noi gin903&ni6 3nh33ivj)ig g i CC^-ig.
65h9 mj96 n0 C0 2-b „Rin^Qhb“ hn630m'bmq3mbq3i^)n g i „ 5 i i 95'bi 3h36b“ 336(401^01-
ojmbqji^hn 5'bnoi; 3nh33i(jn 30 6h36g36i 93'bmqjn0nb 33(i39369n g i 5i9oiny36o-
6 i PEP-nb h35363hi(jnnbicn3nb.
35h6 oii63nghi'biboii6 93 j)539nh36x r' PEP-j)ih6mjbn0i<bi, hoi930 n(j bBhiqjig
5ihgij 96 nb C 0 2-b H C O j-ig, 93d)ig D<33 diiD^r,5- bnb,j39i 9oij9og36b (101501(1^
C'07-nb (|)396ri. 9 ib 933d0Oi n9 o9 ioib (101501(113 HCC^-nb g i C 0 2-b gi6i0n 3016-
U36(jhi|3n36nb, ib333 3i3hnb 9i(j'i0n ^3033(15^;)hnb 30(10163690. ib3 hoi9 , 63^01-
cgn0nb 331130360 601^336 33h3g3mhnb369n ihb36 x oinov)9 nb 90130 C 0 2-b gi
9 nbn bi3oi9336biunoi 30i6u36v|)hiuni 10 (Jl"^-9g3 3Uo9i.
03cbmq)n0nb 33(1393680 cgoi(jmbo6cnj3nb ihihbo6m6nbi g i i5h30i33 n9 nb 5i9m,
Grasses Dicotyledons
6>b. 2.4 C4 9u36ih]36nb 553(1(530361 Rhp. iSjhnyib 6036111330 biqjihSn p> 6nbn gidmyig36x361
53R5hiqjnx 6^63936^ 3 . i(«6n363eni C4 6(536ih336nb %-n 6npn 6>£ibm3:>6aii qjQmhxJn (dih(jb3-
61 bohioin) p> mhQ366ni6 6101601316 3^36^33660 (61(133361 b^hioin) (Teeri & Stowe, 1976; Stowe
& Teeri, 1978; Ehleringer, 1979). C4 6(536ih336nb 05033 3hm3mh(jm ic^m^ncm 33hoi3i3n(j
(Collings & Jones, 1985). fihg. icghnyib ihngx (13500)63630 (Batanouny et al., 1988), 600(13 iljniSn
(Vogel et al., 1986) ei isbîcniSn (Hattersley, 1983) >61^0536300 qjcoihnb 40-90% ihnb C4
8(5361(13. b39nihnex 5693690 2300 6-693 35% bib3R&3&nbi C4-oi, bmpn 2300-3300 6-893
gniUi'bm63n - 20% (Ruthsatz & Hofmann, 1984)
- 69 -
6 ib 30 A?)5 ^nb SDbaartDM 83 fn^ 0 6030 )n0A 060 b B,^fSS8 nd 366
hm8 >9 33(139015 jemhoOo5bv536b qjm^mbnJjoiQ'bnlj 556601(115030365 aj^dcn^cn

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bm&5 555660501. 35h6oijbo0 oh36ob 3 b(j>403qj3j^ 3hn6i 86(13363001356 3016331136,1)30
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C4 8(5365(13015 3fig33 3hoi 33nh50>3bm65b Bih9mig536b 5,bml3nb 6n05hoi 65010
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5hnb 55630(101636300. 6301(13 3bho3, 36g 5 5(po6n96mb 58 815365113015 5h5b5b5h6 o3-
001 015306363(1365(5 - g5&50n b (115033560)560165: 'bhgnb 33(1001980 0)3333h50L|hnb 95-
15385 5-7°C-‘b3 ^333001 35hyoi0300159 8m;J9 3936b 6501 55630015(136563 [ 186 ],
93b5d0m5, 03001333(10 o^^bSmh^nb o63o6 nh3&nb 558m. 5800105 5563ohoi&36x n
C4 bsb 3m&50i5 h5mg36m6nb 333150(136:. on6npn 89h50n 3530b 5g5opbi8ymq33Qn056
6395(136001 15030 05 ,5360560 11350016360b 3o05hoi3036no» (6ib. 2.4 95 65b. 2 .5 ).
560b 553m5, hm8 g 5fihgo0X . 11136056 95 1503 59500380 6565605 C4 bib30i6i0ii
3bm0oi0 8(50(13 h5mg36oi65 (P a sp a lu m con jugation , C y p eru s sp p ., S p a rtin a a n g e lic a .
E c h in o c h lo a c ru s -g a lli) [277 ].
C4 3i536 5h 336 n qjihcnmg 553(1(530363005 3hoi60n56, 55 6b 5330iti36ooi ‘6 5 q j b x n6
3(10160056 8(536*113015 g5 33603(10030)010)00)366 9oihob; 65601(1 ob 3hoi6oo5636b5 95
5301030,53680 0 60 60 083050159 5 3 6 3 9 3 6 5 . 9 5^ 50 & 3fij36b5 g i 6 336 8 0 , (101501(1(5
63b o, C 4 b o 6 ghm 3 n 5h 3ooi5hg365. gc>3ob50i3ob C 0 2-n6 65(101306 C 4 5^5 5 p 3 m -
^360005 035(1300^001356015 18 bb35g5bb35 oijibob 2000-093 6563016580, 55-
h 30 i33 A n a c y s tis n idu lan s-dn [51 , 190 ], Byonb '601503(1010 £3(140013560 815365b3
(E le o c h a ris , H y d r illa ) 060)36603(159 000306366 C 0 2-6 (50,501305,b 0 5 3n 5hb3& x ^
P E P -35h 6 m jb o 0 5<bnb b i93503& oai, 656 3oh 65gn b 6560100101 5606005(5056 30 5 6 9 3 6 6
0 6 5 3 3 3 3 (13 9 3 6 8 0 8ymqjo hn630oi'bo>6oq}oibq35Oi3ih6oijbo05‘b 5. C4 8(5365(13360
8(153059 5 3 6 3 9 3 6 5 P oaceae< K > (01035556(15636(1015, 8 5 5., S ijh n b 03h659o, 6 0 8 0 6 -
gn, 033^,30 9 5 b535650n 5i 3hu3036x n 81153500 650*6013560 015365(13), C y p e ra c e a e
(0 6 0 0 6 36 (16 0 ), P o rtu la c e a e (95693(10636(160), A m a ra n th a cea e (3O 505y5636hcn5 ),
Altitude [m]
fob. 2 5 i ] 5nnb dtfQ&itvjxcTiixIn C4 8u;)foh33&nb 9;j9unh3&i (%-n 6npn &nrodibng:>6) g*

(?13C bngnga) rfbfn^n3] 6nb 8^365 bn9i(ji(Vib 'bhQ}bcni6 ;)hcru>g (Tieszen et a l., 1 9 7 9 )
- 70 -
Elution profile
6sb. 2.6 gmoxyib g i PEP iihbmJjnpiVib Shmojngn Atriplex resea (C4), Atriplex triangu
laris (C3) gi 3icn Fj 3nbhng3n (Osmond et al., 1980)
Chenopodiaceae (6 s(3shjicni0 bbo6 hcru>) g i Euphorbiaceae (hdni6 >bo6 hoii) 0135-

b^60 n. 0138(5}, ‘bm^nohoi 535h9n (Atriplex, Kochia, Euphorbia, Flaveria, Cyperus,
Panicum) 9bmomg ohcnox11 bibom&obn 55632,3013636} 59 ^nSb. 65013005, hm9
33mQ3i3fii C3-gb6 C 4-i36 bi3b36noi Sh^hobocn oym, hbg5^6 53639365 55(19595-
3ipn (C3-C4 8350393(10) 0301(19360, (1^803680(5 C4 9 (5365h336nb50i3nb g i 95 b5bo}-
0136300 3nbljmQm5n3hn pi 6nmjn9n3hn 6n9i6oi3nb36o6n 9395)1361101 65303659
ihnb 5i63ncnah3630n (6ib. 2.6).
Flaveria-b 535(190 (m3. Asteraceae, 5b(jh5b36hoi5) 5(p6o9630n5 0156950156m-
6noin 5igib30i F. cronquistii-Qb& (C3) F. ramosissima-b 95 F. floridana-b (C 3-C4)
5530001 F. trin ervab^ b (C4). 0350393b C3-C4 bibombobb 903330136365 55(130133
Panicum milioides 95 M oricandia arvensis [26 ]. ohonbi 95 n9^33 n6gn3ng303&3n
6ib 9 nh65gob ibn 9 nQiynnb ^n3n 5 i 63oai5h36nb bb 35g 5bb 35 3^ 53^3 93od03&5 556-
bb 3533&30n nymb (bn9 n6 gnb qjmoi0369n C-4 bn6 ghm9 n 930966335 9bm0m9 Jbm-
300015 9nqj3h36i3nh36nb5b) [ 38 ]. nb 9300)0365 nu3036m93b bn6ioi0nb 553036001
{Flaveria brownii) [166], 56 3hcnn 55(138111956 0omh30n 9u365hnb 55p5b3pnoi
{Eleocharis vivipara-b b0303Oinb cgnh05 803330136365 C4, Bypnb qjmh05 ^n C3
915365(1336^.
C 0 2-nb <30^5(505 CAM 015365(133690

8353360b bibnoi C 0 2-nb 9m95h5536nb5 95 65b9nh6yo36nb bn6ai3<bnb5ai3nb
9 ioin 550my3636nb g(?3-^583110 9mho53m6ob 3<bh36330b5ym<359 531500363905
j0mhm305bli)36nb 930(5330° 35h3936nb 3533mp36b 555669301 b5^05m 9mi53pm6ob
b505h55m bn3h(53- C A M 915365(13015 95qjm(jmbn6 oi3‘b3630 mh556m36b b6mh39
ib30in 553630365 5^301 - nbn6n 930155336 9090 <bm9 ob 039m9qj36n 3jh390&nb
- 71 -
Gabaofnwem a n b a ^ a w ett a a c o ^ o enamoafta&ob ffân da6;>
fob. 2.7 CAM d(j35ih 3363n CCh-nb

33013 nbjdnb, &>{>33&nb dmdhimbnb, Bj^nb}
gi bsbidj&cnb gifchnsj&nb ^nSrvjhn gtpn^hn
arje^bn (Kluge & Ting, 1978; Griffiths, 1991).
CAM 3u36*h]] 69n CC^-bitKni (J3(vib
N ight Day ou^nbjJohij&ioii B^bib^b nb. Neales ( 1975)
93015330 Jcmh36jn9bb i6 gngn bi0ihi5m biou>3b36nb 9jm63 Djf^gcni 9h33&b

(»t>DiXD6i!>fT'£>n*J bihnbbn": nb. 6ib. 2.10) îh6m6nb 9gi336nbi g i 6y0nbioi3nb.
CAM 9i536ih3369n bn663039n 9ioin qjihoimp 5ibb6n0n 6i5336nb &'bncn C 0 2
93gnb 33h3p363n g i nj qj3h936j) PEP->)ih&mjbn0i'bnb gib3ih36nai jt^ a n h g jb b
qgmbqjm36 rn0 -3nh33i^bI hm930nu 936935 NADH2 - gi 9 01^936300 Sipi^-gjlnghm-
536i‘bnb 9m6i6np3m6ncn iCPG53&i 6 i0 i^ ig . PEP-b ndp33i 50nj)ri0ribn; 9i 0i^nb
6ih 9 mj96 nb 93bi6i9nbiQ J0mhm30iblj369n 9ynhg3&i bibi33&pnb 936^330016^
(6ib. 2.7). cgnjbiynnb 3nh330n 3hmp3j^n - 9i0nb 6gi3i 93gnb 3ii3m09n gi 33113-
gnb 633680 5hm3g3&i. i 9 nb 936935 (?i9 nb 5i 69i 30m6 i 9 n 33(1391"^ 6336nb 9 gi-
3ni6m6i o ii6 p io ii6 9i^ 30m6b (pH 6(jnh036i 5-gi 6 3-003 ). gn0non, gpnb bn-
6icn03b3 9i0i^n 5igiig5n0g3&i bi6n6ii[n0g35m 9n6 ih013036001 - 3ii3m0ngi6 yn-
^Rbn00n g i j0mhm30ib^369n, bigitf nb p3iih6mjbn0nhg36i. 5i9m6cni3nb3qj-
03&30n C 0 2 9oiin6oiJ936i RuBP-b 9n3h g i i(?g636i 6ib9nh6y036i9g3. gpnb fciS-
9i 30m6 i 9 n 0^9 im 6 b 6mh9 i 03hn C3 yn ^n . 9i0i^nb Shm&habx Qi90ib oii6 b g ^ b
35h3gnb 6336nb 0gi3ni6ni6nb 5 i<bhgi (8gi3ni6m6nb gc>3-(T>i93hn hn^9 n); CO^-nb
33h3g9ngi im B^B^hiuni 3l539i, 6i533&n nbb636i, ibi0n C 0 2 93gnb 5ih39mg^6
g i 3i i 30nhg36i PEP-b.
9R3036icrii g(p3-^63(111 ou>66n3g33hm6i (COo-nb 9 oii6 cnj3 i, 9s)i3ni6m6nb
I53i036igm6i, ibnQngi^ni) y330i C A M 9(536^380 3hn6 ^n330ig 6 ih 333x n i; 0 i£,-
hi9 i9ibo ii6i33 36gi x?n6 n96 mb nb gngn 5i6bb3i336i6n, hiy 33n0R633i bn6icr>-
03bi g i bn663039n C 0 2-nb 3 ou>6 cnj3 nb n6ji36bn3m6ibi g i bi65hd0n3Pidi9n g i
- 72 -
s a b a o fi& a m 0 3 ^ 7i?ccf>G^gn
6ab. 2.8 Agave deserti-b bb;)ieibb3i

sbiyibi gi Vn9nb 8(536^330 CO2-
ki'bcru. (J3pi 24 biicnnb SiSdopb^. ,jn3-
n^tin CAM hr^Jn 3006936* 'bhgibhx
ib ii 3 n (Nobel, 1988)
Control
Time of day
6ib. 2.9 CAM hryjSnb 0693^21 Mesembryanthemum crystallinum-dn 6nigi5nb pi3pi83&nb

'bhp:>b<ns6 3(10119 (Winter & Luttge, 1976; Winter, 1985)
- 73 -
66b30 A 66 ^ 0 b 9 n b 96 (?»a66 9 9 6 6 ^ 0 6030 )00 ff>0 &0b P 6 ff)0 n d 966
hi(j 5i63nhm63&xni 9ij36ihnb bibQmbnoi, 3nhn6 ] 6 ncn (6ypnon 3^3633-

Cymgi, ^ g ^ h i ^ jh i , gpnb bi6j)hd0n3m6i, j>i63ncnih36nb qjibi) (6ib. 2.8). 3b
5s5bb3i336i6n agndo^bi ^sSmnbi^mb C02-nb (?i3 nbi gi gpnb <3oii6 cnj9 nb hn^-
bm6hn3n 9 if<3363b03 bnb 93q]ihg3bnoi.
CAM 0^536^301} 6 ^ 3 3 3 0 6 i 6 n03 n (bîôoB^hn p^hmb 0 Jm63 ^ ^ 3 6 0 ,
^5^3360, iom], m3. Bromeliaceae g i m3. Orchidaceae (jig3ihnb3bh6n)) 9gi3Hi6n-
6nb gpQ-piQjhn hn^Qx^bi 83066633^ y330i6inh 3nhmb3b3n; bb3i CAM 3u3-
6ih33&9n, 39cni3h3big m3. Aizoaceae g i m3. Crassulaceae -nb bib3m&3&9n, CAM
5bib ny363636 3bm0mg j)3i03ncn i6 6nigi5nb jfihbn gi0gi33bncn g,i63nhmb3b3-
0n b^h3bnb biSib^bmg (6ib. 2 .9 ). Mesembryanthemum crystallium-dn îhbn
gi00i33&i 06333b Crassulaceae -oii 03vj)i6m0n'b9nbioi3nb gi9ibibnicn3630n qj^h-
336(|)3&nb de novo bn6cn3'bb (nb. 6ib. 6 .4 .).
CAM 3(j36ih336n b9 nhig 53b3g3&i 06 nh 6nigi53b'b3, ^1130133 nb3cn j>ih30m
3nhm63 &9 n, bigitf 33 hnmg30ig nynb 63 i 03 i. (jhiTOnj^nbi g i b3b(jhm5nt)3&nb
b3j)3036lj|36001 gibib03&30 3 3 0 3 6 ^ g i 3gib6m3b9n y3O0 i JjiJoobn, ^ 3 0 1 3 3
Degree of succulence [g HjOdm2]
Bib. 8.10 53ih Aeomum-dn CAM 3>ji 6mcn<b0nb gjmhajbn bjjxOîjm&nb bihnbbot>6 iS39nh9 n
(Pilon-Smiths et al., 1991)
- 74 -
6abaofnM em (33g?a
Asclepiadiaceae g3 Euphorbiaceae<K> mjib^&nb bib^mbicni CAM

8(3363(13366 6031)30136363. Brom eliaceae g i Orchidaceae-n b mj3b36ob b3b3m630i3
50-60% 35(130133 C A M 6(53631133603; 2>CeD06b3, 6(1363660, 6y306ihRb 395,003660
9 mbihpn Crassulaceae- ob 63(181^18395369360, biôhm36nb 638oib33336o, C A M
6(5363(13013 86533b 83O36m0ob6b 383(03363636. 93303 8bj,33b 53(138™ 3ohm6369n
6(5363(133606301306 63(1536003603 63b6oh&3gob 3bo8o03(5oob Shm^bob b3qj3b3h3-
63g 9363609363 ghm9o: ("36001 CC^-ob <530^3(503, gpobooi j)0 <30^01136300 CO2-
ob 533my36363. 38 fcbooi 6(5363(13 oj86nb 63b6 oh63gob 63(1356 ob3, hm6 01330036
0150036b 6900b g303015000b 53630013(13606 hob^b. dgo3ho 533p3ob3b, hmg3b3(5
6353360 g3b3h30O3 gc’3(j g3 £>363(5, b36oij3ob ghmb 63h8 mj86o0o CO 2 2)3033
90130601^8363 j0Pihm303b(!)36nb 603b; 38001 C 0 2-ob 63036bob 6363(1636363 bg3&3
03, 08333 ©hmb, obm5363 6y30o. 63(1013003, CC>2-ob 3b 03b3h30O (502100 ,,033^03“
835(136 ob 3bh36330ymcgb 6(5363(1360 6366oh&30ob 6363(1^363636 03 35(130133
q3mlj)fTibo6oi3b3ho 3^3(13006 03(533b ^3h6 o bbo33ho 3&3(i500b 0383^3636300
8nj83Q36ob3536, (139536 536369330003 8m(5oh2)Xf'h 3 C 0 2 8mj83g36b hm^mh^
303johni636nb b36m0 mm 3ju 33omho.
25 mj3bob 20000-b3 8 30 ° b3b3mb3 oy363&b CAM-b 63b3nh63gob 3bo8o0o-
h3bob30i3ob [82 , 243 ]. 6301 3 mhob 3ho36 303hogmg3O036o, hmsmhtf, 83530ooi3g,
bb33g3bb33 3^003000 P o ly p ad iaceae g3 Isoetes-ob 6ygob b3b3m636o;
3hoi0366o3636o: Orchidaceae, Bromeliaceae, A gavaceae, Liliaceae 630130001,
35(130133 01(10366036013 m53b36ob (C ac tac ea e, C rassu laceae, A izoaceae,
Portulaceae, Eu ph orbiaceae, Piperaceae, A sclep iadaceae, G eran iaceae,
Gesneriaceae, Rubiaceae, A steraceae g3 U533) 63(18018395360360. CA M 6303601-
0ob8o 93od03&3 936mogj3h50mb 6(5363hob 8bm0m0 53(12)33300 mh536m36ooi:
<53(^1010108(5336 Ererea indica-b (m3. A sclepiadaceae) 03010103660 C02-ob 6301-
3ob3&3 b03&3 C3*ob ftbooi, 6obo yomh^bob (p3hdooi, bmh(5036, 636036 6360-
03660 2)0 3b 5(101(5360 C A M ^booi bmh(5O300363 [ 130 ]. 6308036 b3bm69 o 3b
8 (5363h3 (hni03b3(5 ob 63 q3moi0 O0 O3 ) 38 5booi 35 hm336b &3b3 oh630 ob gog (1301-
0360163b, 835(138 89h30 b3bm63 o qjoioim053(5336o0o 3(5363(13 8o83hoi33b CA M
5b3b, hm830O(5 63039 3033^03(103 6y0ob 03bm53nb o>330b3bhobooi. y3303 C A M
8(5363hnb 0363b3bo3oi3630o oi3nb3633 63010 03&30O 8m8oibm3630m63 53(13301639-
6o, h3(5 6301 b333303&3b 3d033b 033(101010 633h(5300636 ob30i 3210100150311
6033660, (101501(10(533 33oq)no3hn 305opb36ymqj30o, 2>C9 DD&n' bh3836o. C3 8(5363-
(13360936 CAM-b3 53g3b303 60363 oi3603oi36m6ooi, C3-CAM 6303363(106 5330001
(63b. 2.10). 036m0oib, 3(16360166 63|j>30b!33(5og)o2)3ho 536bb3333636o, hm8036o(5
03836360301363003 01005010(1010330 6903660 633(1(53936390 CAM 6(5363(13360-
b3cn3ob (Crassula aquatica g3 Littorella uniflora).
2.1.2. <3fWlb36(Jl:l3:>: 6C flin C ^D h fl 6*b2)
bo&3oi03b 3 jomhm303bo36ob 638(5339 8(5363(1330 jbm3o03&9 o ^ 4 -6 3^ 0 30 0 0 6

33(1393606 53hg3, (10180360(5 3h 630(53336 ho63Qmbm6ocgmbq33O-2)3h6mjbo03b3-
mjbo5363b 3b) 0 o6gn63h3m6 b <3 mombo6oi3boi36 633(^036300 3hm(53bo, hm8300O(5
6oomjm60ho30o 63601^306 8(3533630, 6013060^363 0 2 03 01330630309363 CO2,
835h38 8ob536 536bb33336ooi, bo663039o 3b 3(101(5360 699363. 0 2/C0 2 53boi3
- 75 -
G a b g o tfw m anbaartaw ^ aa^ô 603 m006060 b ^ff>anda6a
Gab. 2 .11 RuBP mjbn536 *'bsb 9* RuBP iphbmjbn-

ps'bib cn*6*q)5h0m{xib ^31^035^ 3*656*9116* 9*
0 15 30 Pa 6*b3nhmfg*5j)nb 3*h^n*Q]h 6633*601*6 i* 30nh9 n
C 0 2 partial pressure (Lawlor, 1993)
U 3 p ib b r» 6 ic n Q n b 1 13 6 0 1 5 3 * * 6 3 c g m ^ m b 3 5 c n j3 ;> 3 6 m g 3 b .
c g m ^ m b 3 6cn j3n b 3 nh 330* g b 3 & b ^ h *^ b 6 *h 0 m *g 5 3 6 b h n b jp m 'b m b n c g m b q j^ n , hm -
a^Qb^LS 3 3 3 d Q n * * h i 9 *h^m C 0 2-n b , ih * 0 3 g 0 2-nb ij^ U ^ m h n u nym b . 0 n n 3 h o i3 b b
hi gi656igb, hn630m‘bm6ncgmbc9b^n g*nb0nfi3&* qjmbqjm50 n[33hn6 nb 63*3*9 0^>
grnbcgm50 nim0 *^* 0 . 5 *h30m9 n CC^-nb g i 02-nb 330 u330m6 * <33(1336^ hn630m-
'bm bnqgm bqj*^) i*h & m jb n Q *'b n b 0 3 3 3 3 m 6 n o i *h 3 5 3 0 fih 3 6 b 2> *h b m jb n 0n h 3& n b * (cgm -
^mbn6cn3‘bn) g* g*g*653nb (b36cnj3*) 3hoin3hoi33a2*h036*b. g*656*gnb 3*(?*pn
6*b. 2.12 C4 9u 36*h3369n <gm(jmb36aij3nb 5cn>)mc*03tin 6'bnb bj30*,j3hn 5*3 m b*bx 3&>. CC i * c 3 n-
6 nb (jnica PGA S-ajpibqjm&oni^h*^; PEP qjmbqjm36mc3nh33*jn. an(jmjfTi6 phnnp*6 5*3mymq)ncn
CO2, 6*6bb3*336ncn C3 8(536^336110*5*6, 9*811633 gjnjbnh93&* 93(bmq)n(Vtb 3jh 3p9 n (Hess, 1991 ))
- 76 -
31(1130103(10 6633} b3cb 36yoi6b 0301,1)0^360^31^ C 0 2-nb 8i(T>ipn -
cgoiljoibn6oi3'bbb (61b. 2 .11) . hig5i6 qjoibcgo^pnioipiujnb 6 ih 8 oi;j36i giSmjjngab}-
0ni 3103060b unt^Sn hn630oicboi6nq3mbq3iljnb 3hmg3j(jni‘b3, gm^mb^Bcn^nbib
Oi-nb 8 on 6oij3 i g i C 0 2-nb 5i3oiyoiqji rfbhggbs ogm^mbnB^^hn i j(j,n3m6nb
i3i(n036ibcn}6 3(10119, hibiy 0301,1,016011 6ij,:>gnb 5igng;}6i 5i6i3nhm636b.
qgoibqjoispnioipi^n 5pnioi0iljig gi (goibqjiîg j0oihoi30ib^3&9n nbpnf^bb.
5pnimp^n j0mhm30ib(j36ngi6 5191195009361 33hmjbnboi8368n, giibcoi^bnoi
3nljmjfTi6ghn36nbb30i DShxDC io ^ ih ^ c ^ ^ b S n , (10130360(3 830(31336 50030101^01-
jbogibib, g i l*jhi6bi8n6i'b36b. 33hovjbnboi3363n 50030101^0 03165361
50omjbo0iî9, hm330 O(5 330g35 16 3oi0ni6i9 935hignhg36i C 0 2-ig (oijbipi^nb
biqj3b]hnb 5130001) 16 ijhi6bi3o6nh36ob 83g05ig 6ihginj3636i 500(30619 (61b.
2 . 12). 50fiun6n 33hoijbnbm836ngi6 3n(joijoi6ghn363n (16 jpoihoi3Qib&)60 n)
5igignb, bign5 3 nbn oihn 601033391 3(1019361 g i 6ih3mnJ3636i 3hoin 3 rT'03 i X i
b3ho6o. i 3 ib cni6 bg33b C 0 2-nb 5i3oiyoiqji. bohn6n Ri3hoi3nb i8n6oi3g:>3iaii
U30i 3 o i6 g3'bi3 n6 nh3£)nb 9 o3 g 35 6 ihginj3636 i 6pn(53hivi)ig (SnghoijbnSnlî^nb
biC53b3hob 6i30noi). 500(33(11,331 03od03&i 5160131901b qjmbqjoihnpnh^&i gi
gi36h36g3b 3i03n6nb unj>0b i6 83nd036i 5180^363630 nj63b big 3 ^ bb3i
bi^nhoi3&nbioi3nb.
6 3 6 3 6 !^ 3nhm63&3n (3i3h8n 0 2-ob SihuniQ^hn 66331 21 kPa, pC0 2-obi - 35
Pa, o6^36bo3hn 516101361, ,33833(11,33(11 20°- 30 °C ) C3 8(336^3360 ^ 6 10 10 3 ^ 3 ih
5160^010336 C 0 2-b [2 2 3 ]. i 3 6(336ih3368n qjoi,3oib36aij3i 601(1(50399361 3bm0mg
8330180336 33h3g363n g i 6 ih 3 mj36 o0 o C 0 2 (13030^0(19361 33<bmqjn0nb 33(1393680
(6ib. 2.12). 3b "biq3i65n“ 3<bh36330ymqjb qjm,joib36avj3nb 8393519 6 ih3 mj9 6 npn
C 0 2-nb g iiih 5i3ib g i 3b 5i6i3nhoi636b i 3 815361(13011 3 ip i0 3hmg3;jljn3poi6ib.
Sihoiipm, 50030100,33(1 33^160100^390 031(15361 C 0 2-nb 5 :^ 33 3 x 0 hioig36oi-
6i 20-50 %), 3i5h i3 (101501(1(5 Ri6b, 0301,30^3601^31 6ih3mig536b ,,1313003630 601-
hoi^36ib“ . ob ihnb i8n6oi- g i 1560801316 8913360116 gi|)i39 nh3&30n 336tboi‘b3hn
33^)i6oi0n'b3nb 5163^013606 63(1,3000. 5ihgi i 3 nbi, îh6n 5i6ioi36nbib 3ib (b3 51-
9190b 393^(10163606 613190b 6i6n0n, 3b 3 a 5i6bi330ih 36001 30 PS II, gi'bni-
636nbi5i6 ni5i3b qjoi(jmbn6ai3'bnb i 3ihi^b.
2.1.3. 363h6nnb d ^ a F iy m ^ WiW&PlODh 3hmi3DbD6gn
(101501(1(5 13601600m, i6i6oi03hn 3hm(53b36n 516130(10163636 6n3oin3h36ioii bn6-

013'bb 81336111380, i 3 6n3ain3h36icrii j)i vji6oi0n'b3nb 16 3 gi8pnbib 30 5i3moyoiq)i
363h5ni, (101630013 3'bh36330yoiqjb 33(13980 3n8gn6ih3 83,3160103(1 3(1011536366. i 3
h3ij(3n36ob b36b,3hi^)b 61(18011953636 6ib8nh6yp36n 16 (jbn336n. 16 6n3-
oin3h36ioii 3i‘b3h5oi630o giSpnbib 50nôi0n(bnb 16 bi86ib8nh6igni6 (jjh36bnb)
Uoi03o, I5h30i33 (3bn3oi3i6 89131011 [3^gi9i653nbib boih(3n30g3&i 6yi06ignb an6-
9iaii6oi6noin, biq33b3h36hn30 61801(300361 g i 363h5nnb 5i6oii3nb3qj036i. 6yi0-
619b 03(1013636 3nhngn6- 16 q3pi3n663i03oili)ng36n. 363h5nnb D33iJ3bn 6i6npn
3nn(Ti36i 6yi06ignb 5 i g i ki)i6nbib 8nb bi6oi0oioi ij(333ôih'b3. 3n,ioijoi6ghn30n
b36oij3nbib i 3 ij(333li)mhb i^,3 mbqj3h30n 9i656ign 6ih3oiig536b: 303j^hoi636nb
bi^hi6b3oihôi 3i^3noi 5ig n g 5O 0 36 x n 6yi06ign 8nli)0ij0i6ghni8n 3Qhoig36i
gi656igb. 18 3hoii33b3n 6ih3oinj3636i ATP (b36oij3nain 31^33(10 0 30 ^030 ^0361).
363h5nnoi 3gngihn b b 3i qjmbqji^bn 6ih3oinj8636i 50nj)oi0o‘bnb 3hoi(33b9n
- 77 -
B a b a o A M m 8nb86(f>oto aa^ô Goamoorssoiou pÂanda66
Ijn^bmpSn (b^b^hi^x0 3^3Dhn qjmbq}mhnp36b). 8oipni6ig 3hm(j3bnb j>i3ri-

bi3ipn 3ose2»36b 36 3mpn ATP, 2 8mpn GTPgi AG = - 2B7MWb vjmp 36îp3nib
(= -686 i*p>c) 3hon 0mp &p3ii»rt)ib3.
b^h^bx* gib ni 636 x & SmbQhQbx Jbm3np3&8 n £)h8 n6 ip 3hn X 3i!)hfn^h>6 b-
3mh^n 83ndp3&i fci6bb3i 336x n- HCN-ncnb giCO-cnn nBSnbnhjbbgn
6nb Shmu^bngiB 5idmcnnd3nb &bnoi 6ihn8ihoimb. *3 uni6ngh3bnb^36£)pf> ^36-
cnJ3nb ghmb gi653i-i(Ti0536ncnn QSgh^nnb gngn 6i6npn 5ihginj3636i bnotbmg.
OJb^hjSiCDh 3nhm&363n igfcnpn ij3b i 3 Qî^mpghn &bnb bi68m»)PD iiyvjrfbi-
(jnib (3i&., bmjjnghoin Araceaeb ^phmb y3i3npfn6nbib), hibiU ou>6 bg33b ^38-
^ghi^^hnb bi5hd6m6n 0m3i^36i. 3b y i 863h36nb 8mbng3ib J>i6i3nhm&36b. yn;>6-
ngh3bnb^,36^)Xn b36<nj3i y^330cn3f*b Rihoixto 3m36nqj36nb 3hmi53b3n 3ymqgn
6iymq336nb jjpnSij^jhnx b36orj3i8n, qj3b33&8n j>n nb 83ndp3&i 6^^63330 nJ63b
NO3 - i6 NH4 - nm63&nb bnîhbnoi. i8 b36cnj3ib R33XD^^n3 o’iB bg33b
gibftbignb 8ou>6o)j8nbi gi C 0 2-nb fciSmymqjnb 96n8363pm3i6n &iijljn33&i [127].
93pnpnb ghmb 6yip6igb n3hoi3&b bb3igibb3i mhj)i63pn 6iyhcnn, i6i3hm63-
pn b36oij3nbib j>n ihirnhj^x^ nm636n, 8*5., 6n^hi^3&n g i b3pop^36n. hig5i5
03bidpm 5i6bb3i33&i6n 6yip6i0bi g i gb6j)di0b 8mhnb 5^0036001 330ni, 3ngh3
6yip6igbi g i bb3i gi3gi653x ibDîbD^5 Smhnb, 3b îî&mp3hn 3hmu3b3bn
ndp33i 5i(5np36nc7> 6 ^ 3 6 363hj,nib, 3ngh3 i3hm&3pn b36cnJ3i, hmdpnb 3<33j(!oh-
m6i 30-40%-b 33ig 636b.
jjpn^mpmbnbi gi pn3m63 gi 3i un^pnb ftihgi ihb3&m&b 5P3imbnb gi 3 pnb
03bi83 j>bi, hm33pbi(j ^qjhm b3 nhig 3b3g3&noi 3 ipipn gnq£)h36(3nii5nnb 3jm63
0U36ih3358n. 3b fcbi ^ x ^ b S m b b &p3imbmq)mbq}i^nb 3nhgi3nh gig i653ib
NAD+-nb 3ig>n3h NADP+-nb ip gj^n cn ; g3iph&mjbnpnh3&nb 8 3 9 3 ^ 9 6 ih3 mnj0 -
63&i hnbxrnbm-B-qjmbqji^n, hm33pnu 638935 gi6&3ncnn ^^m bm jgm bqji^hn
yn^pnb g,i3pncn nb33 hn63pmbm-6-cgmbq)iîg 5ihginjd63&i. 363h5nnb 3hmg3j(3n-
nb oi3ipbibhnbnoi 3b g>bi 6ij)p3606n3363pfn3i6ni 9(j36ihnb 03(ii&mpnli3&r>cn
3bh3633pymqjibcni6 930ih36noi. gi6&3ncnn 336^mbmq)mbq}ij>3hn ynj>pn bmhunxg-
3&i ijn^mbmpbi g i jpmhm3pib^,353n. nb 3 ^nghmg ihnb 0 ij>i38 nh36 x n XPebD^ncn
336jjmbmq)mb<3ij)3h (îp3n6nb) unj>pcni6, hm33poii6i(j 3 ib &33hn bi3hoim h3ij-
I3ni ij3b. mhn33 yn^pb oii3nbn 6npn 03ij3b NADP+-nb h3b3h3nb 33 J36 i 8 n.
2 .2 . b 3 b O \5 15 3 W a i5 D 6 ^ h O D 6 9 fl
2 .2 . 1. 6^banhnh^66nbi e:> 3^666ônb 1530^

6ib8nh6ignb 3ngi35h x x n l53Pi 6n3cnn3h36icni i^ 3 mbqj3h x I530ib o ii 6
gii)i38nh36x n:> 6*<boni U3pnb bi83ip36ncn. 8^36^33680 3b 3pn6g36i C 0 2-nbi g i
0 2-nb ii3pncn 3^36ihnb 8ngi gi j>ih3 ih3<>ii 3mhnb. îbcni q)m^mbn&or>3<b3hn
U3pnbib 8(536^3 noi3nb3&b C 0 2-b g i j^mymqjb 0 2-b, fciboii b36cnJ3ncnn
U3pnbib ]>n gni83j)hip3hig bi6n6iip>393j)m 3hmu3b36n bg3&i: 81536^3 n^nb^&b
0 2-b, îdmymqjb C0 2-b. cnncnni33pn i 3 mh 3hmu3bcni5i6n, 013 ghmnb 6 ^ ^ 3 3 3 0
8m6ii>330>8n 3hcnn „3m66nb“ 83mh3b, 83ndp3&i {>i6nbib(p3hr\b 8r«3mj,mbn6cn3<b 3
- 78 -
(VttOto 03^ 91306^330
9l536ihob qjmcn0nb ^g iin h ft^ . hmgQbxs qjmombn6oi3'bnb Shm^bSn 3cnin6oij036i

^qjhm CO2, 3ogh3 n9 b3ghm30ig {jiQrrmypiqjb b36cnj3noi 3hmi53b3&3n, 3b
îh&o 3oii6oij0 i (j6m6r>0ni hmj,mh(j C 0 2-nb bnoxn 9 oii6 cnj0 i ^63 63om-q}m-
Ombn6oi3<bo (Phn). n^n 5i6b b 3i3g36i C 0 2-nb oÔ^D^n 9 ou>6 cnj0 nbij)i6 , hm030-
bbU ^300ihn(j ^63 6 h30 fn-q3m(!)i,nbn6 oi3'bb ^63 5hmb-q}mombn6o>3<bb 360193636
(Phg). »p»^6rncjh 3hmu3b360n C 0 2-nb 5 ^ 6 ^ 3 3 ^ 3 0 ° Bih0 mj86 nb j,i9 m jjhmb-qjm-
lynbn6ot3(bn 33odo3&i 9 bm0 mg 0nibom3&oai 5 ^9 mncn3 iQmb. g n (i)nbn5 cn3'b3h ibo-
0n0i(inibou>6 9 ^ 3 8 ^ 3 6 3 0 0 bij)no*b3&ob 33b6 i 30 nbib ^3330361103 bij)9 ihnbni
63om-qjmombo6o)3'bnb 5i6bit>(p3hi, hitf fci'boii (53Qnb j>s‘bm03noi ihnb 33 bx>03 -
&30 n.
ibn0nQi(3nnbMn3nb ihib 30 b iy h x 3nhm&363n qgfnombn6oi3<b3hn iJon3m&i 33 nd-
03&i 3unh3 oymb gi ih 5igiiîh6mb qjm(i)r,ib35cnj3nb n6o36bn3mbib. i 3 ghmb
3jb33hn036o3n ih i(pn6 n963&i C 0 2-nb ihy i 03&i g i ihy 3 i 03&i. ib3cn boo3i-
ynib, hmg3bi(5 3cni6oiJ030n gi 5i0mymqjn0n C 0 2-nb himg36m6i 3hcn0i63OMib
Ompni, 36mg36i C0 2-&:>bot> U30nb biyn0336bi(}nm 63h,yi0O.
2.2.i.i en<3Jbnnb 3hni53bD6n
C 0 2-nbi g i 0 2-nb ohi6b3mhob ip63hb qjnyib gnqo'bnnb j,i6m6n:
dm / dt = -D »A • (dC/dx) (2.5)
gnqj3<bnnb bnfijih3 (himg36m6hn3ig 6ih0mg536npn dm-noi ghmnb dt 33i 03g 3n)

gi0mi>ng3630ni gnoj^'bnnb © 3 9 6 0 3 ^ 3 (D) g i rfbhg36i yn6u360hiunxn &hign36-
onbi (dC/dx) g i 5i'boii i33pnb i6 3 gnqo'bnnb qjihcnm&ob (A) &igng3&iboii6
3hoiip.
goq)3<bnnb 03g0n3i g i 0ming3& x f 'i hm&mhu 0mgnqg35gnh3 6n3cnn3h3&ioii
oi3nb3636‘b3, ^ 3 3 3 ^3^33, hm03p3ni5 bg3&i gnqj3<bni; 3i3h3n C 0 2-nbi g i 0 2-nb
gnq^'bni giib0m3&na> 104- j 3h 3qjhm bShiqjig bg3&i, 3ngh3 6yig3 n. 01336ih33n
j^'boii i530nb 5i5bi'b[p3hnbiOi3nb 03nd036i 5i3m3ny36mon qjn^nb *>i6m6ob
5i 0ihon33& x n <3 nh3i [7 0 ]:
J=A C/ S r (2.6)
jjibnb 5i0^m0n 6 iiig n J (gnq33<bnob 6 i$ig o ) 9 i 0X fT|^ ) 6 ^ 3 Si^hbi g i

33h3g3n h3ij(3nnb 5i6bmh(5n303&nb 3&i6b 9 mhnb ^6(53601^150300 ^hign3600b
(A C ) j>igng36iborii6 3hoiig g i i 03& x m^^ 0oqj3<bnnbig0o 6n6i(pm6i<b3 (S r)
9mj03gn ©01300 hnj,o qjijomh3dnb 2»i3036001.
2.2.1.2 iP16l536i!>h:>l5fDCn 2>hên36(M1
gi3303i<n, hm0 <3 mo|T,^n6 cn3<':,n^ 33935^9 Jcmhm5pibo363n C 0 2 i0mn63hi g i

0nomjm6ghn369n b36cnj3nb 3hm(S3b3n 0 2-nb i,m6o36ohijjni 6 x i d 03 9 i 3(ii- 3 i 3 n6
- 79 -
GibaOfni^ÔU 9Db06(f>Oto 3 3 6 6 ^ 0 6030)03fo350b P6A 8D d8G 6
6ib. S.13 C3m^nibn6<n]'bnb n6l)36bn3m2mb

CP CiM^335 Cijj,
giSmyigjfrjpo&i ^jlvjgSnhnbj&Sn CO->-nb
Leaf internal COz concentration im5(}o5lihbiyib'b3 (Lange et al., 1987)
s9 mho 55"bob 31*161536^115150300 5(159036^,360 5i6nbi‘bc'3h]6i 3(5365hob f^hâmd-

1333C ih 30 n 05010 3m6(536j(h5i5onai.
b0303Oinb 9 [5365(13015 0o6ob'b395 <*1115561*1360 9565659001 35I1559 5(1056 0m-
9^^3536306(1. ypmh^nb j3hjob, 03hj6nb 35h36jn0 ob, 6311000 li>ml336nb 63C63 Cnb 5
95 65ymqjob 363563, 03^1*^060131331159 5^03110 305b(jO936 n 33h300mhob369n 55-
3mymqj36 9565659b. yr*i33po 0bb3O0O r*ih556mb50i3nb q]mli)mbn6oi3‘bnb 3b 3033-
j^n, 93 bid 0 m5 3q}hm 06003630^135605, 309(13 65b3 nh65gnb 653036P165. hm^mh^
656b, 95656590 303(1m 03^59 530050 q)3b336b5 95 0060^3385 P3hm36b. 0136(55
nb3cnn 0(5365(13360, hm5mho|j55 01301110 930qj5h5 [39 ] 95 03hy56o [214 ], 9565659b
0013636 oi3 h0 m9 ocg3<bonb 3hm(53bnb 03 mb36ncn, hmg3b5(5 95656590 0mdh5m6b
33h3g0mhob30 bob(i3053n <gmcn036b5 95 C)3b336b (56 <33b3h36b) 3 mhob 5 1 ^ 3 6 3 -
00 oi3h03Qo 5(159036,^ 800511013036001. 6 mh0503h 3ohm6360 o 0oljmjm6ghn3Qn
b36oij3nb5an3nb 95656590b 593^35(33(10 3016(536(^15(505 0359536b 1-3 % . 3qjhm 95-
6500 3(^6(536^5(50360 50(snh36b b36ortj3ob n6^36bo3m65b 95 5cg3hb3&b 0931)33-
6ob IbhQ5b. (nb. 01530 6 .2 .3 .).
636361103 3nhm6363n C 0 2-ob 30161536(3(15(5iixn 51159036^0 55h3 5h3b5 95 qjn-
oim0b 3 mhob d50lb 3 3(5oh35. C 0 2-ob b53fn3336b5|5om 3m6(536l3h5(5n5. hm0 Qnb ghm-
b5(5 55^015 (5305 ,,^365“ 958(^1309363005 3nli330 hn50o 35h6rvjbo0oh3&nb b3hb<b3
(C3, C4). 50ob 550m, COo-ob 51159036140 55(50036001 05(j>50n5 C4, 3ogh3 C3 9 l3D-
65(133630. C 0 2-nb b53m0336b5(5om 3Pi6(536(!(h5i500b 91*163 C3 0(53651133690 95801-
309363005 556501360b 06^36^3(^65133 (05p50O 060336301*16360 95659 5565013-
65^3), lj3033h5(j3h5lb3 (lj3033h5l33hob 05(43&5boi56 3110159 0^9365), 33360b,
6y0ocn 3<bh36330ymq3nb 3ohm63&<b3 ((^13^00300956 5595^5 06333b 5(p6o963Qfi
05fi3363600b "bh95b) 95 03(356m0o'b0ob 9p3-(n503h 95 b3<bm63h (535036590165^13
[ 10].
C 0 2-ocn 0(j365hob 0m05h5536ob 55^ 9 5 505(p036b 5bo0O05(5O5b C 0 2-non
q3ml|,(Tibo6cn3'bnb 553311360b gm63093 (65b. 2. 13). C 0 2-nb 3qJ3i!>nb 550^1065^3300
011390b 95bhncri65 A/Cj 550mb5v453b C 0 2-ob qgmli)rribo6ori3'b3h 03oi3ob36:»b 95
65h0 m59536b 0535h6mjbnpnh3659n og3h036lj 36nb 5j(yi3i*i6 nb b5< bm0b. 50nb 556m
C 0 2-oo> 0(j365hnb 9ro05h5536ob 559093651x1156 3110159, cgm^mbo6oi3<bob bnRj5h3
3qjhm b6h5q}59 05vJXfT'6b C4. 309113 C3 0(53651133630, hm030ou> 0moibm36nQ365
0 0 ^ 3 305ymq)O09365 0bmpm9 050o6 , hn93b5(j 3 obn 31*16(536(3(15(505 005(^63 3b
1000 jiM *1. qjmli)mbo6or>3‘bob bnRj5h3b C 0 2-ob 5533h36ob 3ohm6363o 3m^36 (jn3 h
gm^mbn6oi3<b3h 365hb (Php„t) 361*193636.
- 80 -
03^ 9(3066<n330
3i3h9o C0 2-nb 030i533C«n6nb b3om363hig fcibhgobib C3 61336^13360 2-3*33h

030 COz-b 9ou>6otJi336, 309(13 63636*103 3nhm6369o. 636^(^3360 'bm&33h i 0ib
013363636 0mbi3Cfib6m6ob îbhgob 0n'b6ncn. bMn63h;)63o CC^-nb ^061536(^^(30-
ob s0spQ36no> (CC>2-not 5i6myo3h3&i) 3m0og3hob, fcm&hnbi gi 0901010^3^60
6mb(i)6330ob 9 mb:>3b0n 03nd036i &imh0iag3b.
2.2. 1.3 6^b0nhnh9^66nb a w e:> 6n6:>cmo6n anufi qjnoincgn

an dh^n bn w b
j0mhrn3Qibli36nbii36 0mdhbm6nb CO j-b 6n6i(pm6iCTu> 0oi30n b3hnnb

5^9>Qib3s 36936:). cgmoimQ^ g i qjmoim^n CC^-ob bi(i)hi6b3mh,jm fcbi g i
606^^16360, hfn03QOii(3 ob 63936:1 i 0 6^6336360^ 6ib. 2 .14-‘b3.
0(536ihnb b 3g i 3ohoii6 ibpmb jpbnb 0m03j>30io ii (330i 9363C3^DCn;>- l »
bi'bc'3hm brnQfib 3b 6n6i(j'm6i (ra) îBbiôih 361-101 0>9 n6, hmg3bi(j
qjnoipnb lb3gi3nh,b3 nj0636i 3i3hnb 30mdh:>m 0h3, h;>u, hfnj,mh^ Ri6b, 3ojhm 0m-
bb0mg63cni, 013 0(536ili3b gngn, g i( " ih x n0363b3npn qjmoi036o ijsb . <b3gi-
3nh3(yi 6n6ii(n0g35m6i 0ynhg36i 3i3hnb 0mdhim6nbib g i b h x ^ ig 306n03630m
bp36i, hmg3bi(5 jihnb bnRjih3 3-5 0 b"1 i£'633b (6ib. 2 .15).
cgmoim09n fcibnb 33tT'G>33nbiai3nb 33nh3303bn 06n03630m6i 6ij,336b ij30i.
6>b. 2.14 cgmonmpdn C O j-nb j)m 6(j36l5hiunnb

jhsen^B^n g i giîh^bnb h3'bnb(j;)6 £ x y T>di
q)m^mbn6cn]‘bnb ghmb. UE 'bo pi ]3ng3h-
3nbn; PP S^bhnbjbhn 5ih;)6jn d i; SP
(nh]6Qnb36hn iih^&Jndi; LEj33gi jSng^h-
3nbn; N P C cjryjrnbnBooVjhiQ ih iiiy ijh n
33h]go6n: B L b ib i‘bc*3hm 3h ].
qjm^mbnBarrj'bnb ghmb C 0 2-nb im 6tj;)6ljhs(jn-
nb ^hipnÊ^yi S yih g jb i 6ih 39m0(53x 3i;)h b i
(C A ) g i ]3h]p9nQ i Si^hb (Cj) 3mhnb. $ih -
6mjbn(vih3&nb ig^ngib 6n6n3iQ3hn yn6(536-
(jh i^ n ii (C c x )- 3sh]g3mhrtb bn3h(}]3n CO2
9;jpnb ih i Sbmpfng jB jic m g i^ 3m bq£)hripi6,
ih id x cnjncn ^jh^gD ^ngiîli. b ig iy 050
5hm39]6i Qn^jmBghrrjpn b]6or)3nbi (CRM)
g i gm^mb36oij3nb 9]pD5i 6 (gi63hnp36noi
nb. Sestak et al., 1971; C hartier & Bethenod,
1977; Cowan et al., 1992)
- 81 -
66b30 (f»&«^0b 8 Db 96 fo066 ^ 83 (*)6 C?0 G030)OD(n060b P6ft8nd8Gi
hmQQbiy 6*60 0ai0 Oi6ig ^ n ii, bij^hn gnqj^'bnnb Bnbipmbi (rs) 8 n6 n8i 03hni g i
gi8m yi93b30ni bi53cni 'bmQi'bo g i 3icnn &i60ij)3bob bnbSnh^'b] (nb. ubh. 2 2 ) .
013 bi&D g ib ^ h x n i, 6n6iic>0g3£)mbi 6i63bi'bc’3h ]cig gngni. b03O3<nnb 0153-
6ih]369n 6ib 9 ohmhsji6j)nb i30>niX D ^n 9oii6aij8i 306n93630fTu>- cnâî bnb63033n
33h ^ 9 m hnbx bn3hu33o pi&hm33b30o b3&cnj3ncnn C 0 2 i 3O>ni30nbi Qi 33hnp3h-
03Qn jbm3O036nb g,:>3pncn goqj36gnh36b îhâmSn 3016(536^1*150300 6hign36vj)nb
0n3ihoixo&nai. C02-nb 3n3ihoi bij^ho gnq^bnnb 3n6n3i 03hn BnBipribi gib ip n i
3b n i6 n libnî^nb bi0iboi363bbi g i 33(1330 330^3(13630, nb 93gih3&ncn 3i(pi0ni
fihgnpnb p i b^03hmgn03h 3 ih ig 0 0 3 i6 3 0i536ih3Oii qjmcn0369n, 6n6oi3i6oii 6n&-
3363n. C 3 3i536ih3369n, 013 0iC7in b i j ^ b n 0oi0ni6ig p n ii, big^hn gna^'bnnb
6n6ipm6i yoi330oi3nb 6i303boi qjmoi0ob 8056001 C 0 2-nb j,igi3m dhi336nb bi3h-
cnm 6n6ipm6:><b3, C 4 0 ^ 36 ^ 336 8 0 30 3ngi 6n6i i p 3g32>m6i n 3g 36 ig S^nhji, hm9
6i53&nb b3h30nb 2>ibb6n0oibob bihnbbn C02-nb &igiig5o03bob 0 i 0 n3nj,nh3b3 0 n
qjij^mhn bg3&i (6ib. 2.16).
CC>2-nb 5igi9mdhi336nb 3 ogi 6o6i(j>oibi 3. 6. 6ihR36n 6n6i(pmbi (rf) 339 -
536^ hi3g36n03 im03m636^nbij,i6: 1) jp'bnb 33(p633nbig3n 33h3gnb cnbn3hn aj>-
'bnb 6 n6 ipm 5 i, 2 ) j0oihoi30ibli)3b8n C 0 2-nb ibn3n0i{jnnoi j>i63 nhmb3b xn 3- 6.
iih6mjbnpnh36nb 6n6i(pri6i (rcx) g i 3 ) <301^0^06013^3110 5ihgij963&nb nb 3 i-
0n0 n^nh3&30 n 0mj0393bi, hiy 5 i 0 m633X ni 3nh330ig h3iji303b90 363h2,onbi g i
gi653i-i(?g&36noin 3J3n3i035^36nb ihibii0ihnbn 3hm03j(jnnoi - 3.6. 15 ^ 6 36 1^
6n6ipfn6i (rexc).
qgfnvjmbn6cn3'bnb 3hoii53b3n C 0 2-nb y330i“b3 3i(j>i0n 3ih(5Oi03 hn 6633i x T n'
hnub36i qjmonpnb 3cni3ih &ih3 9h30n. jjibnb 5>igi3oidhi33bnbig0n 03g0n3n 6n6i-
(j>m6nb 0 3g 35ig ajmowncJJn C 0 2-nb j)fn6u35j>hitjni (Cj) 8150(1036;, 5ih 3301815330
3i3h9n 0 nb 933 b330 m6 ibcni6 93gih3booi. i 3 nb 5i0m C02-nb 3ih(3ni03hn 6 6 3 3 i
35h390mhnb360n j)b63nhm63&30ni 3mb3ih363pn C02-noi g i 0nb bi6ii53pmp
5ih3gi6 930mb30n C 0 2-ncn. 5 ^ 3 g i 9 ngi 3nhoib3bnb 9 3(530 ^ 3 6^53360 b8nhig>
h3ij,nh3636 n0j,3ihig, hm0 3i3h9o CC^-ob 8011330300 6633nbib 0 nbo 3ih|yii03 hn
fob. B.15 ishnb {>^3036001 6ib3nhmhgi65nb ynSmSîiyinb 9^3300136^ qjmoxyib ^ 3pil3nhcni6

ib(vn9g36ih3 l>3hnb 3h33n (Grace, 1977; Roden & Pearcy, 1993)
- 82 -
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nb 3hm3mh(3n30i 9 . b }6 } 8 ^ ^ 3 3 6 0 0 1 3'bh3 6 3 3 0 yoiq3ob o i^ n S n 'b ^ n o b ^ 36 9 36130 }
9 36 ihfi36 36 x n:>< C;/Ca 8303^ 9 3 6 ^ 8 3980 30 }, 6 3 33 0 3 6 (10 3 ob 0,7 3^6 0 0 139 36 } [3 0 ].
01301^1030^9 . j0oihoi30 }b vi)36 9 n C 0 2 -ob 3ih(3o i0 3 ho 6 6 3 3 } (pc) 6 30 ^ 8 9 3 36 9 }
8 (30119360193^ 3}j,h i9 qjyjô}, hm8 C 3 8 i536ih 3 3 6 9 o 6 3 6 3 6 hn3 3 nhoi6 3 6 9 o, 06^ 36 -
bo3 h j>} 6 }cn36}‘b3 (3, 3 3 h3g8 oihob3 6 3 o C 0 2-ob SihunbQ^hn 6 6 3 3 } (pc) ih 8130(19 3 -
6 } j0 mhm3pib^36 9 o CC^-ob 3}hi3O}03hn 6 6 330 b 9 3 6 (30(13606 3hm3mhi3n30}g ; P</Pj
9 3 qj}h9 3 6 ob b iS ^ ^ m 8 6 0 8 36 30 0 16 } 3^60013936^ 0 ,7-b; Pq/Pj 3 9 hob 9 ^ ^ 0 0 13 6 0 0 1
0 ,5-b [2 9 ],
Photon flux density [nm ol-nr2-s'1]
61b. 2.16 C3 gi C4 80361(1301} gmoicaiiSn C02-nb ^sîhj&nb h3'bnbli 35>j x fT'î (ra> rs)' 6ihR^6n
h3<bnbli36(j x m* i (rr) Q} 6ovjm-ajm,jmbn6cir)<bn (P ^ ; SjBj^nhn bi'bn) bn6icnonb 3i,53i>iboii6
ii33nh3n. C3 0u36ih33n q]fn,jmbn6or)‘bnb bn6ioi(yioi îjjhQ&nbib rr 0jbi9S633ig £>hdmi>b rs,
3i 3n6 hmip C4 c^ B ih ^ n , Snhnjnoi, nj,n hi3g;)6}g33 6^^0360} (Ludlow & Wilson, 1971)
2 .2 .1.4 6 ^ 01^ l53Cflb h363 C^l3fl^ 6 i 6 D36 flb 3 (lDh
6}j,ob b3h30ob {,}bb6o0fn6nb 3bhoh36ocn 8 (336 ^ 36 333d0O} 03010101080 C 0 2-

nb 83b30ob h36X nh3^ - R}8i3iiO 33(139360 839803 8mdh}m6}9o }ho}6 - }8nb
9 39 35}g 6}530u> b3h30o b}6 306(1013936}, b}6, Snhojnoi, qjihoioig nbb63&}.
636ob8o3h 801836(590 qjmcn0ob R}8i3iio 35h3936n ihibghoib }ho}6 p o i 3hai6}oho
b}hobbooi. 3b J,}6b}j)30ih36n0i 3006936} b,4 ( ^ 3 0 b0(j)3:>(303680 (6}b. 2.17).
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- 83 -
66 b30 f n d ^ 0b 8Db86rnOM ^ 33 ^ 6 ^ 0 6 n3m00fo360 b g ^ 3 n d 866
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width
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munis-b qjmcnmpSn Sîppnbib (Smith et al.,
1989). &>630&nb i 6 i^mdru>, {pBpijjbnb
cniSnbjb^hjbsSn pi q^36j^nm6nh^)6i nb.
Terashima (1992), Pospishilova & Santrucek
( 1994)
2)0 16 ^ (13 0 30 3 0 ( 1 0 1 6 3 6 0 1 3 6 8 3 5 3 3 6 0 b 8 3 0 3 6 3 0 g a b b 3 i p i b b 3 i m 6 i , 3 0 c n 3> 0 b i'b h n b n c n ,

o in o im 3X 0 6 9 0 30 9 0 0 (5 830866333 ( u b h .2 . 2 ) . 3 9 5 n 3 3 6 3 0 m 3 i 6 3 b 36 :> o m 9 n 3 h 3 iR -
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0003 6 ih 3 m b 0 5 3 6 b . 6 3 5 3 ( 10 g n q 3 3 <b o n b 6 o 6 33(n 0 g 35m & 3 0 ^ (10 3 6 3 6350b b 3 h x nb
53009360b 3 ^ 3 0 16 3 6 (5 0 3 9 3 113 0 , 9nbn 8 3 6 113 6 3 6 3 0 0 0 i b 3b n 3Oi3 6 x n 1/rs - 6350b
53003(13690163 (gg) y i 6350b 5bbb6oQm6ob 3oh933oh3hoi3oihi5n3903 . 6350b 9ijbn-
© 39 3(10 5 3 ^ 3 6 0 9 0 16 3 , h iy 0 30 m ])0 9 3 6 3 0 0 3 0ob cgm h 03 b 3 9 3 R 302)3 0 ° 3 3 (13 9 3 6 1^
i3 9 C 3 & o b o i 3 n b 3 6 3 5 ‘b 3 , 536 b 3<b (n 3 h 33 b 03 b 9 n jp b o b 0 3 b 3p n b b o K Jih o b bQ 03
<b(P 33 h b . ( 1 0 ^ 0 1 6 ( 1 0 3 3 9 3b 5 3 8 0 ^ 3 0 3 6 3 hm j,m hi3 6 3 5 0 b 8 3 ^ 0 8 3 9 3 ( 1 0 5 3 8 0 3 ( 13 6 9 0 1-
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(j»03 33 h j n b 8 ^ 0 16 3 0 3 0 10 10 19 8 15 3 3 6 0 9 3 o h m 3 oi) 30 n o y ° b b 3 3 6 o b < 3 0 1 0 19 3 6 0 . 3 b 03 R-
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6 3 (13 3 6 0 0 .
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b3h303&ob 3h3b 360193636. 3b b3h30ob bodi)3ho3ob3 (qjoiongob <b3033nhob lmm2-
^ 3 63530*3 (13010360163) 03 63(13906 03jbo03p3ho bnj,i6nb 630(13396 63(1801305366
(6350b 033(1010). 0(5363(13013 30(133936016380 b3h30Oi3 qjihcnn qjoio^nb <b3033nhnb
0 ,5-1,2%-n3, 0138(33 ohoi3o2> x ° 8(5363(133680 036 83od03&3 3% -b 0r»3(?6noib.
3b 83^33636390 536b3j>30ih36oot 9363903 b32)XD60Dh 0^ b2>O3hoi0 oihq3X qjmov
03680. b3])XD6036ob b33bo0n03(5ooi 01(15360136^ )3-
I1033 9363903. boi0m 363803010190360 93 831139863363 3333 6 3^ 0 1336 0 8(5363(133-
60b 03010193606 635336b 8boigoig 019633 53bb6ob 363hn 533660301.
6353013 9oidh3m6nb qyVbnoioniftn^hn 83^360^380

6353360b 53bb63-g3b3h33 b0ih(50399363 R38*130 33(1393663 9 3 8 30 10 oi368b036n
33n03h8x n 33hxD 6ob 66333013 Soihnb 3hb3630O b3b33m6ob 63833-
016001. hoi03b3(5 638200 33(1393680 00^501(10 34311636b 33 0 9 3(18 x 0 33(13 9 3 6 ^ 30
- 84 -
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6 *.
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^hi6 b3mh(inb b*03*o36noi 601(1(5030936*. ^hdmg, hm93b*(5 K+ 5*g*9ob oi*6-
9b036o 33(139360916 6*9i3,5 33(1393690, 6 *53 0(31361. *9nb 938935 6ih8m586o0o
836,506003(10 gnb6*c*6bn 01*309*6 *0150036* *(1101(15*6390 *6om636ob (6*5., C 1-)
5*9**950036001, m h 5*6 x n *6om636ob (6*5., 8191,50b) 6*h9 mj96ooi *6 3hm^m6on*
5*6oi*3ob3<33036ooi (pH 0 ^ 9 3 6 * 6153360b 5*p36ob*b). hm5mh(5 3^ym6*, 6 *533-
60b 5*(j>36ob 3hmi33b9o 06093630013*60 (10190 3*0(5o;39ob *hb36ooi |jo^m30*<b93ho
Ca2+-ob 5016,5(10106 g*5330Q36*h36x 93b30*b* 9 * >>*o0ni9xn6ooi 9 ob 5*001601*-
3ob3qj036*b 353013606, (1010300(5 *66(5o'bn6 0g*3*6cn*6 (A B A ) 31101*9 8015839366.
om6cn* j)h*6b3nhj(0 91801,50936x^1 363h500b 8 1(115^ 3 (ATP), dibfr) 5 13 9 3 6 1b
169366 369015363(10 b*bo56*0m 6030103(136360 9* 303350101(1360 (0*5., qgo^mSmh-
9016360). *853*11*9, 9pob 5 *68* 30m6*9o 5*bb6*-9*bjh3ob*90o 6*5ob 3 b*gymoj6i
0(53036* 9obo 15,5030160b, 5*63oai*h36ob, 6,5(13606 9 * *9*3,51(5006 895m9*h3m6ob
93b*6*8ob*9. 030,501301(18016360 0600363001316 5*30 36*b *693636 6*5ob *5,503-
hm6*'b3: A B A 66606 om63h *hb36b, hooi*(5 *0(5nh36b 6*5301* 6311300b 516660901-
60b b*hobbb, (50,501506360 2)0 Sohnjooi, 190936b 51(3136061900 8*oi 8fb*9ymqj6*b.
8(536*11301* 3*h*<bo^)36ob 803(1 5*9mymcgo0o ^mjbo^^ho 6030103I136360 bb3*g*bb-
31531(119 8mj9393&b 6*5301* 9mdh*rri6i'b3. ‘601503(1010 9*015*60, 9*5., Fusicoccum
a m y g d a lii- th ,50156060 qj3'bnim(5o6o (AB A -b 16,51501606,50) 6*53b „*930 36b“ q>o*
8g5rri8*h3rri6*9o, h*(5 06333b 9^36*hob ^6m 6*b, Helminthosporum mayalis- ob qjo-
^m^,mjbo6o 30 6150b g*060*b 06333b.
6150b 0PK#i*mbnb 3016^3(1^00

0138(5* 6*53360 h3*5oh3636 9h*3*0 bb3*g*bb3* cgij^m h^, 9*oi 9 rridhirn6 sb
dohooi*9*9 mho 91015160 i63bho536b; 3b36oi C 0 2 9 * H20 . CO2-001 3016,3110100
bmh(5O30936* 33h3p9 m hobx bo3h(539o 9obo 3*h(5o i 03ho 66330b (5300036001. 0obo
5130361 516^3301(136001 83^8663301 bo663030o, hm93bi(5 33op3h8obo 313(180
COr ob bb3i9ibb3i im6(536^hi(50ob 5130361b 51601590b. hm93bn5 3i3h9o C 0 2-ob
31(1(50193(10 66331 150-250 Pa-'b3 9i(pi0m, 6153360 gib ^h x^*; hm93b*(5 ob o9
90163^3 9*69* 3(539*, obo6o 0(3136*. bo6*oi03<b3 35h399m hobx bo3h(530o C 0 2-ob
3*h(5o*03ho 6633* 3(530*, (1*95*6 050 8mob0*h36* egoi^mboBo^'bob 3(101(53690. ob,
(1018 6*53360 p o s* bo6*oi03(b3, 06003630013*6600*9 5*630(101636300* C 0 2-ob
61503601606 *hi3oh9i3oho 5*3036001 (0130(51 i j ihnb 606101006 3oh9i3oho 5*303-
6*15). 6150b 5i(p361^3 C 0 2-ob 3016(536,5(1115006 5130361 5*6613301(136001 6*010*9
39069361 CAM 015361(133600. 0 ioio 6153360 p o ii Q>i0 noi, hmg3bi(5 C 0 2-ob 3ih-
(50193(10 66331 35h390mhnb369o 3(5381 0191,506 06,536603(119 6ih0mj06 ob 0393519
91 p o n 9(313, hm93bi(5 0191,506 91890b 83935*9 6i0oiymc3O0O C 0 2, 5hm393&*
33(139801(1063680, b *6 *8 ob 613(101306 03895m8 03^*6m0o‘b09o (ob. 616. 2 .7 ).
3016,5(10100 C 0 2-ooi 0113b 06366 6y0ob 93030(50,506 3nhm6369n, hm93b*(5 qjmov
03680 S i^ x ^ & b ABA-b 030(5330m6i, (11(5 630830003019 8015839366 61853,5 33(13-
93680 mb0mh3530*(5oob Shm^Qb’b j. *0ob 8393519, 6153360b 516660b 361(10 0116-
910116 5330019361 91 039935, hm93bi(5 9(536ih3b 6y0ob 9090 613036016* 9*3835-
(1361, obo6o 33(3>*h h3*5oh3636 5*h30m 0315,301(1011 cb30m503936*'b3 9 * (3x1* (163-
6016.
6153360b 0mdhim6 ob H7O-010 3016,5(10100 51(1519 51001061,5361 6y0nb 93030(50-
,506 30(10163680, (1019361(5 A B A -b ^ 6 (5 36 ^ 1(50 1 <3010101930 n(bh936i 9* 6*853,50
35(1393606 mb0mh3530*i5n* 836(13^936*. 83935*9, ^>*5301* 5*bb6o0oi6ob 36*ho
- 85 -
(jbhr^n 2.8 qjmonenb j33e> ‘bjpiSnh'b] bi&jcii {>s6e:>53&nb bnb3 nh;), b^nb bn{,i63 gi gphcnmbn, 6i&nb 0n6n6 bcDhn en^bn^hn 6n6:,:>(y>ag3;)m&:> C 0 2-nb
6n9 ihoi (0hi 3iCn ^ly^hnb anbogjncn). 5i6bb3i30&i qjmcnpnb 'bjgi gi j339^> abih^jibnb bi&nb gnqjjbnjhn 6n6ii(?6g3&m&nb 3iR3]636(xni 3 mhnb nb.: Napp-
Zinn ( 1988), Bolhar-Nordenkarapf & Draxler ( 1993).
6 ^b30 fn6 ^)^0i> anbaortoto
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5i0C'n'bni636Q36'b3 g i hR3&ni6 giîiÔ Sg&rnQit^mbiSn.
bi^nb b3h3Qob fcibb&nvnbnb bihnbbto 9 mj0 3 pn qjij^ynh^bn

bi&nb &ibb6n0mbnb bihnbbn h 35X nhg3bi bi 5 D6y3DtSCn3 1330^9 ° Snhmbsbncn
(ijbh. 2 .3 ), oi30(3i R i S ^ n 33h3g3bnb 9mj03g3bi gi9m ing36xni 9 icn 9 n6 ij>i6
9g5m9ih3mbi'b3(3. 5ih39m qjij^mhoii 3hoimbgn3n lb39mj93g3bncn, ^ 3 3 x 3 ^ 0 3 , bi-
j,nb b3hxn biSîgmg ihnb (pni. n93nicnig g i nbny bi 59 m x 3 ghmnb j,i 69 i 30 m-
bi 3 n ihnb bij,33bn 9cn0ni6ig pni, higj)i6 8 3 ^ 9 n33nioini n9 3nhmb3bnb cni5-
b33ghi, hm9o3bnu b30b 36ymb36 bij/ib b h x &ibb6ib. (6ib. 2 . 18 ). 03nh3b
9bhn3, 3jb^h30i03h 3nhmb3b3n, hm903bnu in d x3bb bijtfb 9oi0ni6ig ginb^hmb,
cnb6i|j bib5hd0n3i9, bij>9 img b9 nhni, J>i6bi])30ih3bncn 63i03nbib.
9 ngi g i j ljmh3bb 9 mhnb, i 9 9bhn3, 396n93630m3i63bni cr|n^n3mh9m63bn. ibb
Ijnbnb 9gi3i, ogi^n^nb 9s)i3i, un^myi6n63&n g i 3nb3hn0n&3bn 93gnb “bhgnbi g i
5i63ncnih3bnb 9 ih 353pnh3b 30 bi53h bnbvj 39i 9 n. bi&3 bb3igibb3ij)3ihig np3bi
ibi^nbi g i 5iB3nor»ih3bnb qjibnb 9nb3g3ncn - qjmcnpnb cgmh9nh3bnb, h33hmg3J-
I5n30n qjibnb gi6y3bnbib g i bnb3h39n.
(jbhnpn 23 jphjSm qjyj^mhj&nb &13C36} C3 gi C4 015361(13011 615,33611b fp>bb5i'b3 (Milbum, 1979)
5ih 39m cjiJjm hn iih ja m ojij^m hnb 6ij>nb iS ih i^ n b 3ib^>n
qjmljmbn6cn3'b3hip q9fn^mbn6oi3'bnb bn6i<nonori &i30h3-

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b39mJ33g36s Ligustrum japonicum-nb 6153361^
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- 87 -
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61; (1101936016110319 nb 03nd036i 5i0oiobil!)oib 6 3 6 3 6 ^ 3 3nhoi6368n CC>2-nb 6301-
3nb3&nb ^315003033*10 8ijbo8io3ho 86n936x fT'£>,1Qr' (Phmax).
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bi6&hd0n3n ghnin6 5168130016180 518(1830363060 ihni6 (815., <bn0o3ho bih^yonb
<301010108(53360 63360 g i 63fij3&o, 5310 30160 h35om636nb 091*101010(53360 9(536^.-
h33&o), q)ml!)oibn6oi36ob 3b qjnih0i 83bido363coi 86083630013160 nyoib 6 1 b -
3oh6ignb 6 10 16 60 6 0131(^31^06001. 3b bi8ihoiooi6oi 15I130133 8 6 316 3 8 3 3 1 6 3
6^01033606101306115: 8ihononi, Jomh36jo8nb <30100160601363*10 5180(1613100 0(5n-
h3i, 8 i5 h i8 nb bi3dihobni Ii3b3nhiooih30n 9161311150b 5 1( 13 3 3 x 0 616000b 3 3 -
bi3b 36ig. 1 8 Ri8ri6ioi3icb i8oii3h36b 53n8h36o, 301010013160 6 136360 g i o n -
J3636n, hrri83ooiii5 q9ml!)f,ibn6oi3'6n6 di0ni6 gi6ion 6oRJih3 161601013601. 6b3i>
<391^0111360116 3hong 3b 5i6 3n h ri6 36 x n i bubnSnoiynni oih5i6oi36nb d io n ^ 6
8150*13 6n6jnoi, onJ3636nb 8 3 8 0 1 6 3 3 3 :^ 3n ^31500)03*10 goq)3<b3ho 6n6m'oi6noi Qi>
ou>03b8o 1303*1010*1010903300 003016001603606 gngo hioig36oi6ooi.
6yoo6 815361*13360 J96oi6 5i 66b3i 336x 151053 353036: 3o i6Jo'T>6 nb 6yi09(536^-
*>336nb 8163363630011 8301*1361 ^3*iôrT13i 6 8(5361*13360116 5^ 6 30 36 30 *11. 3 n 6 in -
g i6 ou>3nb3q )cig 0(5boi3h36n 3*ioi33hxni6n 6yio0l536ih336n 083010110 ihm6 1 9 3 -
J3i0 3h ig 3*6*1363309010)0060 8063*1103*10 6030103*13636001 g i i 3 nooi3 qjm^rrv-
bn6oi3<bnb digni6 gi6io 96n83630(,i636b ndo330i6. 0 0 36 x ^ 3^ 0 n0obi, h m 0
6y(yib ‘601503*101 0o3*ioiq)oodo q9mooibo6oi3'bob bo6jih3 b3 x 3or'o^ 6hgo0nb 6 ^ q -
160136360b 1 8 8163363630b 31600130361, 6 id n h xn 6yon6 8(5361*13360 bi3*ionmQ
CC>2-nb 830i3n636nb 916100 361*1001 bi6nioig36i. i8nb 3*ioi-3*icnn 8063601 i h > -
m h 5i6 x o 6ib3 oh6 igooi 81010 i53gn 80181*115361. 8 n 3b 3g i3ig ndnbi, hoi9 0Q5 ^h n
- 88 -
cV)80D<!> 03 ^ 3(3 3 6 ^ 3 9 0
Ijbhncn 2.4 6»m-q)m1i>mt)n6or)'bnb b i93 som 3iJjn 9 iCDhn 3ifi33636e36(i m3tjn9iC3h

5nhmi>363n (3hi3ign i3,ynhnb 3nb3g 3fioi)
: ■•■■:.'/■■ ' ' ■. .:• '-7■: 63^rtw2mlyfibn6cn3<bnb rt&^36bn3iTi6i

%j6* 3«i> J63q3n U8mcn CO7 8&C02 &h (93h.
er263-> aib^boff’
b0X3wnl> aiaBilrnftn
6iCibei363&n
C4 ^5^ h 30&n 30-60(70) 60-140
c3
3(Di66nib 3hoiB(Yn3i6n 8(536^336(1 20-40(60)
MCii^X0 9(536ih33&n 6icrt3b363n 20-40 30-60
3bni5n 3i6fy,i^3&nb SjVngoôbn 20-30(40) 3060
905636(1^ gi bi6i3nhm 'bmpnb 81536^336(1 20-30
jp'bi0jb3C0b &3mcgnj3&n 15-20 25-40
6cnnb 8(536^336(1 15-30 25-60
9iC>i(T6iCib33pm6i 10-20 30-40
fihgnpnb i8(ji6n 8^36^336(1 (2)5-10 10-30
89hign 5i6nîl«i36nb 8(536ih336(1 15-30 15-40
}hijni3cn 3(536^3360 8-20
9ihu3(^36360 gi nbc3&n 5-15(20) 8-35
<33b3nb 330niihi'bnvj36n (1)4-7
P3hnib 339n3ihi'bnlj3{)n 2-8
CAM 9(j36^336(1 bn6i<7iQ3'b3 (2)5-12 0.3-2
CAM 81536^336(1 bn66x 39n 6-10(20) 1-1.5
b3-3u36ih336(i pi 63^^36(1
^n 3ni3(yi JOCi^X" 3(536^336(1 10-15
lyim3ni3cn 8(536^3360 b3j(53bnnb 3nh3x bîgni^ 12-20(25)
ljhm3ni3(yi (Vii636n (lxi6ioxyib qjmcnQ36n) 15-20
,jhmJnj3pn 63n8ni6n (jynb b^3(536ih3cni bn6iOTpnb ojmaiQ36n 10-16 10-25
^hmini3(vi 63n3ni6n ,jynb b3-6(536ih3mi Rhgn<vib O)moip36n 5-7 5-8
,jhm3ni3cn 63n3ni6n ijynb b3-8(536ih3cni 3iC+i3big
giRhgncgpn qjpioip36n 1.5-3 (5)
cn6npn txn8n3hn h350(^636nbi pi b361jhm3nj3&nb
gihcnmq)mo^>m3i6 8(536ih3<Tti bnbicnpob qjmonc3&n 6-12(20)
OT6npn <bm8n3hn h35nm636nbi gi b3&»$hm3fV|3&nb
(gihcnmg3PKJi{;m3i6 8(536ih3C7ii fihpnpnb <3^01036/1 2-4
gjmoimg8(5336n b336nb bn6idQnb a 01^360 10-15(25)
cgnKji(iig3(5336(i b336nb Khgnpob 03(^01036(1 W>
6(1638(5336(1 b336n 8-10 10-20
3ihig863i63 6n63m363&n 3-6(15) 3-18
8i65hnb jijnb b3-8(536^3360 4-8(12)
33hnmg3gig SShi^n h3jnm636nb b>>c3hmojnc36n 4-10(16) 3-10(18)
3ig836(i 4-10(20)
&;>3&3i3&n 4-6
3gi66nb 3(5nh3 ,bm8nb 6^360 (3)10-15(30) (4)8-15(35)
ii3&ghnb jmbgihi q)moing8(5336n63fij36n 6-15 15-30
l)36phnb Jm6gihi 8ihig063i63 63^^360 3-6(10) 4-10
- 89 -
6ab30r t t o m 9nb&>(f>0iW> aa^Ô 6030)03rt3M)b P ^ a n d 865
Ubhncn 2.4 (5*{>hd3e3&*) 6>yo-q3m^ml>n5o>3<bnb b*33*om 3*Jbn9*03ho 6*8336360360 nijâscijh

iihjSm 3ohm63&6o (Sh*3*oo îjmhnb 806393001)
63l5m-ojmljrobn6cn3<bnb 06^^103(^6*
8(536^3013 553030 (i8mcn CO, 85 CO, xh (33h_
9"26&:1 8*b*)bon*
ihnS^rTfchbfl^bn
$308(1360 pn* 3*60(^360916 6-10
53081360 gsfiheo03po 3*60^3609*6 2-5
b*3bj6o 2-3 0.6-3.5
90^36360 0.3-2(5) 0.3-2.5(4)
Bycob 8(536*113360 9* 6y*e3(536sh336fl
^*m6nb 8(536*113360 12-25(30)
9î 6sho Bypob Si^hmgn^jdo (5)7-10(25)
cnonmhicjho bm6ob 93hs 69*06(536^3360 2-6 5-30(50)
30^&kyn63ho 69*08(536*113360 (2)10-30
(^/O^t, CH bor1^ *
(5bhooo fi.5 63lyn-qyn^mbn6cn3cbob 8*jbo8*03ho 06^36^3106* m3lyi8*C3h 3ohm&3&8o (Bjorkman et

al., 1972; Seeley & Kammereck, 1977; Patterson & Duke, 1979; ) smond et al., 1982; Nelson, 1984 ;
Marek, 1988; Ceulemans & Saugier, 1991; Nobel, 1991b; Duftene & Saugier, 1993)
6(536^3 H Smcn C 0 2 a-e 63-’
C 4 6(536*113360
Cenchrus ciliaris 68
Permisetum typhoides 64
Saccharum-ob 53*hnb Sn&hngj&o 64
Sorghum sudanese 57
Zea mays 55
Tidestromia oblongifolia 50
c 3 8(536*113360 (6*c*bm3636n)
Camissonia claviformis 60
Triticum boeticum 45
Typha latifolia 43
Oxyza sativa 40
Helianthus arruus 28
Glycine max 27
Eichhornia crassipes 20
CAM 3(536ih 33&o
Agave mapisaga 34
Agave fourcroydes 23
Opuntiaficus-indica 20
C3 63-8(536*113360
Salix-cfo 53*hob b*b3m636o 20-35
Populus tristis 30
Populus-ob 53*hob b*b3m63&n 20-25
Hevea brasiliensis 20-26
Elais guirteensis 20-25
Fraxinus pensylvanica 20-25
Ailanthus altissima 20
Prosopis glandulosa 20
Eucalyptus pauciflora 15-20
Pinus sylvestris 17
- 90 -
03^5 30366fn3<30
6y*0O 33mji3b 9**600136001 ldO-jjh 830 CO2-6. 309(13 Syjhn, 6yonb 8(536^301*
qjmcnQ^bSn 3 nb 3 30*633*6 b^nhg3b* 5*(5np3booi 30^01 gogn ghm, 3Qph3
b3 xD°>nb 3(536*1130)* q)moi03 b0 n CC^-nb 03 b x * b - 5^hgi *8nb*, H CO ' -nb
5*g*3oidh*33bnb 83J*6nb8n 80i,6*hn 6yoob 3sjJhmq)nli)363n 3qghoi u x^P >^nb
5*63001*1136300, 3ngh3 b ^ n b 6y*o3(536*1133630.
gml5mbn6cn3lbnb 36*hob 6 D636 ni)I^l;>e 5 *63 nho\b3bxg3jon3cn
3 (536ih 336nb 33hR33ob b*03*O3b*b 8 o6 *oi8 fnj33 g 3b*3 o, 83b*c>3o»b*b* g i 830-
y333m6 i 3 n. cgm^mbnbcn^bnb bnRj*h30i* b33i5ocgo3hn 5*6663*3363606 S n ^ V jb n
36g* 333360101 qjmcncnb boh^D^nb oi*3nb363h3b3b6n, n8 *3 n, 013 h*8-
g36*g *g3og*g 3 mdh*oibb 5*bn 33 h3g3 mhnbx bnbo383b3n, 6*533606 qgmhd*b*
g* 3 nb 5*6*60036*80, i*h6ov)6n0nh36nb <33(1636001* h*oip36oi6*bi g* *jon3m6*3n.
(ibhnpfl S.6 COj-îtCTi (i3Ci 33hb3ilb (Populus) bt>3igibb3i y v i 63n O ioipnyijh 3nhm-
6^69n (Ceulemans et al., 1980)
3jhb3nb jjc<n6n 63^rw2fn(jribri5or)'bnb ftsgnb grtapbrrjhri 3ngi 6ihfi36ft

aijbn&jcm 6n6»p3eoamfe> "
" 3mo>i COi a-sga -1 68 bd-1
Beaupre 12.0 3.12 8.03

Unal 11.5 2.78 8.45
Trichobel 11.1 3.25 7.08
Ghov 10.6 1.65 14.08
Italica 9.8 1.77 9.74
Robusta 9.5 2.09 11.36
Columbia River 7.0 0.73 21.28
Nigra Ghov 3.8 1.50 33.56
2 .2 .3 9 no>mjm6 e h x n W 6 cnj 3 nb b SD b n qjfD h n ^ n 3 n 6 i
81536*11301* b36ov)3noin *Jon3mb* Q*0myig3bxn* 3 *oi im6bonoXftDh ijn36*

gi 536mon363; nb 5*6bb3*336xn* bb3*g*bb3* 01(15*6016* g* 5*0my363bxn 636-
b^hi^nb, *5(130133 8(536*hnb 5*63noish36nb cg*bnb, 8nb *Jon3ro6nb gm6nb* g*,
*51130133 5ih30m 3nhm&3&nb 606393001. 53630033(10 5*6663*336360 5*63nhfo&363-
pn* i*o*6m03h g* 3*653001 3hoi(53b3b3n 8016*6003 386063630013*63^ 033(183603-
bnb - n'6mq33h836(!)3bnb *hb3bmbnon. 8*Jti3bnb 8*p*0n 3301533001b*, h*b*(5
33 ndp3b* *95000 ^016936 C 0 2-nb 06036603(10 *bn8np*unnb*b *6 g*b*0 03833(1-
*03h*b3 b*b*03b0ob p*3 pnb 33g3b*g, 06333b b36oij3ob 5*dpn3h3b*b, 8 *3 n6
hoig3b*i5 ("*63 9* 6 *b3 nh6 y03bnb g*6*go 330U330mbob*b b36oij3ob 06036^301-
6* 01*69*01*6 d^nhgjbi. 63601^3* dp03hg3b*, hmg3b*u oj8636* 830*60103(10
363h5oob 5*<bhgnb bs^nhm^b*, 8*5., bhgnb, 001601* 830i3ob36ob *6 boh3bnb
- 91 -
6^b3 0 6 6 ^ ^ 0 b anbSifoOM 8 3 6 6 ^ 0 6 ()3 (J)03(f>3 &0U P^ 6 8 nd 96 ^
gidQ33nb ^hmij^b^bnbi^nb.
9(536^3360 <^>6663339361^6 3hoi9 i 630 iob5i 6 b^fxn^now ^03016001: bib^m-
6hn3n 5^6663^336^ 1 :10-gi 6 l:20-9g3 03hy3m6b. Qmhajmpm^n^hn, ^rncgn'bnm-
Qmjjn^hn g i g}36junm6iC3hn (jn336 nb 93gih 3&i ybigymqjb (ubh. 2 .7 .), hm9
9 bj,i3b 3nhm6369 n 63601^306 o6lj36bn3m6i 6iQibm3i6 9^36^33690, 5i6bi2>3ov
(136001 bBhiojig 9 bihg bib3Pi&369o mh33h 3qjhm 9 ipicni, 309(13 qjmcnmQ0[5336
633690. i 9 3ii6i6i63ccrii qjmoiQ3&n 30 biSîQmg 630133b 303(1™ i j ^ h i g
63601^336, 3ngh3 9ihig963i63 91536^33606 biibn9 opi(yim fnhj,i6m36o. 3hcnbi g i
o9i33 i6 ib0fn995m9 353033690 bo6 ioi0 ob9 my3ih30o 91536^3360 63601^336
(20®C-b3) 3qjhm 06,436603(^9, 3ogh3 fihgoQobi9 lji 66 o, ihj^nyibi gi 03(?>cn
(5bhopn 2.7 bo6630ob (9o,ynjm69hn3cn) b^5cnj3nb o6,j36bo3m63 £)h^0m3a6 8(5363113013 'bhpsbhx

qjmcnQ369n p j 9 nb bs6ynb qj>'bi3 n SOOC-'bj (S h ijip n S3j)fnhnb 6nb353001) ( 9n,im;jm6phn3(yi
b^6oij3nb n6^36bn3m6:>3 n C3 p:> C4 9536^33680 h:>n63 :>hb3&nain 5^6663^336^ ^0600636^.;
Byrd et al., 1992)
COj-nb fciSmymcgnb
at535ih30«> 363^ (i3(X>n COj a6 C02 bh (83h.
a-%^1 3ibi)bcn‘!
iXiU^3Cn 2>i0^bm3i6n 630^360 2-6 3-8
33C3hn 6ipibm3i6fi bo63oi0ob8my33hxn 815363(13360 3-5 5-8
3393(10 &303bm336n fihpnonb 38^360 8(5363(13360 1-3 2-5
339n3ihi'bnli)36n 3 - 5 (8) -
3mprrOsh:>'bnl!)3&n 3-6 -
(ijhmSnix0 b336nb bn63Oi0nb qjmoi03&o 0.3 - 0.5 (2) -
jhm3nixn 63360b Rhpnpnb qjm010360 0.05 - 0.2 -
qjmcnm08(5336n 63360b bn63oi0ob q)moi03&o 1-2 3 -4

ojmcnm08(5335o 63360b fihpn0ob cgmcn036n 0.2 - 0.5 1-2
lbm8o3ho bih^ypn6 qjihomoijmcn0fn3i6o
63360b bo6icn0nb <31*1010360 0.8 - 1.4 - 0.7
'bm9n]hn bbh^L^nb q)3hoiniq)moi0m336o
b33bob fihgnQOb 0^010360 0.2 - 0.5 - 0.3
bmhjiQ^ho "bmBnb 83(139863363 6063^13360
63360b bo63oi03boi36
835336300 6063360 0.5 - 0.7 - 1
bmh3303ho 'bm6ob 831139863363 6o63m336n
63360b Rhgo0<ru>5
93633&Xn 6063360 - ~ 0.2
Rhpo0m3oinbi 93 83C>308oiob 6ho(53(5333-nb mjibob
63118(11839536300 5333 qjmoim08153360 63^360 0.6 - 1.5 2-3
Rh9o0m3oiob3 93 83C'303oiob 8ho(53(5333-nb mjibob
63h8m83953630o 5353 831139863363 63^360 0.3 - 1 0.5 - 1.5
39366nb 63^)360 93 63633(139 63^)360 - 0.8 - 1.5 (3)
CAM 8(5363(13360 - (0.2) 0.5 - 1 (2.5)
6y0Ob ^3h^Q(Ti336n 8(5363(13360 - (0.5) 0.8 - 1.2 (1.5)
85 O2 6*1 (99h- 83b3)bor1
9 onob 91536^3360 ^cghm ijô^hig, 309(13 oi6npo 3i3obi. q3Pioi0m3i6 bi36360n

6360^306 o6li)366o3fTi6i gi6ipOi (1-2 9 g, 5"1 bar1), 01^363660 ob b i^ im g cgihcnm
- 92 -
<W8015 0 3 ^ 6 9l33Gii(n0 3 n
bi‘bp 3h369n 0îyQm6b dmhcgmprr^n^hn 3 5 3 6 x 3 6 0 ^ p i 53 m5hiqjn30n 53636-

np3&nb 3 n b x 3no) (0.5 - 5 85 5 '1 ben'1) [28 , 194 ].
153CM X 9l5D53h]cn3 933300360 93 63ynq)36n 63601^3336 3 0 ^ 0 6 ^ 36 ^ ^ 39 ,
3n0h] 03cnn cgmcn036n. li,mti)36nb3 93 5383^636 x0 0)3b336nb b36oij3noin ij,yi3m-
63 nb3cnn333, hmftmhu p3hmbn, 33h$nb yomh^jdn 93 63(1000 j,n 3qjhm
n5v535bn3h3g 63601^336 (63b. 2 . 19). bb33g3bb33 b3b]m63bn b id in g 5i6bb3i393-
6036 (p3hmb b35oiJ3noin 3^03^161101 (2-936 5-693 9 & C 0 2 • Q & 2 ■ ben-1 y333nQm-
6nbib, 10-20 b0 9n303(jhnb 9Jm63 (j>3hmb 338oib333i3n) [209 ].
5383^636300 y0mhii3bn 94 <3363360 dnhnoiigig 6ih0mg536npni ^hjncn,
338603^013 93 (J36vi)hng36 g s S m l^ x n 03hj6nb 33h3g36noi, hm0p36n(j ij^n^hig
b36oiji336. bb3igibb3i bnbjnb ypmhoj3&nb, g i (n3hm36nb b36oij3nb
n5vj)35bn3n6i (0ibnb 311013x^)3 5i0fTiyng3n0n C 0 2 63900 i6 83 hi0 6m6i'b3
5 ig ii 65 ihn936 ncn) 0^0(19363 gni03(jhnb 5igog36iboii6 3honig, hig5i6 ©unhgQ&i
j3hjnb 0ibnb <3ihgm6i 03hj6nb 0ibiboii6. 33hng3h9nb t^giSohnb 3h0 1 3 x ^ 3
5 i i 65ihn036 nbib jjn (33hng3h3 nb {b 3g i 3nhnb 3 hcn3x 'b 3 5 i 0 mymq3Opn C 0 2)
b36oij3nb n6^35bn3m6 i 3U303pn hR36i.
30nb j>i3 m, 0n'bi6336m6n0ni 0j>3gihn 03ji6ni3hn jbm3n036nb gngn him-
036m6noi 030153300 8tf36ih330n mh5i6m36nb 33 b6 i 3pnbib b36otj3nb n6^36bn3m-
6 i 5,i0rnnbi^mb lingnb i6 (3npm3i6n i'bm^nb 339l533Cm6nb 3 n0 ihoi.
5sb. 2.19 Pinus densiflora-b bb3igibb 3i

gniS^^hnb ("3hm]6nb b}6oij3nb n5ii35bn3m6i
eii6{>ihn036xn tBhie 6m6:>b3 (RDW) gi
qjihomd'bj (RA) (Negisi, 1974). Abies
amabilis-nb 9m6nj;)936n nb. Sprugel (1990) Stem diameter [cm]
- 93 -
6 6 b 3flff)W m 9 n b 86 A 066 896^0 G030103(f)060b ff-bff>8nd866
2 .2 .4 6^63nODh36flb (3^ <bflW e } ^ n 3 H 6 n b 9e6na^hDn6flb

6D3C36D bD6Cnj3^b^ Q} <3Fli!>rrlbn6aiD<b,bD
9 ih o u > 0 n i, hmj)mh(j b ] 6 a ij3 n o in ij^ n s m b i, n^ 3 q jm ^ m b n B o ^ 'b n b n6^36bn-

3 m 6 i S ^ B i h n b b ib ^ m b h n an 8 ib ib n :> c n 3 b 0 3 b o i, 0ij)h:>0 nbn 6 n 9 3 9 8 0 3 b n g n g33b b
ih 6 ih 0 m ig 5 3 6 3 6 . 8^36^360 8 n 8 g n 6 ih 3 fc ib o ii jj3Qb 33n d Q 36 b 830153^0r^b bh:>
9 ih ^ m 8nbn n 6 g n 3n g 3iQ 3h n $ i6 3 n c n ih 3 6 o b 5 i 6 9 i 3 p n b 5 9 n, ih i8 3 Q b 3 <bm6nb gi
9C13 -( j'i8 n b b b 3 i g i b b 3 i gh m b itf ^n.
b^fjoifoi gi 8u36ihnb ibiyi

ibiOftibhgi 9(j36ih33bn b36cnji336 3qjhm n6o36bn3hig, 3ngh3 b5 ri3hn;
5i6bii3cnh3bnc7i n6o36bn3hni 9u36ihnb 9‘bihgn 6i6np3bnb b36oij3i. i 9 ghmb
9d p i3hn bnbo^t^hn 833im6nboi3nb 3hm(jfTi3pi'b8i ny363&b ATP-nb b33hig 830
himg36rn6ib, 3ngh3 3b bg3bi R30DC0bhn3n ybn33p8mJ83g3bnb ghmb. bi^nhm3bnb
838cnb333i8n, 3jh3g3bb 833dpnicn b3601^30010 ATP-b 6ih8mJ86nb DiliP 39nh]c
33ji6 n b 3fb 3 5igih o i3i. ip8m6i|j363b9n, 0 ^ 3 3 6 0 6 633hm368n, cgmcnpnb j>i3pnbib
g i 9b ih g 6iymq)3b3n b36orj3nb n6^36bn3mb;> ("im6bli)h3j|jn3pn b36oij3i" i6 3
"bhgnb b36onj3i") 3-10-33(1 iîh&3bb b36oij3nb n3 n6o36bn3mbib, hm83cnu 6333-
03 bhn3 3 nhmb3b3 n i(pn6 n963 bi ("b36oij3i (5bm33p3 mj83 g 3bnb 336 ihfi363bnbicn-
3nb"). jbm3npoii gnq)3h36i5nh3bnb, 8icnn 5i63noiih3bnb gibh3p3bnb 3ihip3p3hig
b36cnj3nb n6o36bn3mbi nb33 iCD^DCmbb g i 6mh8ip3h gm63b 36 (136 0 3^ (6ib.
2.20 g i 2 .2 1 ) . bb 3igibb3i bib3mbnb 8(536ih30ii qjm oip^bi g i 6iymq33b9n 3 m-
63h3bibcni6 g ij)i33nh3b x n giSpnb 3hmu3b3bnb gi6y3&nbib 93ndp3bi ig^npn
3jm5g3b b36cnj3nb n6^36bn3mbnb ghm3bnai 3.6. bhgib. 3b 803010-
0136b 930ibm cnb8 nb 1530003^36^3, hm8o3bnu 3ihjnb 5i3Q33h303bibi g i 6iymqj3-
bnb 8n3h 8 3 li)i6mon'b9nb inhm3i6n 3hmg3jlj3bob (8ig>., 3Oin036nb) j,i8mymcgb3 n
30n6 g3bi.
6ib. ZJBO qjmcnmpa1^336(1 (S]6^nhn bibnoi), 60631^3*60 (361)33^0 bi'bncn) b^jbnb g i cnQhrn^fenb
(63hl5npm3i6n bi'bnon) 9nd)mjri6ghnxn b]6onJ3i bb3igibb3i qn6m(vn6n3h ajibiSn (Eberhardt, 1955 ;
Pisek & Winkler, 1958; Negisi, 1966; Malkina & Tselniker, 1990; Paembonan et al., 1992)
- 94 -
(W80 to 0 3 ^ 903666030
0 . 5'
O)
E 0 25
o
o
12 20 26 ST 44
Duration of development
[ Days alter leaf unfolding ]
6ib. R21 Phaseolus vulgaris-tb 3nh3x^pn ojmonc^dnb jiSpnbib arymjmfighnxn b]6oiJ3nb n6^] 6-
bn3mdi. b]5cnj3nbb (RM) pb bhgnb bx0;)96ymdn b]6oij3nb (RG) Bncjbn bjfjcnfonb bighorn
n6j)36bn3m6i9n (RD) (Kase & Catsky, cit Ticha et al., 1985 9nb3P3noi)
grTHy*ibn6o>3'bn g i 53630013^1363
0y363hnb 5363001^3606 3hm(j3b3n 01530363 8nbn qjm^mbnBo^'bnb 363hn.
ipann53636nb3 93 t^nh^nb 538906 qj3lb39n qjmlj)m6n5oi3‘bo6 3^03(^163 635hd6m6-
039 fi30mhfi3&3 63601^336, hm030n(j 30 cg3(b38n b3303mg nB^Bbr^hnb. 36 536-
3nhm63630O3 n0noi, hm8 cgma)0366 33b 3h 803^6330301 638330m tbm838g3 93
9(joh3 h3m936m6nb 606301936 801360^3336, Jom hm l^b^&nb 3^31*16315 33(1 8393-
h36noi 9363903 93 y>h6mJ6 o0oh363b 3h 803^63303 83b3d0360ri6nb ^033893 (63b.
2.22). 801903639 53630013^36x0 33hjn oi33obo q3m(jmbo6oi3<b3ho 93 b3d 03 &0 m6 ob
83jbo0306 3030336366. <bm503hai 6303bm3368o cgm^mbnBcr^'bnb 3^031^63 6 x
h30936o03 9(nflb3 013 33nhnb 838935 3333 030^ 113636 o6y3&b 93 36330136 3hoi3g
b x 3cghm 93 jqjhm 03936b. qj3b336ob 33gm8ob 9369363893 8(joh3 b6ooi 3gh3
n6y3&3 j0mhmgn0ob 93893 93 ;)0mhm303bli)3&nB 935363(131503, (1363(5 cgm^mbn6-
013'bnb b h x o 936y33(33 8mb933b. 66(130)39 8b3hg 6303bm336 05363(13013 yomh-
0)36ob LS30i 3X n cgmoi036n 93 nd03&3 b35hd6 m603g 536bb333936m9636 q3m(jmbo6-
ai3'b3ho 363hooi, (13(5 83010 9oq33h36uo3ijonb 095m03h3m6ooi33 53630(1(11636300.
3b30-3b30o cgmoi036nb 53803 93 33h3ob 001003639 53630013(1363 3'bh35330 ymq)b
63b9nh639ob 93Oi30b36nb 53636 9 33^ 03 03^3636, 3b3, hm8 0(5363(13360 3333
'bhgnb 33hnmg9 n 3(^63336 C 0 2-ob go jb 3(jnnb b 3383 mg ^336003(1 gm63b (63b.
2 . 2 3 ).
(bm0n3hn bsh^yonb qjmoim00(5336 b33&9n I 93 II qjmljmbob|!)3036o 633636noi
R38my30n63630n3 q)mai036n6 5390ng36 20 gpo6 838935, 363, hm8 qjm^mb-
n6oi3<bnb 3^031^63 3n3b 3(p633b lb 3qjb30nb g3b36ynbnboi3nb. b3 8(5363(13013 bb33-
Q3bb33 b3b3Pi638n qjmli)mbn6oi3<b3hn 3^03^163 86083639^13639 536W53339363
3hai0363Oinb3536 93 3b 538m633X03 gmoi036ob 536300i3h36ob 06g03ng3303hn
hn^Snoi (63b. 2 .24 .). 5363noi3h3&nb 3390063(5006 33hnmg9n ( 30n03jiJ3h x b,33go-
3^3) liyob b3 6(5363(133606 (835., 6333(166390, (1^093, 83b3) cgmoi036nb gjm^m-
- 95 -
e ^ b ao rtd ^ o ij 9 r)b 8i(n36 ^ 8 9 6 6 ^ 0 6030100ff.a&0li ffArt0Fld866
6ib . 2.P S JpmhmScib^yib b^jfcjijhnbi g i qjr,nj^Tibn6oi]‘bnb ]6shnb u3(yiQ36i Phaseolus wlgaris-nb

gmoipnb 516 30 0 1^ 36 0 6 Jcmhmipib^Sn l^jhjJvjjhxn y i 93 r»)636 &)dnb <7ii 6 iqjihgm6i. PG
3p3b^m5pm63p36n (aih(jb36i bjhicnn) (Kutik et al., 1988). Phmax qjm^mbn6cn3bnb SijbnS^dn; ip
jjsS^yhn 5^9mbi3ipn; Xc bn6soipnb biim9335bnjnm 63h1yipn (Catsky & Ticha, 1980). (9^3336 :>
bjhicnn) ^hn 3f t i x n 63fi3n>6n jynb ’bnanshonn Suj&hnb ib:>c5ibhe:> qynonmQSn RuBP jshimjbn-
p i'b i ssjdjcnhQ jbs dpnjh 63^1 (Kursar & Coley, 1992 )
6ab. B^ 3 63^m-ojr»vynbri5oi3'bnb
<>6^36bn3m6ob 9^9^30936x36^
0(j36ihnb J jii'b Q . 63^
0 m^mbn6oi3lbn bmrv>b 3nh33Q,
03b>93 , 3^ 30 13, 9393093 9^
03(jbh3 qjrunmQSn Cb3g> bjhiom).
ojmoipnb ‘broQnb gbQmyi^XD6^
9nb oob56n6Q33hm6iboib5 (j30Q:>
b^hiom ) (W oodw ard, 1976)
- 96 -
•W8016 03^ 90aB6(f>0aD
Betula Acer
Months Months
6*b. ££4 (gm^mbn6o>]brlb b3'bm6]hn i330nC06]6n SnmSjhx t>3 9i}j6ih33n (Betula maximowicziana)
gi bjJu^bnnb 6mpm bîgniBn 9ymqj 633680 (cgmcn(>n3i6n (jyj, n*3m6ni). b^gi b^hicn'b]: ibndn-
giynnb S^hnmgfib 3i 6dnc'b3 qgpKncaftnb 5*631101^361 (hr>i}b33&ri dnjwnwaftb qjmoiQftb 3m'bn(yiib);
831 b^hioi'bo: 0)ml)nibn6or)'bnt> nBtfjbbnsmdi bn6*cnQnoi 6i 3Dh36obi g i m3)yidiC3hn &)03]hi'£|hnb
3nhm6Q63n; j33gi b^hiofbj: 0n,ynjm6gh n xn b]6<nJ3i 20®C-t>3 (Koike, 1990)
6ib. ££5 Solamm melongena-b qfn&rtxtfknjtfoj

6iymq)9bb9 mnihm6nb 6 i 3C3î - Ph 6*yrrajni6n 8)536*-
h33&nb qjnyjmbnBo^'bn; PllQ ]6iy(nqjn 6u36ahnb
ojmîTibnecna'bn; S 6iynqjns6 815361(13311 bijihm'bnb
ihb3i>mi)i; S(j }6iymqjm 9(J36ih33n bijihm'bnb
ihihb3&m5*. g)m^mbn6o>3'b'b3 9mj93gn JOX1
3ynpm6n3hn qjilynhn mhn33 939onb333i5o
3hcn6*nhni (Claussen & Biller, 1977)
- 97 -
GabaOrt&Ôb 9nb9i)(n36i) <56 93rt6=?0 G030)03to060b B^9nd866
b r>6 o i 3 (b 3 h n 3^ 0 3 (116 3 9363o n 3 , 8 3 5 (13 8 nb o in o » J9 n b 3 (5 3 C 3 C n h fijb i q jm c n Q jiin b

5 3 y 3 no>Q 3 6 3 9 g 3 . ^ynb 9 3 (13 9 8 6 3 3 6 3 8 (5 3 6 3 ( 1 3 0 1 3 3 8 (1 3 3 0 3 1 ) m 6 3 8 n q jm ^ m b n E k n ^ 'b ^ h n
3 ^ 3 0 16 3 , b33hcnm 5389n, 5363nhoi6363 <yi3 qjmoi036nb qj36Junfn63p3 hn bj^m bnb
b 3 6 jjh d p n 3 r T i6 n c n 93 600936 693893 0 )3 6 9 3 0 13 6 8 1 3 0 (1 9 3 6 3 .
y333ncm & nb3 9 3 6 3 y m o }8 b b 9 m n 3 lim & n b 0 ^ 3 6 8 0 q jm ^ c n b n 6 cn 3 'b 3 h n 3 j^ n 3 m 6 n b
53d 0 n3 h3& 3 830866333 6360080, 8 3 (1 ( 5 3 0 0 1 3 3 6 3 3 0 ^ 3 (13 6 6 3 93 3303b 8 ( 5 3 6 3 (1 3 3 6 8 0 .
6 3y n q )3 6 n b 8 1*1( 5 0 9 3 6 0 6 3 6 C C ^ -n b 8 3 0 13 0 6 3 6 3 b 3 j , h d 6 rn 6 0 i g 3(538 3 (63b. 2 .2 5 ),
( 13 9 5 3 6 6 3b 8 nh6 y0 3 6 n 8 3 0 18 0 3 (7 3 ( 1 & 3 9 3 3 9 n 6 3 & 3 . q 3 n l5 m 3 m h 3 m 6 3 6 n 5 3 (13 3 3 3 0 ( 10 19 6
0 13 8 3 8 0 16 3 6 3b 3om (jn S n b h3 5 X i b 0 38 o, (1 3 (5, 0 13 3 0 b 8bhn3, 8 3 0 13 6 3 6 9 0 16 0 0 0 3
8 oi0 n36 n 8 (5363hnb 3bn8 n03i5n3h 3(101^360136.
3p b 36 n9 6 33n3 3 5 I1 3 0 1 3 3 q 3m l3 m b o 6 o i 3 ' b 3 h n 3 j^ n 3 m 6 n b b 3 <b m 6 3 h n 15 3 3 0 3 6 3 9 0 16 3 ,
h 3 (5 5 3 6 3 0 (10 16 3 6 3 9 0 3 5 3 (13 8 0 1 3 n h m 6 3 6 o i3 6 9 )5 3 6 3 h n b 3 3 5 3 3 6 0 0 1. 8 3 5 ., <3 0 1 0 1 0 1 0 -
Months Temperature [°C]
63b. 2 i6 a ,b 53(13801 qjsJvjmhjbnb b3lbm63hn 53000360b 5330363 qjm,ynbn6oi;)'bV|. a

qjm^mbn&Kj'bnb r*6^36bn3ro&nb (53000363 Pulmonaria obxcura-dn 93b63hnb J33<ju38o.
93(p3on 90163538360 (b j30O b ib n ) 900*3630°^ b33&nb 83qjm oi0338e3 b3i)3C330 8 5 3 -
63hob ^ 'b s q jb ^ co 'j gjoxj>0369o. hm^mhij i,n ^ 3 8300901010363 93 Jjyjyjd g O 3J>33 bn63cnonb 9 3 6 ^ ^
hjb^hbo 3(p633b, Pulmonaria obscura-b 53(b 3q)b30nb 301010360 053033 93 nbnBn 0530363 Rhpnpnb
38^360 ‘bsqjbjoob <30101036001, boiooi 8015303636001 - 83801650130b <30101036001. 83010
<30iôibn6oi3'bn 538oib3b3Q03 ort>3oo (“b iq jb x o b ) 93 SjS^yihn (83801650180b) 63^36001. 6 ib i'b o b
^3^3 Sbih^b 9 3 8 ^ (^ 3 3 sb3bi3b ,jyob 83<goioio3ob oii68n 8g33hx 303336b (G oryshina, 1969) . b
6830013833^01330100 63633(139638^ Phlomis JruticosaAj bjbw 6 $\sQ go9oihqj30n gpioio3&nb
qjnooibfi6oi3'bn 93 1>,3933h:>£)hnb mS^nS^dnb b3t>ni63hn 53000363 (539363530363). W 'b^donhnb
<301010360 (5hnoo 63080360 30(1016360); S b 3ojb30ob 0)01010360 (39h3on 30(1016360) (Kyparissis &
Manetas, 1993)
8(53360 (jynb 6303bm336n 8(5363h336n 8 3 5 3 3 6 x 0 3 63360b 6 3 9 3 n3h3b368n 5 3 8 3 -

330n bn63oi0nb n6^36bn30i6nb 1533036390163b (63b. 2 .26); 'boi8n3hn b3h^yQnb
8(5363(13013 gjfrioi036nb 93 83h39863363 6n6336nb q3mljmbn6oi3‘b3hn 3 ^ 3 0 1 6 3 'biQ-
oihnb 801369013360636 6(50(19363 839n63(5, hng3b3(5 yn633 33(1 30 9 33 3(1 3(?n6n3-
6363 (63b. 2 .27).
- 98 -
(W8016 03^ 9036^6030
Monltis Months
Bib. £27 b0xo>i93i'b(n3oinl) qjihoiriqjmcnpm3i6n 3ihip963i63 bjQbnb qjr'Vjnibn6ov)'bnb bjbmb^hn

U30np36]6n. Sih^bî b^hxn'b]: Olea europaeab (Rhg. n^pnngbb) qjm^mbn&Jij'bn, jp’bmdnpn b^6-
g ih ^ x 3nhm6369n (600 |J9mgm qjm^mbn 9‘269‘', mS^nSip^hn Bygno' 6^33^361, 30
Pa COi). a 9n9gn6ih3 6gnb ojmoigîn "bxgbxnb gibs 6ynb9 n; b 9n8gn6ih3 Bpnb qjmong^&n ‘biojb-
3gnb impmb 0mjjg3 63ig3nb 339&36: c 9ndgn6sh3 Bgnb qjmcngjbn 939mg{,m0no) 6309056 S^hnmgSn;
d 'bsScnihSn, 3nh33gn yn633&nb 6n6; e p i 9nb 6:>yn633&nb (-5®C-9g3) ^nm gSn; f 6n6i 6gnb
qj(TKnc3t>n jp'biOjbx't’D (Larcher, 1961). 6^33361 b3hxn'b3: Arbutus unedo-b (3mh,i)35sgnnp:>6)
qjmcngoinb j ^ ^ n {,idmbi3x>n. (p; qj3hgm&n, b igjy <y>6iort3&nb 6hqjn3n g i 3mj)ng;)&x 3£>:>
iC<n6n3636s. • 'bhgibhxn qjmcnrifyi; o ibign opcnmgo (Beyschlag et al., 1990)
2.2.5 hD3H qjîbnhOl^ 6^3036^ C 0 2-nb 1330^3
CC^-nb 1530^3 6 ^ 30 36 ^ bbg36b j>ih30mb bb3igibb3i qjij^mhn. qjrn^m-

bn6cn3€n cgm^mjnSrvjhn Shmuobro gi, (jbigni, 3nh330 hnj>9 n nb 9 i 3 mj,n9363 (}ni
9 bnb g)i3mbbn336b‘b3. cgm(jmbn6ai3'bnb 6 nnjn3 n^hn 3hmu3b3&n yi j,i63nhm6363-
Qfii C02-ncn, li3033hi(i)3hncn, 6yoncru> gi 0n63hiO3hn 6030103113636001 3(j36ihnb
3m9shi536non. &i£,336nb {/boai qjmoimQSn C02-nb 03b3c> dohnaiigig QnSn^nh]-
63pm, Fib0i3ii 33(139366(1 5 ycnb Sm^Bunignb Qi^^Snoi. b;j6oij3no> fciboii 1530^3
3 mj03gn qjvj^mh^ngib 1^330^3 36 n9363 pm3i 6 n ^ S ^ h i ^ h ^ . 55(130mb
gi3i6n5d3h36x>n ^mjbn^hn 60301031136360 3ihymqjooiig 3mj03g3636 y33C> 3hm-
I53b‘b3, CC^-fri'boib 15300b R>cn3onoi.
1530^036360 CC^-nb i53C^9n, hmftfrihy bs3:Jj;jbfn h^juru &sh30m 3nhm&3&nb

*bj9 rij9je 36a'bj
5 i>h33nb jjcnSi^hn (bo6iOio3, li 3033hivi)3hb, 3i 3hob 36ob6m6h) 9b 6nigij,nb
(Byicn, 3n63h>c3hn 60301031136360) gy^mhois 5^3036001 3 ^Q6 ihnb 803(1 C 0 2-nb
53ai3nb36i 03ndc36i mhogib 3(101-3(1010 cgmh3 oai g,;>6bni(1(51130936 [2 0 ].
6 i 3D*1 36 n b 8 (139 3 6 0 ib ib o io i3 6 3 6 9 u 3 6 sh o b b i S i b ^ b m j>sh30m q j i j-
(im hoii ^ 3 0 1^ ^ 0 3 9 3 6 ^ 3 , hm 03oou> n 6 ^ 36 b n q jn tp i5 n i 6 ^ i3 D X (b p 3h :> 0 p 3 'b h g n b
ajm ^ m b o 6 ai3'b n b n 6 ^ 3 5 b n 3 m 6 ib ; 6533 369^ xPn-
- 99 -
^0^0 WOto'DCnO
6o36mb, hm8 gn'b^nb bbj>8:>mg cgbhavn q)ih&03&8fl i(j€n 36x n

6ob ^3801^839365 8530633 063331* gihp3333&b. <b38moioij830ob biîjcn^bm
35550001b 6>h9m^0536b CO2-001 95 bo65oi0ooi 5533(136^ 8(139360 (nb. 65b. 2.13
95 2.28). &>53h3&nb 8(139360 bb9350365b 53x193336 953595060101 5 8 5 013 00
E
OJ
o
E
=1
o
o
Photon flux density Photon flux density (PhAR)

[(imol-m^'S'1] [jimol-nr^s'1]
6 ib . 2.28 b n 6io icn b i g i Rhgnpnb q)mooc3&nb C O j-ii'b o ii (3300b Q idm y^jbyyybi bnS±a\y]hy 9 ih -
ljb ;)6 i bjhicnn: Fagus sylvaticah 3ih jn b Sjhnqj^hnxn (bn6icn0nb) g i 3 n g i (fihgnonb) gmcnQ^n.
Rhgnpnb cjmcnQjbnb biim 8336bi(jnm Sjh^nQn ( Ic) g i 3nbn 33bi&:>3nbn b36ar>J3nb n6&)6bn3m6:>
3<3hm g i6 i(V ii, 309(13 bnSxnpnb qjmo»03&nbi (nb. gni{,hs3 nb j33es 6s6nc<i). 9ia m bn6xncncn
6i IDhD5>H5 ( Is) J>i6iat3&nb jqjhm g^dbQ 06^36^3(^61133 bg36i (nb. S ih u b ^ gn i^hid ob b 3 g i 6 i6 -
ncn). ligo&n fi:>05h 3& x n i b ^ S g ih ^ x 3nhm&369n (30°C; 30 Pa C O 2) (R etter, 1965). 8 ^ 3 3 3 6 ^
b^hicnn: jjhm S njjxn 63n9n*6n ,jynb fihgnonb i 9^ i 6 n b3 9(535ihnb Argyrodendron -nb 3ohrra33h n x ,n
(L, 163 bn6ioi0nb) g i 3ngi (Sh, 563 RhpnQob) qjmcnQj&n. 6^301^303336(1 R i^ h 363(^0 CO^-ncn
5i33h36nb 3nhm&363n (Pearcy, cit. Anderson & Osmond. 1987 3nb3g3noi). qjm^mb36cnJ3nb
bn6xnc3'b3 g i 9 m ing 3& x 3&* nb. C h m o ra (1993)
1539 59 0 5 5 ( 13 8 ™ q jijo n h n b Sb b ^ n d ^ p o h ^ b ^ c n 5 5 3 9 3 6 5 bbnSnQ bô^hn 3h m i53b 3& n b

bnRjih3'b3.
5i33h3&ob <b(p3ihn bîjncgn^hn bnpng;p gb 331^9365 8(3365(13015 j(333nb
Qbbbboboi^bpig 5 b3 mg53b. hm^mh^ 63bn, 5533(136^ (bc'3bho 0^(19365
bn6oi3(bob ihm ^bnb 5bbjon33^bbCTu>6 3hcnbg. b 3 8 5 ^ 9 0 5595^5(1636390 gmlb 36nb
3bhymgooio 5539360b 83935*9 6i 30(10&nb 8(13985 831^9365 8osp6omb 8ijbo839b,
hob 838935 cnb6gbcnb6 Qg^hbgo 9^33001365 n6y35b. 3b 330dQ36b 053o6bbmoi 95-
£>590 &b6bcn36nb (ob. 6bb. 2.30) gb 6bhi333QO 3 o6 n8 b03ho 6030103(1363606
OrvjbofQho 2,m6i336l!)hbi5n36ob 81336^3^3 3 mj33936ob 838016333580.
m3on333ob 9 h30o 5b3m bb^ 3b 55(13801 cgb^mhcnb 3y>93h3b 86 o836 x (T’&3&<b 3
- 100 -
&68D16 03^6 8133666330
(J3D‘ 9^ ‘b3embp6‘b]) 3(536ihnb 35hd6m630m6b6. m^^nQic^hn 36606 qjihftco&n

6ih0mbQj,36b bib^mbnb Qimcrtn'bnmQrn^n^hn 3mj6n0m6nb, U30bg 3ohm-
6360^.6 9 nbn 83533606 bicbm9 b. b3CnCD^)i^ qjmlj|mbo6ai3‘bn di0nb6
gb3bbbbobcn3&xo (TOli)n3bp3hn 0(139001 3bb3bm&6 (nb. Sib. 2 .37 ). pnj3536bi gb
q3moiQm3i6 ^363680 C 0 2-nb 3 oib6oij3 n6 rn^^nQbc^hn 3h3gn 6bhi33x^Q 9b^3-
3 nh36x ni 0^93606 6yb0333u33pm6:>6ou>6 (nb. 6b6. 2 .4 5 ).
q)n^yTibn5 oi3'b3hn biyi3m6o6 0n3 r\yih36b ^hjdm qjij^roh^bnoi

db0nb6 n93nboinb, hm3 3han0hmx^0 !J330i 5 bh33 m cgij^mhn 9^93^01^9
81^03936936 C 0 2-nb 33<Ji3o636b'b3; 6333936^3, hm030n03 qjij^mhnb n6li)366n3m-
6b nb30ini, hm3 bq)3hb366 b3 5hmi536b. 013 b303300030n 5bh30m 3ohm636nb 3 mj-
039365 bb50mi,03b, 830130636300 6 b63 nh6 bgn6 hbmg36m6b 306033630019 0ynh93-
6b. 663b bbj03b, 013 5bh30m 3nhm636n 9(j36bhnbbcn3nb 6^(136300 6936b - 3 b3 n6
3 bcnn 'b30mJ03936ncn 3y36bh33n 5bi5n036ooi 63 hom'bxn Hbhymqjncnn d3h3&o 30-
CTibh93&b. gm^mbn6ai3cbnb ghm36ncn pn3n^nh36b 33ndQ3&b 5 b63nhm636 x n nymb,
Ob^bQncnbg, bb53 m])03 35yQm6ncn, ^ 3 b5 , Snhnjncn, 15639 b0 n6 g3 n 6 bb3 nh6 bgn6
330i3n636b 3536b bhbb30bbyh30O 3nhm&36n6 0mj03036nb 6b65hdoo3m6n6 3 hmh-
3mhi5n x ^ 9 - 013 3nhm636n db0nb6 S^b^hnb b6 3 bcnn 3mj03936b dbpnbS 6b65hd0O-
30, 3 b3 n6 bb3cnn (b33mJ33g36nb 336y33d)n6 838935 (013 36 6b3hoim9 8 36 ^ 0 3-
6390 nj63&b) qgml5m6o6ai3'bo6 h3bj^n3bi5nn6b<n3n6 8|536bh36 gngn ghm 6^nhg36b.
b0 0g5m0bh3m6b6 6ljh36n6 3309360iTiJ03936b6 360193636.
cgm^Pibn6oi3'b3hn ^ ^ 30 1 6 0 6 bgb3ly>)5nb
0(536bhnb 6363^ni3hbg q}ojboh3&x<i jl533^ 0 mn|5b3b bhbSbh^m 5bh30m3 n 3 m36-
gbh i53QO03636'b3 3 n6 333bpm h3bj(5nbb, bhb039 3i536bhnb 33bbdo36pm6366 833-
53m6 3n6cn3nb R3330 3nhm63&6 gb b5h3cn33 6 ^ 3 6 6 . hb(5 3q)hm gngnb 3i536bhob
bgb3(5bi5nn6 36ihn, 3 ncn 3qjhm Sbpbgnb cgmlj)m6n6cn3'bn6 5b3m6b3b0n gb 3 nai 30}-
hm 6b65hd0n3n ghmn6 5 b63 b30m6 b3 n 033d0nb 3i536bh36 33n6bhfi36m6 n6 33l530n0
5bh30m 3nhm&363n. bgbS^b^nb 3u36bli33n 3n3gn6bh3 3hmi536oib m3lsn3 o‘6 bunb6b
gb 3bh0m6n'bb|5nb3n 3g5m3bh3m66; hm5mhi5 3hcnn, n63 03mh3 hn5n 335336ncnn
6b6nbcnn6 „6b3bh33336o6“ 5b3m333b336nai 3nnp633b. b0b6cnb6b33 835336b 03m-
li)m6n6cn3'63hn bj,jn3m6nb 30bp03b gm630g3 by3b6b8n y i bh 9g 5m3 bh3m66 , bhb-
039 5bh],33x im 03hm3 n63 n 3m536b6b gb hn6^6 3 mhn6 . b3 nb 0b5h0naibg 03nd-
036b gb3bbbb30moi Rb3i3^n 3311303606 03i!P0 05hd6m6nbh3 bb3b636m h3bji50b -
3b 33h3Q3b(i n630i6bnhbg 3533&nb6 CO2/H2O 33qjbhg36nb 83i530bb, hm3 3i536bh3
„bh(5 3 n38n0m6b gb bh^ 6y3h3n0noi i3936b“ .
2 .2 .5 .1 b f^ O lC flb 6 ^ 3 C 0 6 i a jm ib F lb n e o m rtiD
qjfn^)mbn6ai3'bnb bo6bCJi0nb
013 cgmoipob 3n3h 3 oib6 oij330n 5b3m66n336b db0nb6 3(5nh3b, 6 3 6 0 1 ^ 6 3hm-
(53b8n 5b0mymcgnp3b C 0 2-3 b 33nd036b 5igbb^bh6m6 q3m^m6n5cn3'6n6 3hm(5363n
<3njbnh36xn C02-nb hbmg36m6bb. 36 5b3mn6blj36b C02-n6 5b0mymgncn. hmg3-
bb(5 qjm^mbn6cn3<bnb 33 g 35 bg 8 cnbn6 aij336 b n3 g 36n CO2, hb0g36n(5 5b3 mnymqjb
b36cnj3nbbb gb 5b“6au> I530n6 bp3mfi36b 33h 63h6g36b, b36m636, hm3 3np6330nb
6n6bOi0nb bbjjm3336bbi5nm 336^0.
- 101 -
66 ban6 5 6 ^ 0b 9 nb 96(f>3&6 9 9 6 6 ^ 0 603m0 afoD60b P 6 ff.9 nd 966
bn6>OTcnb b:>j>m3336b:>i5nm 3 ^6 ^ I c 6 ih 3m :> g6 36 b 5^ 0 m b b n 33& n b n3 8 6 1133 6 3 -

0m6ib, hfn330 -
h336n bi^nhm3b36 ^ghm 83^ bn6sox?3b jjm9335bi(jnnbôi3nb, 3ngh3 nbn6n, hm3 -
C3&ni5 6iiC3&bp n6^36bn3hbg b3601^^336. hmg3b:>u Ic n'bhg^b, 3nh3X bi 6353 n
i 66nCn ^ 3 ^ CO2 -nb 330i3nb3&nb Bhqjn3 5igng3bib, hiy d^h^n j^Rbbng^bnbi
g^ q)fn^mbn6ai3'bnb &:>3 mb3enb 3nhgb3nh3hn3mhyn30 m6 i‘b3 3 n30tnai3&b. i8 3^ 3-
b 3 bn6i>oipnb hîjunQbnb bnfijih3 y33C^ g*6:>hfi36nb 3^0080^(13630 qji^jmhig
Ijbhncn S.8 ajmcnpnb 6]^m-q)m,ynbn6an3'bnb gidmyigQ&XD*^ 1*1610103^3 bb3i ^yncm^n^hn qjij-

(jmhjbnb m3ljn639nb Snhm&j&Sn (3hi3icn i3lynhnb 3nb3P3ncn)
b iy i 8336bnsnm bo6ioiooai
atpSahoa):. 5*30511 &i6 >0 )363cyn6s ic &i53h3& i fs
(iflmon gm^mSn 3-® 63 ‘ ’
bOxsomb 9|ufuhx)bn
6ipiti<T'3i6fi 8)3361(13360
C4 0(j36ih 336n 20 - 50 > 1500
3gi66mb 81536^3360 - > 1500
C3 81536^3360 20 - 40 1000 - 1500
33enr>q3nl|)3&n 20 - 40 1000 - 1500
5 i‘biq3b3Cob 53(11030,1,360 10 - 20 300 - 1000
bisnmq3n^3&n 5 - 10 100 - 200 (400 )
8]hj6ni6n 8(1361113360
j|hR3ni,3Qn ^ynb b33&nb bn6icn(^nb qjmoiQ3&n 15 - 25 (400 ) 600 - 1500
^hmSnyjpn ^ynb b33&nb Bhgnpnb cgmcnp3&n 5 - 10 200 - 300
j)hm3n i x n iiyob ibiQjp.'bhgi b336n 2 - 5 50 - 150
qjmoiriQ0u336n qjihorimqjnicnp(n3i6n b33&nbi
pi 63^^36nb bn6scncnb qjmcncj&n 20 - 50 (TOO) 600 - > 1000

q]rriotmc0i5336n qjihonmqjmaxvn31611 b33&nbi
g i &3Rj36nb Rhpnpnb qjmon036n 10 - 15 200 - 500
ihi»5hm3ni,30n 3 ihig 363i 63 qjihoimo3Pioipm3i6n
b336nb bn6ioienb qjma1CD^n 10 - 30 600 - 1000
ihi^hm3ni 3Qn 3 ih19663^63 qjihoinKgmoncpi3i6n
b33&nb fihgOQnb cgmaic3&n 2 - 10 100 - 300
6063^13160 b33&nb bo6ioipnb 6063360 30 - 40 800 - 110 0
6n63R3b6n b33&nb Khgnpnb 606336(1 2 - 10 150 - 200
530811360, ^36360 g i (yiijBjftn
3 bdi6n 3^6nî>36nb 5308(1360 - 50 400 - 600 (8 00 )
Rhgnpni6n 3i6nlji^36ob 530811360 1 - 5 50 - 150
bi3b36n 5 - 20 150 - 300
0(ij3636n 50 - 100 300 - 600
6yc<ib 8(536^13360 g i 6yie 3 (536ih 33 &o
30s6 i 5m63h(i 6yiQ3(536ih 336n - 200 - 500
bi93ipfn bnphSnb 'bp^jnb 6yi(xJi536ih336o 5 - 8 200 - 500
gngn bnphSnb <bc'3nb 6yic3u36ih336o 2 150 - 400
6yhc8n Ridnh3pn £jh^0Pi3i6o 3u36ih 33&n 8 - 20 (30 ) (60 ) 100 - 200 (4 0 0 )
- 102 -
(W8016 03^6 800666330
o
o
o
e
Trues
a
z Sciopnytes
200 400 600 800 1000 1200 1400 1600 1800

Photosynthetic photon flux density [nmolTn’2,s"1]
6ib. 189 bb3J@ibb3i oj)5junm6icohn 363q)nb 815361*13011 63e)(TH3mj)mbn6oi3'bnb bn6ioionb 8*1393-

6n. 6s9pii3C03D6n fiiîh^bxm m3lyi9i03h ^^hivKjhi^ 91 CCb-nb 63636^3 im6tJ36(ihn5ai'b3
(9hs3spn i^mhnb 9nb353(101)
100 - Chlamydomonas
___ __ ___Dinoflagellates
iv , / \ %
o 50 - :> /
E i ( ' ' \ Diatoms
I
Chlorophyceae '
*-PhAR
.S i a
03 0"
100200 400 600 800 1000
Photosynthetic photon flux density
o |jj.molm'2-s'1]
B
JZ
CL -50 J
6bb. 230 ^n3fi3hn 6*13936(1, hm9(v)&no ib ib i3 36 3ps6Jy>i63hft 69*o9&3&>f036 rtb

gm^mUi6oi3'b'b3 UnBiCnpnb 5130361b. 863163 6yipdi}36sh30n (Chlamidomonas reinhardii)
bn6cn3^5fib 910*153631 (169361 19*13; pn i^ds& n b qjmlynbn6ot3<bnboi3nb bn6ioifyib m3lyi939n qjihcnî;
pnBmqjpi5xi(b369n qjr\ynbn6o!3<bnb m3lyi838n 6^x^6331 bn6ioi0nb 3Qhci 3ip ic (16^36^3(061^3
(N eale, 1987; O tt, 1988)
- 103 -
6 6 b 3 0 fn 6 ^ 0 b 8 n b 9 6 (f,0 6 6 9 3 6 ^ 0 603010 aftD& 0 U P 6 A 8 D d 0 6 6
533300636*. j3>6t!)3hn 5*8^166033606 5*90936* ( $ a) 5*6*3ohm&366 b:>3*b3bm

9h3gnb 83330111 * 6 3 3 ^ 3hm3mhi5n30 m6 n6 36*680 („q)mlj)mbo6oi3'bo6 3q)3j(j3hm6}“
<J>i 5*3mnb*»536* 801*601^8300 CO2 -nb 801036001 8mb30o cgm,jm636ob 1 0fnpt>3).
bo6*cn0ob 8(13906 3j)33cnhn *633* 5*630(101636300* bn6*oi0nb ^3*6^3606 iph^n
830130636001. d*0o*6 9*c>*0 5*6*0136*^3 gm^mbn5o»3'bnb 5*0mb*3*0O 0 ^ 9 36 *
306083630^9 *6 bh3036ooi *h 0153036*. *3 91*163^3 qjmlt)mbn6cn3‘b;)hn 3hm(53bn
bo6*oi0ooi 5*33(136300* (Is). C02-nb 3 oi*6 oij8 ob bofij*h3 *3 ob 338935 3333 *c*h
o‘bc'3036* qjm^mjoQn^hn 3(^(53636001 - ob 0n9n(joh936* <33h836(!>30o 3hmu3b36ooi
g* C02-ob 8*11*5001. Ic g* Is *b*b*3b 5*6*oi36nb 3ohm&36b 81536*11301* &3636hn3
* 96 n0b* 0 ymqj303 n g* 0*8*b*bo*oi3630o* 0(536*(i3Oi* 5*6663*336300 l4n336nb*cr>-
30b (tfbh. 2.8).
bb3*9*bb3* ô3ob 01536*11301* bo6*oi0nb 8(1393606 8 3 9 ^ 3 6 * (6*b. 2 .2 9 )
I5b*gymcgb, hri8 C4 3(536:1(13360 *8 86(103 qjhn*9 b*o6&)h3bm*: *63010 8(536^33-
60 (8*5., pm8o, bn3 o6 go) d*0o*6 3*c»S0o j>*6*ai36ob 3ohm6363n(5 30 33b *(p-
63336 bo6*oi0ooi 5*33h36o6 8g5m0*h3m6*b g* 3bm0mg <bm8o3ho 5*6*013606
(jbhncn 8.9 6i 6:>cn363QrTi6 :>bai56 93&33&X <3 'T'<n03^ 3 mhnb jp 6663:13365 (0h:>3;>Qn
i3^rTthnb 3nb3g3noi) (+ ôjhm Sipipn, dpnjhn, 3 3^11; - jqjhm pa6s(vi, Irjbjn, 6sjoo&n; X sh ihnb
&s9m bsoXn shbj&noin bb3im6s)
9 ibil>n 5013630° bn6ônb

ojmcnmcn ! ! apnenmQft
gmoncnb 6 A ib lM ] 6x n
qjmoipnb b53(jnqjn3hn qjshom (tjgiSnhnb qjbhoim&n/Sahipn 6m6 s) - +
3 nps “bxiSnhnb qjshom (33'bmqjn0nb 35h3ea6nb lb3Q>3nhnb
qjihcnn/qjmcnj-inb qjshoin) + -
93'bmqjnonb bnbjj + -
Sipnbig^hn 3ih 35jn6 nb bnbj] + -
6ij)336nb bnb3nh3 + -
Jcmhm3Qib(j36 nb h5mp36m6i qjmcnpnb qjihoft^ + ■
Jpmhm3psl^j36nb Jbtabnbogbx11
IcmhmScil^yib ‘brads - +
b^hm3s / (yiQsymenb 3m(j3Qm&s + -
^nQiimngjbnb hsrnp35m6i jhsffb3 ■ +
Jomhrocgnpnb 9Q3i}3xra6s jQmhm3Qsb|y3n - +
Jpmhmcgnen s / 6 + -
P700 / Jomhmqgncn X X
P680 / jQmhmqjncn + -
CF1 / Jpmhmqjnpn + ■
a ishm^nbn / b >(5hmly>6n - +
0>J3n6n / 3nmQijb:>6^n6n + 55oi3hsJbs5 ljn5n + lb3yjbs6|yi6n - +
XOijhmBjbnb ^hb5b 3mh^nb shbn + -
qO&ftyghn asbsbnscgbx0
qjm^mbnb^33nb sfonsmds + -
3C3Jo>hrr'636nb ^hs5b 3mh^nb bnftjsfy + -

ATP-nb bn6cr>3bnb s^jn3m6s / JprnhrKgnfxi + -
RuBP ^shbmjbncs'bnb 1^ 3 ^ 6 5 + -
J3s6^]hn 5 sSknb*3scn X X
5sh6mjbnQnh36nb Sshyi J03e 3&> + -
gm1ynbn6on3'bnb 36shn + -
9n^mjm6ghnx>n b]6oiJ3s + -
- 104 -
<W80)6 03 * % 8(30666330
Ubhnco 8.10 6no#v>bs g> b^hmb bn6xnenb* e* Shgncnb qjmoric36nb g£>&nsarjbs (Lichtenthaler et
al., 1981; Hoflacher & Bauer, 1982)
9>bibniono6x n Fagus sylvatica Hedera helix
ftipncnb br&xn(yM bn6x»enl> fihgnpnb bftSKnfyib/

Rhen(>nb Rhencfk
qjmcncnb bjgiinhn (bS8) 26.8 48.9 0.6

ojmoicnb bnbj] (:9) 185 93 2 409 22 11.85
qjmoxyib b3x n9Jn3^n ojshmn
(ea26h-, aah. 6m6>) 0.97 2.6 0.37
6^5336nb bnbSnhj (hime36mbi SS2- ^ ) 214 144 1.5
6ij,nb J,i8îhni6m6i (b9 68"1 ) 0.65 0.33 2
jprnhmSpib^jdnb himgT6m6i ojRcnonb 5.09 2.45 2.4
qjihoi'b^ (109 lv5i g3 "®)
JcmhmSpib^ftnb hvng36m6i qjmoi0nb 1.24 1.11 1.1
8r n jx f,'* i '*33 (lO9 h-6j g9"^)
jcmhmgjncnb (a+b) yn6u36)jhiisn:« 1.6 1.9 1.2
(9j, / qjfnompn)
jcmhmqjncnb (a+b) im6(j35(!)Kiijni 8.7 5.5 1.6
0 6 e 9-2 )
Jcmhmgncn a / b 3.9 3.9 1.3 3.3 2.8 1.2
RuBP iphbmjbnps'bnb iiyigm bj
(p, 9mon C0 2 a ^ 1 ) 398 202 2
6jjrn-q3fnj)fnbn6a)3'bftb n5^36bn3m6 s
(n 9mon C 0 2 a-2^ 1 ) 3.5 1.3 2.7 22.3 9.4 2.4
bn6icnpnb biim9336bs(5nm 63h^npn
(30) 9-S) 2.5 1 2.5
bnBiCxyKn 5^331136;. (3^ 9"®) 85 44 1.9
bn6>ocncn &ij 3h3&i
(|i ampn <gm^m6n a’^Ba-1) 600 250 2.4
9nj)fnjm5ehnxn b36cij3s
Oa c o 2 a -% ’ ) 0.5 0.16 3.1
3nhm6o50n ibo0oonh3636 C3 0^36^336^3 116^3660^^9. bi 6o6ii(? 0g 3&fn b i 3i-

b3bm hîjijdbb ^03(^^363636 bn6ioipnb 0n0bhoi 0i(ni005hd6rri6nih3 q}m^mbn6-
cn3'bjhn iiihi^nb 3jm63 6(536^3360 - 3bcnn bn6iOiQnb 0h3gn h ig o ijjo x fn^n-
030b :>03p:>3636b (6ib. 2.30). i 0 363q)b 0(13^3013636^ g ^ h x 0 »*»yr«L> jy33iiynb
g i Byonb 0(536^3360, ^(130133 bb3ipibb3i b3mhm3s6n 01536^3.
ibnOncM yii 9 b n i6 g> piRhoopnp Ib 6 n ^ j3 6 3 n

giRhgnox 3nhro6360 n 03 &33& X 9b36ih 30 i5 qjmoi036o b36otjs336 6 ij)03 &of>-
^36bnjhig, 3ngh3 0boi6, 6^0130 595003680 0m‘bihg 01536^300 ojmoi03&n g i
50oj,m0 yn9336biynnb 0g5m0ih3rn6ib obo6o 2^50036001 65^0360 o6J|(36bn3ri6ob
bo6boio3(b3 ^["63336. bi3hovng, fihgnQ0n 0m'bihgo 6ôibm3636ob g i gi&3hx<1
jjynb Ik Qiib0m36noi 5 0 moo q]m^m6n • 0 "e • 63"1 3ghob. &ihgi i 0 obi, 36Qi
ipo 6 o96 fnb 55(130133, hm0 Rhgooob q)moic36o g i6io &i6ion3&ib 33301 oy363636,
3ogh3 bnBboiQnb qjmoi036o g i diQOi6 0iO3 5(^63336 bnBioxyioi 5533(136 ^ 63I1-
vyiOb. 03nh3b 0bho3, 3x nfTK3 n(4D^,n' hm0Q36ou X 3W 1m® j;>il»^;>5b^fI'K>rri bob£)0ob
gngn 03bid03&c»,»636oorii g i j)ih6mjbn0i'bnb 0i(niQn ij<yi3m6noi biboxng36ni6.
- 105 -
6^ 9066^^013 anbaartow ^ aa^ô 6030)0 0 6 0 ^ 6^6anda6;>
Acer saccharum Prosopis farcta
6ib. 831 bn6ioiQnbi g i Rhgnpnb cjf'ncnp36nb 3nb1yncmj,n3hn b^hjj^hi. mh9bhn3 bn6 icnpnb
qjmcnmpb (163 bn6boi(Mibsg9n mhn33 9bhngi5 3jb 3m6nh3&3cn gjnKnmpn) ^.hîg jp d n b ^ x 0
bjjpn j,ih3 {>ihbnb SJmBj DSngghSnbn (EP), 3hi3i£ 9hni6n9 hn:>6fl g i 3hai9i63cncns6 9^nghmg
0njhnQn 6i5hd3Q3&x33h3g36ni6n S^bhnb^bhn 3ih 36jn0i (PP) g i ph36pnb36hn 3ih 36jn3nb (SP)
bj3Qn jbm3 ri0 n Cb3gi 8^ 3336^ b]hicnn). dpn^h giRhgng3pn 3i 6 n ^ j 3&nb 0(536^3011 qjmonp36 n
5 iamnhfi33 rii6 cnb30n bnbjnon, 03bhob36hn 3di36jn0nb 0mj>C3 3jh 3 g3bnoi gi qji9ih p'h3&cnb3&h
3ih 35 jn3 i 3 n ishîg 5i0mbi^xn 3sh3g 3 mhnbn bn3h(j33&nan (fojgi 8 ^ 3 33 6 1 bjhsom) (Salisbury &
Ross, 1992). igiS^>(innb bS^nqjn^n bjhnbbn 6i 0 mnbij|3&:> "3o>bl>>ni3hm6nb n6g3jjncn‘ : bftSjcnQob
qjmoxyib bnbj3 9n63b Rhgnpnb ojnicnmpnb bnbft gi fcsymcjngn bo6*Ji(Y)b qjrocnenb bnbjoô
(Carpenter & Smith, 1981). ^hmSniX0 63090b ,jy336nb biqjihfojS amtihg g i dg03 h0 RhpncnbigSn
ig i 3,yih3&3p 9(j36 ih 30U> cgmonc3&b 3n3mg3h03gn jbrgncnb (Hy) J339 5163001^36300 ifocn
03'bmqjnpnb oibxn 3 h3 (M) gi gogn 3jh 3g3 mhobn bn3h(533&n (SC) (Lee, 1986)
35301 8mnb3bh36 06,536^3(1 bo63<j»93b 93 03010 031*1,50^06013133110 530^ 33390 15

53^936001 90905 (ubh. 2.9; ubh. 2.10).
3360,5300b 536301360b 3ohm636ob 0o03hari q}36m,5o33ho 3935,531503 bmhunQcg-
365 dnhncniQig a|mcn036ob cgmh0oh36nbi 95 0:>ain 930950100 goq33h36uo3unnb
3hmu3b9o, hmu3 939069365 ob 001(1030191*1503110 (63b. 2 .31), 30^501001503(10 (65b.
2.31), 0oj>hfnboh3Jo3h3cn 9s 6omjo0o3ho (ubh. 2.9) oi33ob363h3636o, hn0O36nu
556b3ohm63636 93639 93 05p 59 556501365^3 C0 2-ob U3Cn^> g303b3bo3oi36x 6n9-
636b. 536fTion^n 5^6b;>'b(p3hi3b >8 5953,55(503(10 3hmu3b3&ob 9o0oooh3&3b: 035.,
5390R(goo36obioi3nb 9303b3bo3oi36x o 0^(1031^191*1503(10 g i 036junm6393ho 6n03&-
oi3ob3&36nb W1390 550m0g(T'536365 03b5d93&xo5 0bmcor*g 03(j>39fl 536301360b 3n-
(10163600 g i Obmcmg buomqjol536b 3^301 363hn 556030015(1016 3J)O03h3b3g fihgoQob-
30,5360 (301019360 936390 536301360b 3ohm63&0n. 0138153, buomg)O036b 033090301
636o9m6ho3 03353016 bo6iOipob 0156950156^6001 0b3hg o6o36bo3m6ib g i b3i>-
03mg 03(t»39 536301363^3 0393(136001 631)9360 936535(150 3Jm6g30».
- 106 -
cW80 to 03 «=U 8 0 3 6 ^ 3 3 0
Time of day
fob. 23B COj-Sjitou (33Qnb g i3minp3&3C3&s 9bnb h ig m u n i^ . C4 0(536183360 q)mv!>mbn6-

cn3'bnboi3nb 3ijxi9ie38ig ny363&36 bn6scnc3b; 0som qjml!>nibo6oi3'bob gpvi^hn 0(1390 oinoiJ9nb 083-
mh36b bo6ioi0(ib 0h3gnb qjmh0ib gi, 5i6bb31336001 C3 8(536183011 <gmlynbn6oi3'bnbi6i6, ih bg]6i
bn6icr(xiai 51338361. C3 0(536183360 deroh 516101361b (513383606 63h^npob ^>33001) ih 01^363636.
1^001(30^360 (bn6iOTQnb 8(5083 h3b3hb36ou6 83633630(1 9^36183360) 10305x183636 83^ C02-b
6o6g9o (gognoi bg83, 16 53016 bi(ni8mb, 15830133 gpob 00 33homg363n, hm(}i obn6o ofihgno36oi6
bb3i 0(53618336001, 16 01369^5 03ig(pnbib dcfvjho c+i36pni6m6ob ghmb), 30gh3 330001030,5360
(bo6ioionb 9my3ih30O 6(536183360), 0138(51 b(5nmq}nlj3&b cfbni6 i8o6g0o ih 933dc(iioi 3300m-
<30^3603001 3<33ll3hig 5i3mny36m6 8ip>ico o6(j36bo3m6ob 516101361
b n Sxn cn b 5 5 3 9 3 6 5 qgmly ,^ nB ai3<bbn5cn3'bn 95569^136001 6hqgn3 956(11309363-
9365805 556iOi36xrT'65bai56 (5533(13606 336^5693) 013 55*6015 15395 5h rib-
(^39365 (15063 6635 cjbjiynhnoi (655., 6y9nb 0ih55n, 03933(15,33(15). C4 6(5365113015
93601633359(1 3b 6086536 065b, hm9 9b (156 595093680 9501 8331^90501 80*905659
559moy36m6 3359(3106 bo65(7i93 (65b. 2 .32; 95 2 .33 ). C3 8(53651133680 cgm^m-
bo6oi3b3hn 55hg5j9636o 5^03(19365, b5659 556501365 5h 805006336 Ig-6. (H136-
93680 55953590 ^650)93 386083639(^9 8mJ83936b 33oomqjn(j36ob cgmo(Tibo6-
oi3bob o6^36bo3ri65'b3, 303(1™ 909 5539365b 56936b 35(1559 558m650X n ojCDd-
O35150360 1539590 8mph369XfT'6ob ghmb. J 330!dn'j b(5nmgo^36o 95 6(190906
<3(0019360 b3 8(5365(13015 35(133606 8056001 5560(59056 558^(090 (535936590 06036-
bn3rn6ob 9bnb bbo336ob 5539365b. 9bob 501065010, 013 nb (158936083 6301b
5hd399365, 06333b <3Piom6o6oi3bo6 06g3j(503h 3hm(53b36b. 0bob bbo336ob
3m6ooi qjmoi0flb 556^(^930956 h58936o93 63010b 836935 qjri^nibo6oi3,bnb 83mh5-
90 (135^0360 653(101306 30(133959 (135^503680 65h8mj86090 NADPH2-065 95 ATP-
b 5338x0113606 3(11*1(53680 (C 0 2-ob 31^00938065(503(10 030^5(505) [ 185 ], 585-
31(159, iggmaigob 803b 33od93&5 559(^19363639 oj63b d903ho 556501360b d59n56
b569n393 03513515. 68oho ^yob ^3391 05h36n6 6(5365(133695, (1(089360(5 5(15-
oi56565ho 5565013606 3nhm&368o 0^936056 (55839836001 0(539365 606501906
6nb3oh3, 6(1909065 95 60650190b 9mho53m65) 83089365 q3m(jmbn6oi3bnb 3qjhm
- 107 -
6 ab 3 0 ( f > w m 99( W 0 60 aCD0 0 ^ 0 6 0 l» Q 6 9 n d 9 6 ,i
Time of day Time of day
fob. 2.33 6>jm-qjm(!>pibn6cn3'bnl> g} 9myi9363^36} ffbnb higftiijni'bj. bn3 n6gnb (C4) 6»m-cgrn-
(jmbn5oi3'bnb nB^Sbr^minb gprrjhn gn6} 9 n^} najfnhjdb brrfbcnpnb 8h3pnb 9003b qjmh9 *>b. Snqjpob
bn6ioicfib qjmcnQ36nb 53^m-qjRlynbn5cr>3'bn 809336} bn6xn0nb 9po3h U3sc3&>>gm6}t> 3bm0 mp 1000
n8mQn qjm^&n 8"s 69"'-nb 33b}6i9nb &}6}0)36}d@3 (Hesketh & Baker, 1967; SchuJze, 1970)
Irradiance [kW m Photon dux density [pmol rn2 s"'J
6}b. 2 3 4 a , b U iQ iO X 0 qjmoncnbi g i 9onpni6n 81536^3300 b io jih n b qjmljmbn6or>3'bnb 9 ^ 6 ^ ^ 1 9 3 6 3 0

6 } bnBienpQ'bj. a bmh&pnb gn cnpribi g i 6soi3bnb (LAI 3 .2 ) qjrry}mbn6ov)'bnb bnB iO ico'bj g a 0 r> y ig 3 -
6 x 3 ^ n b ao g ih ^ b i. 6iO>3bnb <3rryy'ibo6on3'bn LAI-toj, bmpm qyncipob a - 3 nb
q jih o vn ji'b ] (Evans, 1973). b ym33p9bhn3 {,} 6soi36x n 6063116, in o ^ o h ac^ h iG bsSsarjbyyt iVîiO"
6nbs d>ynb j^ m b o b <3mljmbn6on3'bnb 6063116 qjmlyribn6cn3'bn J>ii6& ihn 33& 3[yii 9 nb
<3 >hanft53, ^rn^nbi - 6063360b b ^ o n m q p h o n m i^ , bmpm aynb} - 6nag}&nb 1b 3g i 3nhnb qjshcnm d'bj.
ICN = 606336nb qjfryj,mbn6oi3<bnb bn6:>oicnoi 6 *33^ ^ ; = iiy°^ q)fTi,jmbn6cr>3'bnb bn5 i<nc>non
6^50*1361 (Jarvis & Leverenz, 1983).
- 108 -
&68016 03^6 800666030
6ib. e.35 tiq jb xn b 93hio 33hnng9 n CC^&s'boui 1530 Quercus coccifera-b 3ihj3n 3DhiyiiiC3^n
5hign35^nb Snbjpgnai. 1 bn6iOi(Vib ajmcnpodn 3shjnb bjQi. 6i 6 op9 n; 3 3ihjnb t>0Qi 6b6npnb 9jbi93
3hnb bn6ion(yib qjmoipj&n; 5 Rhgnpnb ajmancjim phjnb 9 ngi 6i 6nc9 n; 7 Bhgncnb qjmoiQ36n 3ih-
jnb j33gi 6»6np3 n, hm9pnb ^3300^^5 LAI 3.5-nb ^mpni (Tenhunen et al., 1989). qjn\ynbn6on3bnb
g i 0 minp36xD 6i qjmoicnb nBiOfCiOr'i'ba nb. Kuroiwa ( 1969)
05 (piQ j,i3m bi3i0b 3 ni(p6 nm6 , 3ngh] 3m bi0mg630ni 08^33 himg36m6nb

bn6 iC7iQnb 5^3(^3636001 oii6i&iho 5i6:>ai36nbib.
qjm(ynbn6ai3'bob bn6ioi0nb f^JyiQ&n 6(536^30^ oii6ibi'bni2>i9Pi363&9o

b^CiDDC qjrncnmo'bQ 9i|)3oh336i3 83(^036^ a n6 3 ny3i6mb 9ibj>36i0g3, hm9
cnnorjmb 3 boi 6 gp3b bo6ioi0ob boîh63i. 9;>5hi3 3b ib 3 ih ihnb. gpob J>i63 i 3-
0 m6 i 9 n 15^05330 <3 rrioi0 i 3 g 3 bo6 ioi 03 b]0 b b 3ig ib b 3i 33016001 b[p633b - 563 hm0,
cgmCTi03&n n33fiiOiig ihoi6 3 ai0 Oi6 ig 5^6^013630*1. 6ihg;> i 3 nbi, 0301010360 Rhgo-
0^336 3hcn0i63cnb. 3b J9 6 nb 5i6ioi36ob (330ig 3nhrr»&36b, hijj 3(j36ih3b 83bid-
0360m6ib idg33b 5i3piny36mb {>i6iai3&ob y330i'b 3 3 ip;>on 63636(1(13(1 06^36-
bn3m6i (6ib. 2 .34 ).
ibynb i336m'bnb bb3igibb3i nih3bnb 03(110103611 J>i6bb3i3g36i q3m^mbn6oi3‘bnb
bi3hoim 6s0nbi30noi. dohooiigig b6 mh3 g 5 pi3 q3moi03 &oi6 6igibri3i6oii oii6i-
bi'bm5igm36i3n q3Pi^mbn6oi3'bn 5i6bij)30ih36ooi fi6(j36b(i3hni 3 î lbm6 i 3 n, bigitf
3cnbn6crrj03&h v|i3g3 3ni(T>6330o bn6icri0nb 3gnp3bn 6i6o0o. ^336811, ig,h30i33
b3crii b3 nh, 3b3q3r'ioi0oi6 3^ 336 8 0 I536^hng*6 gi3mh3630 01630 03363630 qjm^m-
bn6oi3<bnb bn6ioi0noi 5^331136:) Sbmgrrig digoi6 o03nioiig 013 3no(j>633i. 3ih3ob
8(156001 gi li)mli)36nb y33Qilb3 Sipign 0ih3bnb j338 (bigijj 5^663300, 90033^3(10
5i3mbbn336i 3q|hni i(p033b, 30gh3 3bob 3ohgi3nho bbo336n) qjri^mboBo^^ho
5i8mbi3i0O 3(3ohg36i bo6ioigob o6^36bo3n6ob oii6gioii6m&ooio 830ijnh3&ob
3hm3mhj3(i30i 9 . 3ih3ob 805600^ fihgogob i8(ji6o cgmoi036n oii8i3m636 86033-
630fn3i6 hm0b 3j336ihobioi3ob i 3 ihibibih6o3gm bo(j3i(300b gidg33i3o, higgi6
nbo6n 5^0936001 33301, 30gh3 ihiigi3ôh3630n q3moi036o, 3mnb3ih36 b3b^
bn6ioi03b. 3b ihob q3m^m3m03mbî^ni3ho 3033^0, hm03obiu ij3b 6303311300 36 o-
3363gm6i bo6ioigob o6(i>36bo3rn6i3o ihb3&xn gogo 3m6v|,hibli)36ob 83h6og36ob
- 109 -
60ba 0 6 6 , ^ 0U 9 n b 9i(n 066 9 9 6 6 ^ 0 6030)0060601 &6 6 9 P)d966
oi35g 65<bhn6 noi. 30650956 6(190906 <3^319360 559563(13606, 38 hico 353(1065 g i ji-
hnb 6ij)036 5530365b 5560(39056, 1530^9 b9 n6g3 n 0501 650039m 639090 3351)301
dohocnign 363(i53l|)02,xn ©5659001 63015 0m05h5536n6 bijcl^n.
ogryyO(>d6jlyn63hn 3m3xH Sn3^ > qjrvy’ibnBoig'bo

Bycnb <336580 q]Oljm3056;)orTi6n 656099365 bo65oi0ob5 95 6o3^3ho3nb 33(1-
^,oj)503hn 5(159036006 306393001. <3mombo6oi3'bo6 6 o(ph3 n63x n 3h3gn 83365-
658365 qjml5mbn6oi3'bob 60650100b 8(1396 (ob. 65b. 2 .30 ). 69500(3365(133606 336-
016333580 3b 00535608636b m3(jn 939b 3, hm00ob 1)333001 35jbn0503ho qjm^mbn5 -
oi3'bnb5CTi3nb 60650193 5h5b 5 t,05hnbo5 95 3656b, Byoob '639530(10156 5bgmb, 6595(3
3bnb 05(ji05 g^niOobib 009360 bo65ai035, hm3 3b <31*1^1*1062060(13656 06333b.
50n^rn0 <3i*iom6n6ai3'bn6 851)608593110 06^366031*165 n‘bi*i3365 5h5 38359^19 6yon6
Ib3053ohob J330, 5h503g h509365903 3qjhm (ph05g ( 6 n(j+i3 o6 5(163306 bnb6mh3
9581*1^09363005 606501906 06^366031*16565 95 Bycnb 6o8p3hn33lb 3). 3m r(i36c3c
9^33665 95 95 33g5h36ocn 956590 hsgosuoob 63'6m6368o, (11*19365(5 606501906
060366030165 m3ô0306 5^5(16366, 05j6o0503ho cgpn^m6n6ai3'bo Bycn6 '63-
g53nhai56 56cm 81133680 5(*vihni56365. 6o(ph3 o6 85036560156 3hoi5g <31*1^1*1606-
oi3<b3hn 5joo3m65 3133^ 0^5893, 65658 06 ^033660(19365 63601J3001. ^ 3 3 3 6 6 5 -
(3056 6 3 3 x 3 ^ 0 3 595090 5j36 n3 bn(T>h03‘b3, 6595(3 5^6336 '639530(1390 (15905(30-
06 ih i 363036 1%-bi (b5j)f*i3336bii5nm bn(ph03).
2 .2 .5 .2 . viD aÔ h^hflb 6 ^3 CD6 i bD601j 3 iW Q) q 3fTlv!)lTlbn6 CnJb'bD
830561*193(1 3hm(53b36<b3 03933h503hob 5539365 5563nhm63630O5 Jo3o3h jph-

g5j3650i5 ^0630035^3 g i 50 55(195^863680 3m656og3 <33(18360015 5 ^ 3 ^ 6 5 ^ 3 3
3 nbo <b30mj03936ooi. 3560 Smognb ^561*1606 oii65b3ig, (135:113006 606^5(13 k 0^(193-
65 O3053hi^)3hn6 3j63m636(5O503h50. 0 38 ^ 50 3^ 0 6 1 0°-noi 81*1850360656 h3ij(3rt-
ob bnRjihob 5i'bhg5 55 8 ^ 6 5 0 3 6 5 033^3(15,1)3(1390 ^1*130^(3036^001 Q10:
In Q j^ lO /C T rT ^ ln ^ /k !) (2 . 7 )
6595(3, T2-T[ - j)3033hil!)3h x f> 06,53(135905, K 2-Kj - (135;)(3006 336i6i3nbn bnfi-

jih336o. 5ihj)3330 3(3oh3 q35h50363n ^3033(15,53(106 83(3390656 ,53833(1503^30"
yn3qjn(3n36oo 6 ij)05mg 03g0O3 6ogog3g (16365: 1 .4 -2.0 - <33 h836 ,5X h35^ 0 50 15
33hi3036rn6n6ioi3o6 95 1.03 -1.3 - qjn'bojtfho 3(101(3363606501306. 3qjhm gng 035(15-
93680 oDS^hi^hob (339093606 03,5561*193(1 3hpi(53636'b3 553C3606 83665390656
3695 535661*13936, hm0 03056m03h 3hmu36oi5 Q10 0(339365 038^5,53(106 83(3390-
656. 95659 03333h503h36'b3 3b 5539365 gogoi, hig5i6, hm5mh(3 63bo, <33(1636-
0 X n h3ij(3n36n 606^3908000(136390 3hm(5363&05; 3qjhrn 05p 5O ,53033h5O3h:5'b3
050 3(3ohg365, (159556 qgnbn^hn 3hr*i1336360, hmftmhoij, 855., gn<33<bno6, 6nRjih3
69365 8590800^36390.
03833(1503(106 5530365 3nlynjm69hox 636cnj35b3

60663006 563 3 o^mjm6 ghnxn 63601J35 o'bhg36i O3333h503ho6 8503606
83656580659. 5°C-'b3 j333noi 63601^3580 0m656ng3 6635956635 3(11*1(336015 ijô-
- 110 -
366666 L 6016360
Gas Exchange in Plants 107
6ib. 2.36 Pedophyllum peltatum-rk

qyncncodnb 3n(ynjm6phn;x'n b]6cnj3i (bn6-
6xnb b]6onj3i) ‘bhgilxniB gi
(ioS^hi^^hnb JP3C36nb b>5{)hdcn3m6ibaai6
ii39nh3n (Semikhatova, 1974)
3iljnnb 36 3*1^ gngni gi iSn^mQ Qj0-ni3 3i£’i 0ni. 25-30°C-'b3 3qjhm 0ipiO ^38-
33hi(!)3h36<b3 b36cnj3nb ^ 3 0 3 3 ^ ,1 ^ x 0 yi3cgni3n36^n 0(j36ih3cni 33 hi3C3bm6 i 9 n
3U33i 1.5-0g3 gi ^qjhm gi&gi. ^0 33 3qjhm 9i(ni0 (j3033hi^3hi'b3 6nmjn3n3hn
h3ijisn3&n 3n3gn6ih3m6b nb3 bBhiqjig, hm3 33p>ih b3hbg36i 3hmu3bnb 3 ^ 3 6 -
330ymq3i b^ftb^hiijncni gi SQî&mrn^&ncn gi b]6orJ3nb n6^36bn3m&i 0unhg3&i
(6ib. 2.36). ^S^Qhi^h^ftn 50-6CrC 9mhnb i'bni63&b q)3h936^36bi gi 0336j-
UnfTi6 i 03hig 96 n93630 m3i 6 0306hi6x hibiu b36cnJ3nb 336 y 33^ i
6mbg33b.
5i 3035i qjriiynbnCioqftfbQ
q3m(jnbn6oi3(bb3 ^ a ^ h i^ h n b 5530365, dnhnoiigig, 5i 63nhm&3& x n;> DCDilr
hm6nb (ijhi6b3rihl5bi gi 03mh3x 3hm^3b36'b3 9nbn 0mj03g3&ncn. gi&i0, b3&m3-
^n0i0^h ^ 3 6 33 ^ ,^ 36 ^ 3 6ib9nh6ignb cgnjbiynnbi gi i(^g536nb bnfijih3 n<bhg3*
6i lj)3033hivi)3hnb 9 i(j36ibcni6 3hcnig m3^n0 i 03 hn 06 n93630 m6 nb 0 n(p633 i 3 g 3. m3-
ljn0i 03hn ^3033hi^3hnb lb33ncn C 0 2/0 2 33qjihg36i 0unhg3&i; 3b &i0mnbi,!)3&i
q33h036l5 hnb30m‘brn6nq9iTibq3il5 îh6mjGngibi/mJbnj)36i'bnb 6iiih6mjbn0nh36ign
ijijn3m6nb giU30i9n. di0ni6 0i(p:>pn ^3033hi^3h3<bn i0unh3&36 6ib9nh6ignb 03-
îbmpn'bQbi gi 6n3cnn3h3&iou> îBbimh^Sn 9m6i 6n03 bb3igibb3i h3ijun3&b
9mhnb 3hcnn3hcn>p39nhb. i 9nbi gi 0336hi63&oii6 gij>i39nh3&xr» cgmljmjn9n3hn
3hni(33b36nb n63n6nh36nb 93g3&ni cgml5mbn6cn3‘bnb b6hiqjn 936y33^>i.
diQni6 9i(nig lj)3033hiv!)3h36lb3 C02-nb 3 cni6 cnj0 i 3cn0ni6ig 6yg3&i; 013 ©153-
6ih3b i 9nb 930g36 gi3g&3&i b3pbiyh30n 3nhm&3&n g i 013 3 ib ih 0n3(j'ni b ^ 3 j-
- Ill -
G^baOfnb^Ôb 8n b 966356 8 9 6 6 ^ 0 6030)03(f>360U P 6 69 n d 866
Temperature
6ib. 237 C3 9u;)6sh33&9n <gmj)mbn6<Ji3<bnbi pi b^&Trjsnb e i3myip36x 0&i (bD ^^hi^hi^. jp 5bb-
3i33&i 6h3,im-qjmj)mbn5cn3'bbi g i bn6soiCD'b3 b;}6aij3ib Smhnb ihnb ejjfn-Gjm^mbnBonj'bn (j,hia)n-
inb gi9j(hnbxf’ 6i6ncn). bnubnb{>i6 &h3^m-ojrnlimbn6cn3<bnb gi^jnoij&ib ;)0cnb333i bn6icn03'b3
b]6cnj3nb 3 38^^136x5. Topt = 63^m^mly*ibn6or)'bnboi3nb ,j30>)hi&jhnb m^nS^Sn; T , ^ =
i^jhnb i)ihpn6b(v3hn 3n6noi3n (bfnjn3nb 3n6n3] 8 n; bn(jn3nb Qndn^n); Tmax = ^^QSjhi^^hnb îhgn-
6iQ3hn 3ijbn830n (bnjjbnb SiJbnS^Sn; bnybnb QnOn^n)
ii3^DCn g i‘bni63bi, qjm^mbnGo^^hn 0336 ;!^ 630-630^ ipgg^bi.

^ a ^ h i ^ h n b &i3Q06s BQ^m-qjm^mbnBcnQ'b'b^ 3QndQ36i 6i6nbi‘bc'3hmb b>9 n
3ih i 03 (i)hnoir 3b36ni: qjm^mbnBoi^'bnb bn^n3nb ’bp3bhn (^63 63(jn-cgm,ynbn6oi3-
"bnb (j 3033 hiii)3h30 n 9n6n030n), ^ © ^ h i ^ h ^ n m3^n930n 95 bnoi63hn 'b(p3ihn
(563 63^m-cgm^mbn6an3‘bnb ^ a ^ h i ^ l ^ n 0ijbn030n; 6ib. 2 .37 ).
m3 lin9 i 03 h j ^ ^ h i ^ h i o i i 3&i6n. ^ 3333hili)3h30 n 3h3gnb nb 3&i6n, b ig i^
63lijm-<3rnlt)mbn6cn3'bnb n6^36bn3m&i 3 yjbn3 i 03hnb 9 0 %-b iîh63&b, 3 nfi6330 ni
n 3(jn0 s 03 h 366^9. 6 36 36 ^ 3 3nhm63 &3 n C3 3(j36ih 3363 n 63^m<9m^mbrt6ai3‘bnb
l53033hi^ 3h30 n m3^n030n 3qjhm gi6iQnb, 3n9h3 3m^36 ^n3hn q)m^mbn5 cn333h3£>nb 3nhm6363n), hig6i6 (J3033hi^3-
hitb3 n‘bh@36i qjm^rr^Ba^nbi g i 3 n^mjm6 ghn30 n b36crvj3nb (ihi 963 i 63 jbrn3n-
03&3 n) n6^36bn3m636n, hi^ i0unh3&b qjpi(i,nbn6cn3'bnb 6i3mbi3i0b. 153033^^3-
h30n m3^n039n 9(j36ih3ou> b^yncgnjghn 3ibibnioi3630ni, 0 13 3 ^ nb 8306036^
93 nb3i 0 |T|b J>ih30m qjij^mhoii lb 36i3036nat; 8^5., bn6iot0nb gi6ip n6^36bn3m63-
6^ 3 m3ljn3 i 03 h ^ S ^ h i ^ h i o i i 36^60 3qjhm qjihoimi g i nb 03^011*13936^ (538-
^ h i^ h n b 330 ynh36nb 3 n3 ihoi3036 noi. ib30i, ib3 301^3^01, „ 5 ihoib930M mU^nQâb
6ihj,3330n 33nhilj>3bm&i 6iiR6ni - <3CDAKPUn3&n b id i n g qjihcnm
bi‘bp3h363n n6333b cgm^mbn5cn3<bnb nB^&^m &nb 3bm0mg 306093630™ 13300-
03636b.
63^m-q3m^mbn6oi3‘bnb m3^n333 n C 3 0 U36 ih 3Oii 30hi 303 bm&i-
3 n 15 -30 °C 3 mhnb 03hy3m6b (ubh. 2 .1 1). y330i'b3 gi£»i0n m3^n933n 6iifi6ni yn-
3n h36nm636nb 0nj363&b (ihj^njjibi g i 6;>6k:>iD01h3&noi ^6 îhj^nii 9 n). ^3033]-^-
(J3h36n 10° g i 20 °C 3 mhnb m3^n3 i 03 hni b^nriq3n^36nb gi n0 3i536ih33&nbiav
- 112 -
cW80 tt 03 ^ 903656030
Ijbhnon fi.ll 63^Pv«jiTKbiTibo6ai3'bob gi8piyig36x3&i l i s S ^ h i ^ h i ^ C(>2-obi e i bfl6*npo0' 5133(13-

60b 3nhm6363n (a h ijifo i3<bmhob 9n b x 3ncn)
^536*3011 » ]g n ,5 3 3 3 ^ ) ^ 6 ^ aS o h itfjh n b ,53333h i£ £ in b

j^ g s fcp3ihn (<0,503330 (°C-> t>3gi “6(^3ihn (° C )
6xs>tx,i3i6o 8(5361(13360
lib x n 3s6n^j3&nb C4 8(5361113360 +5 - 1 0 3 0 -4 0 ( 50) 50 - 60
iX ib O h X " C3 8(336^3360 -2 - 0 20 - 30 ( 40 ) 40 - 50
33Cnmqj(\j36n -2 - 0 20 - 30 40 - 50
b(jnmqjnj)36n -2 - 0 10 - 20 40
3Qi66mb 8(5361(13360 -5 - 5 20 - 35 (45) 45 - 50 ( 60)
C A M 8(5361^3360 gjnobooi -2 - 0 ( 20) 30 - 40 45 - 50
C A M 9(5361(13360 (ns0non -2 - 0 10 - 15 ( 23) 25 - 30
5i'bK 3b30nb {,3(0050^360
g i 3hcn6p(Ti3636n -5 - -2 10 - 20 30 - 40
9i{n ic 9oinb 9^36^336(1 -6 - -2 15 - 25 38 - 42
6ypo6 ynhS m ojoô -0 ( 15)20 - 30 ( 35) 45 - 52
83(1^60160 8(5361(13360
l5hmUni)3pn g i b36^hm3nj)3on
91(119863163 qjihoiriq)fnori0(>i3i5n
b33&n 0 -5 2 5 -3 0 45 - 50
9i 6j,hnb 3(536^330^61 0 - 5 25 - 30 -4 0
99hipn (1350(1163606
bicohmqjncjhn 8(5361(13360 -5 - -1 20 - 35 42 - 45
bnQn3hn bih^yQob
qjmonmo9(5336n 63360 -3 - -1 20 - 25 40 - 45
91(119863163 6063(113160 63360 -5 - -3 10 - 25 35 - 42
15369(106 3331 63^360 ~ -3 15 - 25 40 - 45
6136360
ih iy iy ib i g»
b36bhjlyii3gn tbm6nb ~ -8 5 - 12 -3 0
lbm9o3ho bdi^ypnb ~ -5 10 - 2 0 30 - 40
00)363611
(5030 h35on<636ob -10 - -15 S - 15 ( 20) 25 - 30
3gi&6m36ob ~ -10 IB - 20 38 - 45
j|hm 3n336ob -2 - 0 -2 0 25 - 35
oim30ob3nhi — 5 0 -1 0 30
ar)3h9(nq9op3hn 20 - 30 45 - 55 65
30b, hm3036rn5 93*3016 igh] j^'biqjbjc'b] (hmg3b*u Sijhnb g*6*-

Qnb). 0]mh3b 9bhn3, 9 bn*6 n iQ&nco&nb 6*0^^36360 , *&h30>33 cn6nQn 3*3nb b3
0l5D6*h33&n 0*jbo0*c3h cgmombnBo^'b^h j)*0mb*3*pb od033ns6 25 -30 °C-'b 3 , bmQm
39* 66 mb 63ftJo6n - 40 °C-b* g* ^qjhm 0 *(?*c 63 ^ 3h*i!Dh*'b3(y 0 h*3*on C4 0 ^ 0-
6*h3 in n6o36bn3h*g *bn3 nonh36 b 3*c>*0 *3 g 36*g, C4 bn6ghm-
9 n 3'bh36330ymq3b (jb3cn sg5nQbs3ymq)3onb *cn3nb36nb 6nmjn3n3h bibobb. 0136-
13* *hb36m6b 63^m-qjfnofT'^r,6cn3^)n'J @i6*0n 0 D9 ^D'l:niD^DCn m^nO^nb 3 Jm63 C4
0l336sh336n(j (3 *&., Spartina-bs 9* Atriplex-nb b*b3m636n), hm3o36ou 1503 h3&nm-
63660 b*bcm636. C A M 3^36*h3363n m3on033n 3*p*0O* bn6*0>Qnb
bo*gnnb &*60 * 30m6 *0 n, 0 *3 n6 hmQ3b*u C 0 2-nb bn663onb qjnjb*un* b 1^(503093-
- 113 -
G6ba 0 6 6 6 ^ 0b 9 nb 06(nD65 0 3 6 6 ^ 0 6030)00606015 ff668m 866
(nb. 2.38 Ferocactus acanthodesSn SiQiOK'b

i>i39X<ih36nb 0i6mj)ne]6xD&s
pi ojrryjm(jn6oi3'bn Opuntia polyacantha-6
nbmpnhj&x dc^^Sc^îlD^n (Nobel, 1977;
Gerwick et al., 1977) Temperature [°C]
65 p i6 n b 95659 ^3933(15^31136133 (65b. 2 .3 8 ).

B3^>m-q}OTvyT)bri5oiQ'bnlj 80603380. ^hmin^Q 8(3365113015 cgmov
03680 q3rn(jmbn6oi3<bn 6yg3&5 95659 959361101 ^3833(15^31136133, dnhncnigig, 850m
j)fiQ5j)fTin936nb bni3n3nb5g6n 35F50O 8g>hd6m63pm6ob j,i 3 m (nb. 01530 6 .2 .2 .3 ).
<bm9n3hn bsh^ypob 8(5365113360 C02-nb 5bn9n05(3n5b 2,56551^01636 0°C-'b3 3qjhm
95659 ^,3a33h5^3h5lb3i5( 35j,h59 hmj,mh(3 jbni3n0363n 65h9mnj963<b5 yo6xn,
cgm^rnbn6oi3'bn 690365. 55'b5q jb x r|b £,3mqjn^36b5 05 05p 5p3 cnob 9^3651133690 3b
b03&5 055b0m36noi -500 -133, 05h50063563 6(5365(13015 gmoiQ36b5 05 60633630 -
3°C-056 -5oC-303 qj5h50363n. ^013093690 yo630ob 55960160b 938935 cgra^mbn6-
013'bob 3hm(33bob 556569365 6305 8089065(13016^ 5bn30p5(500b 55yo63nb93695m-
3 n 033 h3bri5 95301^09363905 55yn63ob b56j,hd0O3ni65b5 05 b5hobb'b3. h5(3 3ghm
0565005 Q^Snb ^3833115^311360, 0 oon 3Gghm b565hd0O3O5 838036 PCOb C 0 2-nb
5bo 8 oQ5 i5onb 5 5 6 bmh(3o3 0 3 6 ob5oi3 ob b5ôhm 0hm 05 3 ooi 3q)hfn 9565905 35jbn-
65b. 2.39 (138936n3 ] 9pni6n pbdnb yn633-

6nb 9389358013639365 qjn^nb
ajnvimbnbon^bnb gn6i3ntp'b3 (Polster &
Fuchs, 1963). ^3hêrn3i 6 9(536:^30^536
556(1635336001, 3mc5h3pn (135^636^
0nj363&b qjm^mbn6oi3<bnb ]6ihn 83663301
9n63b 10®C-b3 95 3qjhm 9565c ilo933h:>-
J,3h36'b3i}. idn^ma nbnbn 5^41131159
6 9 12 15 10 8311013063636 C 0 2-b <bb9oihn6 30(10163630(5
Time of day y i (Kappen, 1993)
- 114 -
<V)8CT>5 03*^6 900666090
0iO3hn 016336360360 (61b. 2 .3 9 ). 615361(13^3 (110036003 (?i0 nb 5 i 69 i 3cm6 i 3 n

yn63fib ^1361*1^03936001 3hfn5h3b3pi9 0^0(19361 C 0 2-nb cgn;|bii5onbio»3nb &i0my3-
536 x n 9Cfib bn6ioi0nb bi65hd0n3m6i.
63vyr>-gm^mbn6oi3'bnb “bêi i>D0 ^3hatf3h x n tU T Sâ 0i(?ic l!)D93 3h;>vbDhD2>'tD.
3nh330 (10580, n63n6nhp36i qjm^mbn6oi3'bnb 3nh33pipn 3hm|j3b36n (nb. ubh.
6 .4 ). 63^)m-q3m^mbn6oi3'bnb ,53033(11,53(1x11 0ijbo033n 83080361 51630^01*101 hm-
5mh^ C02-nb 13300b b i 5m0336 bn 5nrn 3 3 6 ^ (bno^^hn b i 5m3336bnjom ^36^0);
hii3 39jhm 8i(ni0oi q3m(jmbn6oi3lbnb bncn6m0j)hd6m 6xrr' ^ p i hiy ^cghm b6hiqjoi
b}6cnj3nb bnat6m-n6g3i5nh3630n l53Cn03^36n, 0ncn 3cghm bBhicgig ip633b ©15361(13
biim 0336biunm 336 ^ 5.
ibo0npn5ni R 33X (Sibry^yibo^nb (MditabruoiQ&x) ojrn^m-
bn6cn3‘bnbi gi b36oij3nb bnRjih336o ig i3 l5nh363pm 0m(530xf> phmnbio^nb
^33X n iJ3033hi(i3h36nb 0n0ihoi. 0mgxil5n3hn ^,3033 hi^ 3h x r> 19 13 (5 1^ 1 bmh-
^0399361 hi0g3&n03 gpnb, “b m j^ h 50 (110936003 buoinb 5 i 60 i 30m6 i 3 n. 10 3hm-
U3bnb 03bidpm 03ji6o'b036ni: (530n03&36n b36bvj)hi^)nb yn 6 u36v5hit5n369 n - Sij-
h36nb i530 X i l5n;>' hm&mhu ^(5033^13 bi3ib3bm hQiJyni; 5ih 533X n <33(1836,5360b
5 ihgij96 i n'bmq33h036(536ig, (11*183001^5 0 ^ 3 3 01*1^039361, 0i{,hi0 5i6bb3i33630n
,53833(11(53(1300 m3l5n830 n ij30i; 6 nm0306hi636 nb jn0 n3hn 9b b ^ j ^ h x n l53CnCD~
6360.
b35o|j3notn 1 ^ 3 ^ 6 1 0^(19361 bn^nbigOn igi3,5ii5nobi 91 x 3 ^ X m^ J bnov
6mb 0 n9ihcn 1913,511500b 3hnnj3b0n: 1^31531(119 hm0 301^3101, 6nr*ijn0o3ho (1353-
0i^ni 36n6ii(p0g3j)36i h3ijynnb bnfijihnb j)i6bi(be'3hnb qjn'bni3h-jn0n3h *536936-
I5nib. 181b 0 X 36^6 0 n 3y336 i 91610 ,53833(11,531136^3 363h5nob 35301 81*181(1153-
6i, 0 i 9 n6 , hm93bi(3 3 ip i0 ,53033(11,531136^3 b:j6oij3nb bnRjihnb 0116910116 i*i6noi
61*5361 83,5161*100,5360b 3q)hm 3^0161*1003(119 0nb3ih3&nb bi93i036ib 080331. 093-
103(1 030oib333i3n 03,5161*100^0 0093619 bhxyrooyîg ihnb 0m63bhnj,36300,
hm0 b36oij3nb 06,536^31*161 83061(1636361 3(10161 p i 00133 0<T>63'b3 ,53833(11,53-
hnb 839:^36001 qjihoim <31(1503630 (oi3h8m3m83mb,5i'bn). 3b 6163363601 61b.
2 .40 -'b3. b6hicgo 80193911503(10 1913,51(501 dnhncnigip 06n03630m3i6ni 0(5361(13-
61b. 2.40 qjn^nb b(p6m6b(j3636nb bj6cij3nb

n6(j,36bn3miib Sioin 3i6n^yiboi3nb
Qi9ibibnicn36x (Rook, 1969) Temperature [°CJ
- 115 -
66baorfj66ôb anb 9 6 6 oaa a a ^ ^ o 6 O301OD6 D6ou P66and866
6 sb. 8.41 Chamaecyparis obtusa-dn b36or)3nl> jm jcjn ^ n ^ n b Q10-nb 93^301x7)563 6pnb

j>36933C<n639n p>39nU j,3953hi^3hnb 339|3nh36Mxn>6 3(10136 (Hagihara & Hozumi, 1991)
3&nb:>o>3nb, hm0036n|3 0^11936036 gngn g i oouihn U3:>036:>gm6ob

ig^ncQbSn.
b]6o>j3nain ^031*163 0(53036:) ^ 3 0 1 3 3 6Qob bb3^gibb3i ghmnb>oi3nb g:>6i-
bibniOT363Qn ,j3033h:>£)h30 o 3ohm636 ob (53>p36igm6nb 33 bi 6 i 0 nbig (63b. 2 .41 ).
9ihig063s63 0(536:»li336b, (11*160360(5 “b:>0CTU>h9n(5 53635(161*1636 sbn0nQbijnib, nb3-
0136b, hm^mhnu b030cns93i'b(n33cnnb bi03hmqjn036n (3b30gmcn0ni6n 0(536^33-
6n) 36 cn6npn Ii35nm636nb q):>hoimqjmoi0n:>6n 3ihip063i63 61536^3360^, b36-
cnj3ncnn ^0 31*163 03(p30n 3^301 6pnb 13113 b3'bm6360nu. :>8ob 5^0m b36or)3db 06-
J)36bn3m6i 6 :,h.!>333C cgih5 p360 n 03g0 n3n (16363 601300 60nb 5 i 6053 pm6i 9 n.
b36oij3noin 3^031*163 3933^(1(1363003 bib3m6nb 6i3h(33C36nb (13501*161^ b:>63>-
0m ^ 3033 hi^ 3 h36 nb 0 n03hcn. s0 ib 6301039 3g3b^3(i36b nb 033j^n, hm0
613363(133680 b36ovJ33 633(139 b^b^ni, 309(13 'bm8o3ho <bm6nb g i 53663^301(136001
ihj^njjnb 56 83(*i30n 0oinb 8(336:>h3369o [ 128, 239 . ^hn3ni36nb, b36^hm3n^36nb3
g i b030 cnb93 b‘b(p3nb3nh3Oinb 031139063363 b3-0(336bh3Oib qjmoi036ob b36oij3nb
n6 ^ 36bn3m63 20^)C-<b3 03hy3t*i6b 0,1 -0,2 (iSmQn •6‘266"1 , 6^^9063^6301^
60633660 - 0 ,4 -0,5 iiSmpn •6'266'1, ihj^njjnb 3353 63fij36ob <3010103680 - 0 ,6- 1,5
)i8mpn ■0‘26 6 '1 Smhob. 013 38 8i336ih3ai3 b;}6cnj3nb 0 6^36^3016:^ 8333033(193601
“bhgob ^homgob b3833pm ^3633(1:^3(13^ 933116363301, hm6 636nb6n3hn 3i 6 n-
î^nboi3nb b36cnj3ob 06^366031*163 01330363 8bm0mg 0 ,25 -0,4 |i8m0n -O’2 66-1
crp(1503660 [ 132 ]. 3b 6383(101003603 6303br*i336n 6i3363h336nbiai3ob3|3, 35b30133
6ihobs g> 8010b 65363(13013 63g:>h36ob 838016333^8013.
63jy»>0)mîT)bn6oijbnb m i^ o S ^ h ^3033(13^3(13013 36360 g:> ^383311^^3(1300
^(*>3(1360 jpBn^gnb 01*193031503(1 (33000363b 336o^3ôb ^3633(13^3(1300 30(11*163-
6obsg0o 6i336ihob 3t)0o83ô'b3(3nob 83g35^9. 3301031503(10 ig iS^ i^ o i bSohsg 5 s -
8 mb3^ 30 Oi ^("6^^36300 6bigymg36noi 3b3oin 8i*ip30:>i3O3hn l53Cn03^36nb 6063(101.
Oxyria digyna-b 8010b 3^1*1^03360 bnoi6mb ‘b38i*ij83g36:>'b3 (13^2^3636 C 0 2-nb
8013601j0nb ^3833(13^3(1300 m3^n030obi gi ^39^ btr^hob 303(11*1 06^36603(10 53911-
g36ooi, 309113 shjôj,xn 9 * 3ohojooi, boi5n3nb ,b38f*ij83936obib shj^n^jg 3*)mvin-
33660 9^630 ^3833h3^3h36Ib3 ajm^mbnba^bn 0^(1936* ^jqjhoi b6h30339 93 3033^-
- 116 -
(W8016 133^ 803666330
ODhig. 309113 9 oiobi (6sb. 2 .4 1 ). 50933 3(1010 815190010: ^ 3 9 0 39i 65 m9>5 g i

15030 b i 6 i 3ohm b n 60916 se<3&3c n 61(119863163 djfijob A trip le x le n tifo rm is-rh i c (T'-
6360 5160)6313936016 cgmor,ibn6on3<bnb 0 D633hi03 h x n 8(139001 8bm0mg 8i(piQ
i53833hio3h3&b3 bhgnb 338016333190. bn(jbnbi9 oi 6 n 3gi66mb 2,9016360 10 838-
cnb333i3n 16 ^ 13 6 3 6 3 6 0 ip i0 qjm^mbf»6cn3'b3h ijon3m6ib, 9 i 9 n6 hmg3bnj bi6i-
Srihm bmprib 59016360 8ie<ico 03633(1i^jhob 9 18 ^ 0 16 36 39 0 8 mj839360b 8393519
C3m^mbn6oi3'bbb 83330^19 16150(13636. 3 ^ 11^ 53(18 1 90133(1361501(5018 33fK)o36i
5i8mo63omb (5390936360 630rr>-q}mi5mbn603bnb mSonSiCDh 91 81908000113639
^3833(11^3(13630; 0 3b id 0 36x o i 3b 9131390(1363901 2>ih38mb ,53833(1103(1360116.
Oxyria digyna
Arctic ecotype Mountain ecotype
Alaska 68° 56' N Arizona 3414 m
£
T3 10
O)
E
0
5
\ " ----
10
5
^ •
\
°
'■O
----1 3 / 7 X \
\
.
''O _
— 30 /Z4 *C *<Q
\
_1---1--I J----1--- 1—i_l---L. ■ ■ ■__ 1__ 1__ 1__ 1— t- o
Temperature
A triplex lentiform is
fob. 2.42 6»iTHgm^mbn5oi3'bnbb gb pidfib b^fjcnjsnb (6n^mjm5ghnxn b^&jtfonb) Qi9myipo6x 36i

Oxyria diginaAi ihJ^njjX 3i^ ic 8omb hm9x n u rfbhgjbi giiip n
(gtpycbdj; 13/7°C) 91 9i(nion (qco/cÔ1 30/24®7) Snhm&^&Sn (Billings et al.,
1971) Cbxi Bsbi'bn). (jb x (ec^b bxJîpm 43°C) 91 ^hnc bs6i3nhm tmpSn
(gcvib b>3]icn i)0<G;)hi^]h> 23°C) dm'bihpn Atriplex lentiformis (C3) Bjyn-qjpvjmbnbcnjbnb
gb0 m»)ne36x 3S>i (J339i 6iWbn) (Osmond et al., 1980)
- 117 -
66b3 0 6 6 ^ 0 U 8 n b 9 6 ff>066 8 9 6 6 *^0 6030 )0 0 (f»0 6 0 lj P 6 fn 8 n d 3 6 6
2 .2 .5 .3 . C 0 2 -n b 133W 6ycn01 an a^h^636i
6yipn, 6ib9nhmhgi65nb Sbg^big, qjm^mbnBcnj'bnb i3)jnp3&3p b^bb^hi^b 6ih-

9msgj,36b. oij0iji nb 33h bp3bi q}n(jmbn6l53'bnb SipnSn^nhQ^p qjiAjmhig, hig-
5^6 3hmlyn3p:>‘b3nb 9ipipn 3nghbîjnnb 33bi6ihfi]63&pig bi^nhmi Bypnb gngn
himp36m&i. 6ypnb 6iiP3i>m&noi 5^0^163330° DS^Dgoii SnHyjppibnbi qs 0bcnn ^h-
fcnhnb ou>6 bg33b ojfTi^mbnBcn^^hn y^n 3m6nb gi^ 30:> (6sb. 2.43).
3^6530010 3hrt(33b36n b9 cn3ipbi‘bhnbncn 6iip3&35hd6m&n:>h3i, 3ngh] cgm^mbnB-
013^0, g i i 3nb 2>i0m cgm^mf^binhujni gi 3n^pn;Jm6ghn3pn b36cnj3ncnn 3hm(j3b3-
6nb n6^36bn3mbi 3^336b 0bmpmg 3i 3n6, hmp3bii5 33h3goii g ^ n g h i^ m biy
3iPig gngni. i;)ogi6 5i9m6Qn6ih3, Bypnb 6;>>£360161 5i0m3pn6g36i C 0 2-nb qjnj-
bîjnnb cgrn^mb36cnj3iboii6 93q)ihg3&nb aii6giOii6m&ncn 333^nh36b3n.
£)h^pm3i6 9^36^33630 Bypnb goqjnyn^nb 93g3&ni 33nh33p3b ym3pnbi
6 :>533 &nb b 3h3pnb 333n6 hm336 i g i C02-nb U3pnb g ij 33nai3&i. &i533 &nb Qib3h-
6sb. 2.43 gmoipnb b3j,936,jnb 63^™-

q)ml5mbn6on3'bnb 06^366031116^5 91^33001361
Bypnb 5 ih £ 3 ib a ii6 9 15 13 3 ^ 3 6 0 0 1.
q)mljmbn6ar)'bnb 961^93630^13^6(11 6ypnb
50%-ni6n 51(153^ 936935 (Kaiser, 1982) Relative protoplast volume [%]
%
10 0 - Helianthus annuus
CD
75 80 - ■31
0 _ Photo respiralion
s 40 - C l9b3b]9 bnhib ojpioionbi gi Jomhm-
0 W
0) 3csbvj3&nb 63ljm-0)ml!pmbn5<n3'b'b3,
0 Electron transport
JZ - 1 DC
a. 20- Ph (chloropiasts) 300ijhm636ob j)hi6b3mhj|'b3.
Ph (leaf)
Photophosphory-
q}mbq)mhnonh36nb 3hm(53b'b3, Sn^nv
lation jfi6phn3o pi oji,iiynb36oij3nb 06^36-
“f- ' "1“
-0.4 - 0.8 -1.2 -1.6 -2.0 bnsmbÛb] (Boyer & Bowen, 1970;
Keck & Boyer, 1974 ; Ortiz-Lopez et
Water potential [MPa] al., 1991)
- 118 -
5680)6 03^6 900666330
3i 33nd03&i 5i 3mn63nmb ^ h n b gi6i00i (536ni6 m6i 3iu, Qncn^Q^^b, 013 0ib

oii6 it>C53b 6nigi5ob (i»36oi6m6nb 336(301136^. qjmcn0363o Bypnb gibigo 31*1^36-
I50ign 3i3n633 116333b qjm^mbnScn^'bnb n6!Jn6oh3&ib (6ib. 2.44), dohocnigig 303-
j^hmBnb ^hi6b3mh^bi g i qgmbqjmhnoofît^ 3mJ03Q36nb 5i0m. hm^mhu B^bn,
i j 0mJ03936b mhn 03ji6n'b0n: 0ipi005hd6m&nih3 6i533&ob 9Jm63 0^36^33680
(0hi 3i 0n b3 0(336ih 3) C 02 -ob 3ou>6oij3i 0(3ohg3b i 6ij,336nb b3h30nb 33306*1m-
3360b 83936^?. &33h 6i0ibm 3i6bi gi ^ h m g jo ^ x bib3m6363n $0 3b 3oh330
hoj,3o j,i3m63330Oi 03î6m03hn n63n6nh36ncn.
3fnny)0n3nghxn 0U36ih3360b ^nghi^Hjnnb bihnbbo gi C0 2-ob [5305

3noj>n0R3ngh3C 01536^33680, hm^nhniiii 6yi00|j36^3360, 01^36360 g i bi3b3-
6n, 3hm^m30i'b0nb Soghiîyoob bihobbo 5ih30mb ^36oi6m6ob 93bi6i0oboi (ob.
aii30 4. 1). 3b 6(536^3360 bBhiqjig 061*1336 6yi0b, hm^mh^ i 0ob 8Qbid03&-
0m6i 0n3U30ion g i ^ 3 3 3 fijihi imhoijQ3&36 0ib. i 0 n^m3 0ioin 6yi0030(5330m6i
(33^036^91*16b i 3n6gob 33bi6i0nbig bi 63mj)P3 33homg363o g i 3i3h3o Bygnb mhcn-
j0nb 330i333Qm6iboii6 6m6ib6mhm6i 9o o3ymqj36i.
3moj)n0m3ogh30o bib3m&36nb qjm^mbo6ort3<bob gi0mj>og3&3p36ib 6yi00 30(333-
0m6iboii6 5i0mbibi336 3.6. m3ô030ob 0h3gocn. jbri3O0363n C02-ob 33^63-
3obig0o gogn Qntgjbjhn 6o6ii(p3g3j,m6i, hm030bi(3 3b3g36ooi 8i(?i0o 6yi0330(3-
330m6nb 0jm63 00^3636^ g i bi3b363n, 30o6g36i 63^m-qjm^mbo6cn3cbnb gicnh-
5353i3n (6ib. 2.45). i3ib o ii6 i33, 0(336ihob 5i3m3hm6nb gi6y36obau>6i33 0ib 3n
oii6pioii6 0yohg36i hmj,mh|3 b36onJ3ob, ib333 q3m^mbo6cn3'bnb n6^36bo3m636n.
bib3m636o j>i6bb3i3g36oi6 5i3hnb <gihgm6ncno ^36ni6m6nb 6^33636036001
(RH), hn030<b 3g i (3 6yg3bi C 0 2-ob 3301301)36^ b03p3oiob 6yi00U36ih 336oboi3ob
0i0n0n^,oh3630oi 7090 % RH, Qoj363&oboi3ob - 80-96% RH, bi3b36obioi3ob ob,
6333036*103, 90%-‘b 3 03,joi [ 187]. 063i 63 6yip 3i336ihob 830153300 0^36360 6ij)-
03605hd6m6nih36o ihoi6 g33oghi^>i|3nob 9o0ihcn, 3ogh3 03hj-063i63 q9m^m6om5-
1^360, higj)i6 i0 3i>i6 ibj)630363 o mb0mb3hig ijô^hn 2^0301636^350 83gih36ocn
I3n^>ii [131]. cni0mqjo^)3bob qjri^mbn6oi3‘b3ho iiih i^ o îhîg ihnb 835336300
Water content (g H2O g'1DM) Water content (g H20 -g'1DM)
6ib. 2.45 Bypnb 336(5330^606 6i3CD6i 3mnyxvn3nghxn 9(506ih 33&nb qjtn^mbn&j^'bbs Qi bn663(Yib
bj&onJsnb n6v)36bn3mdi'b3. pnjoBSn (Ramalia maciformis) g i bi3b9n (Barbula aurea) 63^>6nc3hn
3hm|33b36n sibn^hpj&i 3nghi^jnnb qjibiSn; <koi J>i3m9hm6ibcnb6 3hotog, CC>2-nb 33<Ji3nb3&i
3 i 33crhig iC3&X m^ (Lange etal., 1987; Rundel & Lange, 1980)
- 119 -
6 ,bb3 n6 6 ^ o b anbaafoow 93 < n ^ o 6030i 00 r^0 anu &6ff)anda6^
jjhognb 6 y ic 939 b3X m&nlj b^0*1 0^ bi2|9 img fciSnibi^x 5390036360156. hmg^b^
9 oicni6 ig 2,i9m99hi0o oiig^bn *6 <301010^2,360 2)30^3 i CPn@63Eb6 bmpQ^ 6yi0-
939l533Cm6ib, <30100^06013^6 Shmij^bnb h35:10035505 b x h59936083 bixn9 n 9 o-
np533i; dnhotnsgig 3b ho6xoi‘boi6ncr)mbq)ily2>ih6oijbo05bob 6301506013^001 ihob
5563(1(101636300. 963563 6 yi 09 u36 ih 336nb 83853300 9^363606 33O3hoi33h30 b5b-
301636b Bycnb mhoijQob 330i3nb36i 833800501 036ni6o 3^311095655 jjn, bncnbob
bibnoi 6yonb ihihb36m6nb 838016333580 - ibQoi 30(101636805 nbn6n 0693636
005305601 93,1^.60103(1 3hmu3b36b gi ^>£>63336 C 0 2-nb 93936001 6i056bb.
CnJ3636b 5,59013938539501 bibsh^Qbcm oii3nb363h36i6n, (101893605 jp 6 >3nhm-
63636 C 0 2-nb 3033^3(1 83013053656 8501 jbm3n0oii 8i(?i0O 6yi 0038533001611b
33(100193680. 65b. 2 .46-133 6563363603 39i66mb 0nj3&3n C 0 2-nb 15300b gc^ho
gn650n])b (5^0183360 fiiOih3&xni ^ cnj36oboi3ob 636361103 3ohm6363o). Q>i9 ob
5368339016380 cri503bn ooi3nb36b Byigb ^360560 ^53(10956, 39(13 gncncn 6580-
9565(5. 9 bnb i 9 mb30ob 938935 90^360 9bm0oig bi 9 o b33aiob 5569530016580 ibo9 -
n0oh36b C 0 2-b 93658, bi6 i 9 539018(1365 95 5i9i5hg36i.
5iboii U3pa ^Dh^0m3»6 91536^33680 6y0ob g^qjnuryjnb 3nh0163680

91139b, hm93 cni5 5p 63 hb 9590121(19363036^ Bycnb >|3(i53 ob bnfijih3b3 gi 5 iboi3
5303 b 9 oihnb, mhn 2>hnon.!DCn 63(1,1,000 3j 3b: bp^h o qjoiooibobo^bob 9ijbo9 iC3 h
83b5dc36coi6ib5 gi i(j>g536ncn 363(1 b 9 oihnb g i fciboii 15390b 6x013360 63(1^090
Tim e of day
6sb. 2.46 j g i 66 mb enj363n (Ramalia maciformis) CC^-j^bcni U3c>nbi qs Bycob 330jj3xm 6nb
ecnjh n pn6i 9n>p. ps3noi Qnj36n 6y>Qb n(^)6b 6idnb b j9]i(v){>n<7i,
9^&h:>a pnpnoi n{,n 3 i3 3
5i 8m39hi(yu). 53hnpig gi3^hnb3CO ih]: CC^-nb 330i3nb36i; 8530 ih ]: CC^-nb 3sh v
CDCDhie e i 3 ^hnbxn ih]: Cfi9nb 33hnrnpn; 0130*0 3I13: Qfpnb ^hnmgn (Lange et al., 1970)
- 120 -
<V>80to 03*^6 81306.^090
(nb. 6*b. 2.48). <bc>3*ho annp633i 6yonb nbôwi b^h3bnb*b, 6*53360

n6y3&36 9*63(13*6. 60h*o 3*3^33 ym33ooi3ob 06936* Bypnb 301,1)36150*006
3a 6n9353Cm 93615*1113606*6 (^360*6 3 * 3 ^ 3 *1*6936*963 3<3hm 530*6). 3*3ho6
9*6*00 (J36n*5m6nb 6o6*hoi 6*3*63601 h3*Jun* 5*6663*336300* b*b3m636ob
606393001. 013 ^013003660 6y*on 9*8*^5936* 3oIi33qo 3(10603300 B3hlyioo6 5*3-
qo6 336935, 5*'bcn* (i3c* 66h*qj*g *(>19536*.
5*<bcn* 153006 630*^3*60 63^ 0 0 0 5*6o6*'bp3h36* 6*533606 6h3co *6 oiooijdo6
bh3on 9*63(13001 9* Byonb 6*iC36m6nb 3ohg*3nhn 5*3036001 3hryyn30*<b8*'b3;
*8*b *95000 *^36 86010019 (3010103606 36n9363om3*5n 93309(1*^*15006*6. 0136(5*
<bfri533h 36 8p50i8*(i30i6* 3001*11936*, *0*9536* 0530*6366 9* 6003(10 5*8016(101606
636935 5*hi333on 30(10163660 cjoHy’i6o6oi3,bo6 b*6yo6o 36*ho 33^ ^*6^ 936
*("9536* (6*b. 2.47).
5*8oi3hoi6o6 1501,03606 5*68301(136*6 6306036* 63935*9 6m3y336 6(536*1106
5*6016(101016* qjm(5mbo6oi3'b3hn 3(101(5363606 *g*3|i)*(503h0 601930*15006 8301636001.
5jb. 2.47 A j,i 8n 9 hm6nb jo jo iy 1 Abies

halsamea-b 3(1010 6(vib i(p0m6nj363&<lb
6O0)l™ 3 miim^n6°i3,b'b3 (Clark, 1961)
6sb. 2.48 93t)nqjn03n 6yonb 3m^]6un*pnb

Qij330crt3&nb 5130361 COj-nb
330i3nb36i'b3. S b3iX36tl36n; M 33‘bmo}-
0^360 (6i{;ibm3i6n mh(vj66ni6;)&n, 6ih-
^30^136360 91 336093(10 h35nm636nb
b336n); Sc b30o336np3ho 91 b3601(109300
h3jnn636nb bioohmojncjhn 636^360 91
6336(1; X J}3hmq)n(j3&n (ihng3pn
h350m6]6nb 650*13013636(1, 63^360 gi
b33&n) (3hi3ipn i3,jmhnb 6^61(336011
606393001)
- 121 -
66b30t*»56« m 8n b 36(nDf>6 ^ 3366 g?0 6030)03<n3&0b ff6fn3P)d366
...^
Agave
a
s%
I fe 2
0d \
7 d \\
13 d '
8 a 0
8 12 16 20 24
tub. B A ? CC>2-&i'bais UJCi CAM 9036^33680 63^03'* } 3hnmg9n. 9ih(jbD6i 6 ib iV i: 336363x^9°

diWtehen o3nqjn^;jhn mhjnp3nb ( Schomburgkia humboldtiana) C O j^ b o fe U3Cn'> <y<3n3hn epvi^hn
gnCidni^ 63060^6 33hnme9n (0 ) p i 63^03°^ ( • ) 3nhm&3&9n (Griffiths et al., 1989). 9^3336} 6jb-
itjn : s&i3nb C 0 2-6i'b<7t> (jx> 6ycnoi ish jn 9m9 jh i53&ol) 3nhr»63&3 n (imBvihmpn) g i 63iC3obib (7-
13 ep>3 36yomg) (Nobel, 1988). 6yenb 930)0(50^06 3nhro&363n 6153360 p & jh x n i 9o0 C° eC*1^
5i 69i 3pm6 ian g i CC^nb 830130636* p i 5 i 9 mymo)i SjbidoQjgcre) 9bfn(xnQ (pb8nb biiO>368o.
Bypnb 6ij>Q36ni6obip0n C 0 2-nb 15300b 0{,hd6m630m6b, h>u ‘b 30 moii(n6 o36 x n

mhn ihn(j n i x n 9ifi336360ob bngng30o wibibjbi, bibQmbnb gi0ibiboiav)6xn
6 n9 i 6 ni (6ib. 2.48). Rhgnpnbi06 i 6 n 81536*113360 (b^nmqjn^^dn) g i b3-0i536ih]au>
0hb3son bib3m6i 3i093h3bip 0j,hd6m6nih3i 6ypob 306 n836 x < T' giiih&3ob 0n-
Sihaiitf jjn. 0301(13 0bho3, bn6icnQnb0my3ih300i 3(536ih33d0i g i 6iQibpi3i60i
0^15300136360* 03ndQ36i 5i3dQm6 umjpg 013 633*1:^ 06n 0 36xfT'3 :>6 6ycob 93030-
I5fi^,b - 3b gi0mjjne3&30ni b ^ m & i ^ g i 'bhenb 30*101636^3. >0 o»3i0bi<bhnbncn,
C4 g i C3 015361*1301* 9 mhnb 5*6^53*3361 306o936x 'T'i. *101501*113 0mbi0mg6x n
oyoi, 08h*0o ie 6 nQbi0ymogxnb, 15*130133 3916601b 815361*133613 dnhnoiigig 0i(j>i-
Qn 0i0n0n^nh3630n 016336360360 1^301. 6ypnb 93030150^06 30*1330033 6096360b
5i0mR36nboii6i33 CAM 01536**13360 b3hi336 011301601 615336b ©01300 gcpnb
2)i60 i 30 m6 i 0 o (6ib. 2.49). b36cnJ3ob ghmb 6ih0oij06o0o C 0 2 51*1301 ip ih
61b. 2.50 Glycinm ax-So 5ycob 93030^^001

{>i9m6333cn 6>jrn-g)m^mbo6on3'bnb 338^36^
Time of day 9dpnb 5i59i3pm 6iao (Raw son, 1978)
122 -
<W8 (J)6 0 3 ^ 6 9 0 0 6 ^ 0 3 0
539moymq)3, 9303»jnb 939cnb3b 935(1013000 C 0 2 3)30^3 539 0 ^ 36 36 3 qjoiombob-

cn^'bSn.
53i03nb 5330363 6 3(joi-<30ilj,oibn5ai3'b‘b3 9 3(^ 0 36 3 3bb6n0 nJ63b Bycnb 6303-
6bnb 53cn330nb6n636noi. Byonb 33hymqjncnn 63036bob j^bhgnbib 8 5 363b3013
6 yonb 501^36503905 b x Dghm 33(1yoicgocno 69363 p i 6353360 5 gpnb 9301(13 63b3-
3ih9o b x 3°5hri 9 3 ^ 9 g i b365hd 90339 063(136036 (63b. 2.50). 63b. 2 .51-b3
63633636(13 6yonb 93q)n50onb3g9n bb3393bb33 95hd6m63om6nb 9^0163 95363*13013
002-53^013 5300b 9(3*03(10 9 n639 nj)3.
63b9nh63gnb 6 ^9 ^ y i 93 Bycob (ufojphfen

53hg3 o9 nb3, hm9 C 0 2-ob 9 oi36oij03 95363110b 903(1 b01(151109363 635336001,
nbo6 o 3<b(i363xyfT1<3D6 Bycnb nhoijoob 90idh30i63b qjmo^nb 8093 3(10936 53)13-
9 m9533 o 3(1380. C 0 2 hm9 9 oi36oij3b, 95363(139 33150036039 3693 539oiyoib 93
933)3h5mb Bysco; 013 ob 9339j5oh36b Byonb 539mymqj3b, COj-nb 633)3905 936-
5oh9363. 3b 3hoi03hoi]p39oho d30O36 39(13 oj6i 938663300 [91, 151] 93 h05601-
6ho339 539oiob303 h0i50ih5 3bo9n035nob 93033119363 9 0^93(136390 Bycnb h30V
936oi63boi36. 33)363b3)63c b36b ob93(i3&3 ^3(19060: <3T»lynbn6oi3'b0n Byonb 53801-
yo6o6nb •xyril^“ihm6nb 93^33636300 (W U EPh) - 3b 3hob 939013)0936x3^3
q3oilj)Oibo6ai3'bb3 93 oh36b5oh3503b 3 oihnb:
WUEPh= Ph/Tr (ji9oi0n CC>29"269'1 / jx9oico H20 . 9-2 . 69-’ ) (2 .8)
q30i(joibn6oi3<b9o Bycnb 5i9oiy3636ob 3*33^3^0163 (10(36016(10339 539oib3^33b
6 9 12 15 18h
Tim e of day
63b. 2.51 635330b 3g3d6m9o (nbhi3Qo) Bypob 930511|yi,yib3gdn bb33g3bb3i b36bol)3hm6ob 0Jm63
9u35ih3360n 63tyn-qjml)robo6ai3<bob gjpn^hn [33000363611. ESypoai 33301 9m93h3536x bsb3m63&3n
63d)m-q3mj)mbn6oi3'bnb 933(131*13 8306096363 Qbmcng 33ip(nnb b i30i3&9n. qjrymboBo^'bnb 116^36-
bn3m 6i 39(13 gopioi 6mh9nb <33(15036303, bmpm 9339e>ob3b bgjryyi 3j3b do03h 933h3bo3b. *13015031101
bib3n633n qjn^mbnBoq'bnb 931*1(13 9(jnh3 3nyi 3(3x16096363 b:>e>39fnb bb3CTt363n (Schulze et a l., 1972)
- 123 -
G^banfn&^^nb anbaaADto aa^ô 6030100^05011 s ^ a n d a e ^
qjmoDrri09 n 8rt00n6**i3 5**601* 153C^>-

o8ob*oi3ob, *1018 5*3mohoi5bm6 5**13001 30*101636001 5*630*101636300 C 0 2-<i6 *
Qi H20 -b j,m6u36,!)hbi5nxn 5*1*0036^3606 001^03936* 9*, *5*130133, 81536*11301*
b^h3jjP)3hxn 9^> 0336^503*10 8*6*60*0136036001 5*6b*<bc'3*i3on 3.6. b^nqjn^jhn
gnq)3<bnnb g*b *6*60*01360*9 b*^nhoi* 6*680*101*19*6506* 9* 6yoob oihoijoob 6>g>
0oidh*336ob*g0o 6o6**(?8g350i&*oii 3303**193606 [9 7 ] S r H2°/ E r ^ 0 (*6
*1360016*01* 3303**193606* 5*1*-20/ îng6i. *10^36*15 6*53360 0ot0o*6*0
po**, C 3 8(536**1301* 38(1*303601606*01306 6n6**(?89350i6*oia 8309^936^ 0,1 -0 ,4
39*106, b3 8)536**1301* 38*1*303601606*01306 - 0 ,5-b, C4 8(536**133680 *3 8*633636-
0368b 831^036* 0 ,8-b 8o*(n6noi6 [64 , 121 ].
gng33<bnnb 30*1016360 15300b oih 3*101(53680 *h*og36li3*io*. 5*113 1*3*106 C 0 2-nb
im6i536v!)hiun3c fi j)hign36vjn jooihoi30*6^3606 C02-nb 3016(536,3*1*15030 6 * 1^ 3 6 -
(101*6 839^ 36001 5*150036001 6*30360*, 309*13 H20 -b 3oi6i536(jh*(5o30o 5 *^ 0 3 6 -
tjo gjfTicncnb 3 og* *h3b* 9* 5**13 *j>8oibq}3*ioib 3 mhob 5**i3 3*3*iob 6yonoi 5*33*13-
6*893. OTD 3*3hnb (j,3833h*(i3h* 20 °C , 6oi0oi 8303**19360010 <1360*6016* 50 % ,
6yonb mhonjoob 5*1*9036^0 9**600136001 20-33*1 *("38*^36* C02-ob 5*1*9036^6.
Sbmofng *8nb 5*0m 6yon6 *01*101^036* 8o8go6*li30i6b 5*150036001 303*101 66*1*03*9,
309*13 C 0 2-ob 9 oi*6oiJ8 *. 5*hg* *8ob*, 6yoo6 8(50*13 ‘boiSob 80103330360 900336-
goh3&36 1,5-33*1 303*101 66h*q3*g, 309*13 C 0 2-ob gogo 80103330360 - 3*1016*0*10
5*i*0O36l3nb 30*10163680. *hb36oi6b *5*130133 03360*836^3*10 5*6^3*336*60 go033-
tinob 5<b368oi5: 33h 3*1010, 5b*, *101830015 369* 5*o*hoib C 0 2-8 * jooihoiSpib^bSn
8oib33gh*8g3, 303*101 5*id30O* g*, 8301*1315, 6b6 *h3 o C 0 2 8oidh*oi6b 5*^0036001
303*101 630*. *80^018 q9oiôibo6oi3'b3n 6yoob 5*8oiy3636nb 3<33djj3*ioi6* y01330-
0130b n153036* 01330 153*036*90* 0og33<bonb 3hoi(53b'b3 8mj830o 03*^01*1360.
h01036*15 6*53360 8cri0o*6*g po*s, 030101006 8056001 *(1636300 goq33<bo3*in
5o6**(ji0g35oi6nb 5*801 C 0 2-nb 3 oi*6 oij8 * 0080^0*^36* 303*101 83(3*g, 309*13 ^ * 6 -
0.20
a Quercus stellata
a Quercus alba
Ph, Ph/Tr Tr • Acer saccharum
■ on t ph. o Juglans nigra
ST 0.15 ■
60 ■ b &q
Q.
—H
« §5
0o
<o
CL
C. £© E
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40 -
Ph/Tr
O -■12 SCM
o X
*ai_
#
V ^ -
3 20 - cn 0.05
... , II
ft
FC PWP
0.00
25 20 15 10 5 -1 -2
Soil moisture [%] Leaf water potential [MPa]
6sb. 2.56 90hip 3nhR&3&6n qjm^>rnbn6cri3Vi6oi3n6 5ypob 5180^363606 J33ii3hfi6*. 8^15636*

b^Kicnn: 6ypo6 {p9myj536nb pifcicn 3<33lj:)h|T'&s (Aho et al., 1979). 3^ 3336 * b^hjom: 6*5336nb
gsb^h3nb b ih jto Bypnb 516011063606 jghm 3yn6m6n3hn p i 3<33:J<J3hn 9 15100010 (Ni & Pallardy.
1991). 9hi3ipn 6631 8151000*1 nb. Larcher ( 1963a, 1965) pi Mullerstael ( 1% 8 )
- 124 -
<W8(Jtt 03^6 803666330
b3nhi(jni. i 9 oîcbibhnbooi, C3 91536^33690, 9 icno gi&i0n jphbmJbnQi'b^hn ij-

^n3m6nb g^m, 6i$03&ig b 30b iy h x 8gj>m8ih3m&i9n n8ymq)360^6 C 4 9i536i-
1133601^6 939^36001. bi3j)3cn3bm ])m93hrTi9 nbn 5 ypnb 9mb9ih36ibi g i C 0 2-nb
3oii6 cnj9 ib 9 mhnb 9nnp633i 9 b9 n6 , hmg3biu 6^53360 6i6o0m6hn3 pnii. ib3cnni
9g5n9ih3n6i 6y0nb Q3gn(5nonb j>i63ncnih36nb gib i 6 ynb9 n, hmg3bi|j (530nb mhn-
33 3hm(53bn 3J)33 cn9nonh363cni g i 93qjihg3&i qjmombnBo^bn/^hiBSnhiuni 39i(p-
C3bni (6ib. 2 .52 ). hmg3bii5 &ij,33&n cnnoij9 nb gib^hxni, 93ojihg3&i qjmom-
bn6oi3'bn/li)hi63nhiuni dgn3h 3(i39 i, 3n6ingi6 C 0 2-nb 93 b30 i 5^0036001 3qjhm
9 3 ^ 9 9unhQ36i, 3ngh3 ohi6 b3ohiuoi gi higj,i6 Bycob mhcnJcn 5i6ij)hdm&b &i-
9mb30 ib i3iin^30nb j>bncn v>»hi6b 3ohiuoi).
cgmombn5cn3'b9n 6ycob 5i9my3636nb 3q)3jo3hm6ib3 6i3030ib ibg36b ^0n9i-
^hn 3nhm63?>n. pncob igh 330 biicn3b9 n WUEph 6 33x 3^ *0 3, 9 3 3 0 ^ 3 9 i p i 0 ni,
bi6 i 9 Si3hn b id in g ^36ni6ni g i cgmii)mbn6cn3'b9iu j>i9 i i 9 i bn6ioi03(b3 3^33
0ni(p6ni 9ijbn9i03h 96n93630ni6ib. gpnb 5 i 69 i 30 m6i9n 3i3hn cn&36i 9bnb bbn-
33611b jp3036ncn, 9 nbn 0)36ni6m6i i03&30m6b, imhovj03&b obhg3&i 3i3hnb ^3^-
63036om6nb 5i 9 m g i 93qjihg3&i cgm^mbn6ori3‘bn/l5hi63 nhii5ni 9ynhg36i. i 9 9 q-
qjihg36nb 939(5nh36i b9 nhig gi»,i39nh3630ni m6lt)rTi&363<bcni6. higj>i6 qjmom-
bn5cn3'bnb 3<33tohm6i m6om$363bnb 9i6dn0<b3 9unhg36i, WUEph -0(3 i5h3cn33
31539i - &33h bib3m6i9n 3b 39cnb333i y3i3n0m6ib ( 9 ij>., bn9 n6 gn [5 ]), b m j^ h
3b bg3&i 6iymqjnb 5i63nou>h3&nb 33hnmg3n. b^h^bn ^1130133 }9unli36b W UEph
- 3b ihnb 3q33j(in- hm930 ni5 b^h30133 cni3b nR36b b(jh3boii6 ii39nh9n.
3im0nf,nnb, bmqxnb 933h63m6nbi gi 93iiy333rTi&nb 9n'b63&nbicn3nb fri^n-
03bncn 30jhm n6ogmh9it5n30 ni 63gpn 9ibnb 3hmg3jijnnb 903(1 9 mb9 ih36 x n 6y0nb
himg36m6ibcni6 Qi9m3ng3&3036nb 9iR3363&30n bhgnb 9 o i x n ^hnmgnb {,i59i3-
0m6i9n, 3ngh3 ftiboii 15300b 3hcnj3hign îSbib^h n b 9^336360360. qjmombnB-
oi3b9n 6y0nb g>i9my3636nb 3qj3Jli)3hm6i 939g3j,6inhig &i9mnbib36i:
6ih9mJ06ncn 63900 mhj^xn 3bbi [j,h 09hipo 6m6i • H20 ]

WUEph=-------------------------------------------------------------------------------------- ( 2 .9 )
9mb9 ih3&xn 6y>pn
6ih9rnj96 n0n 63ggn 9 ib i g i 9mb9ih3630n 6yi0n 93nd036i &i9 mnbibmb

3hcn330o 9 i536ihnbioi3nb i6 9 u36 ih30ii 901300 oii6ibibm&i6m36nboi3nb: 33V
6 ^ 36 30 939cnb333i 9 n ( 3oh330ign 3hrng3Jon30n 6 nb By0nb j)i3 my363&nb 3093^3^
m6i) 63ggn mhj,i6xn 9 ibnb 3hmg3Juni 5i6nbn03&i u36m‘bob 9cri30n qjihcnm6nb
9 n9 ihcn (nb. qjmh930 i 2 . 19 ) g i 9 mb9ih36300 6y0nb himg36m6i j)i6obi<bc’3h36i
bh30o 33i3m^hi6b3ohiunob bngogncn (nb. cni3n 4 .4 .1 ). W UEPh 6ih9miQ536b &3g-
0n 9 ibnb 9i^36nbi g i 6y0nb gi6ibih3nb n6vi,36hiC3h j)i 9 mbi(i)X36^> ghmnb
bi6&hd0n3n 33hnmgnb 5 i 69 i3gm6i9n, hm930 ni5 hi9g36n93 i3nhngi6 'bhgnb b h x
33hnmgi9g3 am n ^b. ftibcni 15300b oii6i<3ihgm6icni (WUEph i6 5h C 02/&hH20 )
im633hbni jjm633hbnxn ogij(t|mh36nb 93933m6 noi 3 Qnd03&i &i9 my363630 oJ63b
6n6ib6ihn 93qjib36nbioi3nb, 9 ij,hi 9 nb ih ihnb bi3b36ncn b3b(!)n. i 9 nb 9nb3boi
nb, hm9 9(536ih3cni 63gcn 9 ibnb 3hmg3Juni g i 9 m^ng3&30 ni ihi 9 ih(i,m jpboii
L330nb n6^36bn3m6ib3, ihi93g 3q)hm 93îg C 0 2-nb i530nb 6i0i6bb3 (nb. oii3o
2 .3 .1) gi ibn9n0i^3&nb 5i60ij,36nb b33unqjnj)3hm6ib3. Bycnb gi6ibihsn Bih9 m-
J96 n0n 63g0n 9 ibnb cnnoim3x 3hcn3x b 3 93hy3m6b bib3m&36nbi g i 309360b
9nb3g3ncn gi gngigii gi9m2>ng3630n 5i63noiih36ob n6gn3ng3i03h 9g5m9ih3-
m6ib3, 6ihj)im6nb bnb9nh3b3 g i &ih39m 3nhm&36b3, dnhncnigig Bycooi 3 ^ 3 6 -
330ymq3ibi gi imhcnjc36nb bnRjih3b3. 013 b x 3Ainm63 h9 i n^nb 33h3X "
- 125 -
66 ba 0 rt 56 ^ 0b 8nb9.iff.3to 89 rt ^ 0 G030)00 ft 060 u B 6 fo9 r)d866
6ob WUEp^, 8*b 8330)00* 9;phRnrnb bibjmiijbn *6 308360 5^3801 3nhoi63&nb

93b*b*3nb*g *6 38h*p *95n0b*8yoiqj308n 5*9oinoi3*0oib 8oih6y3nb*oi3nb bi^nhm
Byonb 11*01936016* (ybh. 2.12).
Ijbhncn 2.12 Bycnb 5*9^^3636^ 3gj3ii3hoi6i Shmp^jiyiocn&obtnsnb (9h*3*On S3,ynhnb 9nb3p3noi)

ais^ha fch 33h. 6m6i ift-1 H20
C4 6^36.^3360 3-5
C3 6xy>bm3i6n 815361*13360
9ih(53(vT<3636n 1.5 - 2
3ihynb636n 1.3 - 1.4
i»Njmcgnpn 1.5 - 2.5
afb^b^cfbnhi (sbicîtihgs) 3.6
Sbob^ffbnhi (80^313003) 1.5
CAM 9|}36bh336n 6 - 15 (30)
C3 03hj6nb6n 8(5361*13360
i3(»n3nh363(y> ,jhm3ni3eo
q)ihoimqjmcnpm3i6n 63360 1-2
'601803*10 bih^yfob
qjihoimqjTTOxy'iSn 6336(1 3-5
'>4C3^m®nCD*io 63^360 3-6
6063013160 63360 3-5
bjnnb 3^ 3]6ii 3.5
2 .2 . S A . C 0 2-flb 133W 0} a n 6 3 hiCDhfl J)3 3 6 i
bij)33bn 5n3onn3h3636ncn 3'bh36330yoiqj* bb3*g*bb3* ftbocn 3mj8 3036b qjm^m-

bn6oi3'bbi g* b36oij3*'b3. R33XD^n3' 60*9*580 *h 3303R633* dohnoi*g b*333d
60301031136*01* b3hnm b*i33& 6030103^13-
6*01* b3Cfn363h*g 83^*6001. 3o63h*03hn 6030*^36360 801^8303636 6*b9 nh6 *gnb
83^*6mon'b8ob 06^36^3016*^3 hoifcoihu 383*0^0- *^333 lbhg*b* 3oihq}oi5363'b'b3
5*3036ob 83833016001.
8o63h*03ho 6o3oio3h3636ob 6ooijo3o3ho 5*3036* qjml!)Oibo6oi3'bb* g* 636-
01^3^3 5*63nho>636300* o9ncn, hm8 obo6o 9390*6 hoiôihu 0)311336^360^ *0333
305836,11360b 636*95360016*90 *6 383*0019 8016*600301636 qjoiôibo6oi3b3h 3hm-
133b3680 h0i50ih(j 9*oio *^03*^0111360 (ybh. 2.13). iboiô. hoiôihy (jO036nb* 9*
jooihmqjocob 3ho*-3hcno dohncugo $018301636^0 *3(50036x0* i5nc*.boiog36nb* 9*
eg3(1836^360b cgoih9oh36obi0i30b. 3^35^383*, hoi8 *hb36oi6b inhg*3oho g*3 my>-
9363936* 3010193680 **boiôb 833y3X rT'6*b* 9* jooihmqyxyib* (b*83*omg 50 ^n-
9*301093110 *'boi^n 3hoi 6oig jooihmcgoQ^) 6* ho630m<boi6n(3o»bcg*v!)3*h6mj*ip*'b*b
838lS330m6*b 8oihob (6*b. 2 .53). mhoimcgoibqpiyi 8300b 3631150001 3gog*hn 6*3h-
01360b - ATP, jjhooi'boi-, 336ôi‘bov 9* 33^01^0103oibo]*^360b 838*95360016*90.
- 126 -
(jbhnpn 2.13 9n63hiQh 6n3om3h3 &ioni hmpn gm(ynb<i6 o>3'b3 n (Natt, 1975; Gerwick, 1982; Marschner,
1986)
y«83m-^naan>
<336*5*1 636*36(1 p> b^ldn- yrain ^w bn 91
a «**N ^ S S S S ^
jcmhmacib^nb bdhltahnb 900)3(136- N s M pe Mg, (Mn), K , Fe,
Unicoi pi b*i6ncnbnjni ’ Zn
qjm^mjn&vîn 3hmy3bn Mn Mg, C l, (Mo) Cl
3C3ijhm63&nl> *hi6b3mhlyi Mg, P, S, Fe, Cu, ^ K , Mg

pi gm^rigrnbqjmhn(Yih36i (Ca)
C 0 2-M> qjn^Hjni ei p Mg, Mn, K, Zn
iiC3noob (yiico
sbn9ncs\136nl> ,)hi6b3mh*n. p g
bsbs936pnb bnboq'bn
6153360b 9mdhin6i ___________________________________________________ K, Cl
i h i m h j , i 6 x n cgm b gm h n g n g hm p b o»s9 b9 m6 b i i 0 3 n 6 n b (jni)p0n, i& h 3 0 1 3 3 0 3 ( ^ 6 ^ -

0n(|)36nbi gi ^bndnQ^^bnb ^hi5 b3mh^9 n; cgmbagi^îbnb g3qjnijn(yi 30n6g36i
jQmhm3 0 i b ^ 3 6 3 n i b n 9 n0 i li 3& nb ( b i j i h m b i g i b i b i 0 3 b 3 0 n ) g i 5 hm3 3 6 i 9 n, h i( j i j -
33noi3&b gni^mbnfjcrybb bb3^dbhn3 b30biyh3p 3nhm6369rnj in (6ib. 2 .54). K+,
Ca*f, Mg2+ g i Cl* fiih o i30 n :> g p g i^ iS n b (& ig:>0rodh:>33& nb) 3hf*nj3b3& 0n (Jornhm -
30ib(i)369n; K+ gi Cl" ^ ( 1 3 0 1 3 3 fibdj^n 3 3 h 3 g 3 & n b 9 m d h im 6 3 6 9 n ) g i i j 3 0 i b ^ v j j -
(iD ^ X n 9 5 n 9 3 5 3 p m 6 i tg m ^ m b n b a ^ V jh n i 3 i h i ^ 9 n. On^h ( 113 0 39 3 6 ,13 2 ^ d n h n o i i g ig
i3 l5 n0 3 6 3 C ni hm&mhu 9 3 ^>>6 rT»0 n,b 0 9 n 9 m6 i 6 n0 3 0 3 3 (19 3 6 ^ 0 ^ y n - q jijljfn h 3 6 n b 8 3 -
9 ig { ,3 6 3 0 n 6 i6 n 0 n . 9 n 6 3 h i0 3 h 6 n 30W 3h3£>i0ii 6 i] ) C 3 6 m 6 i0 sS
9 ^ 3 6 ih n b 0 n 3 h 0 3 0 V
3 ^ 3 6 3 0 3 0 3 8 3 6 ^ 0 ^ 3 hm3 m h(jn3 0 fn6 nb g i h p 3 3 3 i 0 0 3 n d 0 3 d i n9 fnJ0 3 g m b jgm h m -
qjnpnb 6 3 9 15 3 3 0 ^ 16 ^ 3 gi j0m hm 30ib<i3£>nb h n ijb 3<b 3 , b r i 0 i b i g i D O d h ib ^ lrjj,!}-
h i(b 3 ; 3 b , t)3hdmg, 3 b 3 & i n9 3 0 3 0 3 6 ) 1 3 6 ^ 1} ( 8 ^ 5 ., h ]^ i6 i). hm 0 036 n u o i3 n o im 6 i h
Bib. 233 a,b y n l^ ijjn i

Triticum aestivum-9 n ibpyjob
339(S33Co6ibi 9 1 (a) jtynhm-
qjncnb 339u33cm6 sb 3mhnb p i
(b) ibfn^nb 939li33C T'6 i b i g i
RuBP-iih6mJbnci'bnb
ijijn3nn6ib 9mhnb (Evans,
1983) Total leal nitrogen (mmol-nr2]
- 127 -
6dbaort&d<?ob anb&ufti&d 9 3 r n ^ o eoamoaâ&ob &6 (n9 Dd 866
30 40
Chlorella 0.50 mM P,
0
0 1.2 2.4 3.6 4.8 6.0
Intercellular phosphate concentration 0 10 20 30 40
[10 " iimol P-unv3] Internal COz partial pressure [Pa]
fob. S34a,b 6>jm^mtfnbn6on3'bnb n6^]6bn3m&r>b gi9nyi936xD 6 i qjmbqjmhncn 9rodih>&]&i*>]. a

86316 ] 6 i)i0®|j] 5 bh]an qjmbgmhnb ljh ]e9n gi im6(3] 6^hb(ini g i 6 ]^ro-g)m^robn6a*3tn (Senft, 1978>.
b y>30nhn C 0 2-nb ibnSnpiijnibi g i qjmbojrrihncn iih^be (bb6ih3n 0.50 mM Pj) Qi (££*9 (bWSihSn
0.05 mM Pj) am0 bhh5 ]bnb 5nhm6]63n (Larcher et al., 1989)
33Qn>6 jemhmqjnpob 0mQ3i3enb 339636n0m6i9n. OijôpOobi g i hyi6nb 6i^C3-

6m6i (16333b jgmhm'bb, hm03 c 3 ijj 8300)036^ C02-nb 33 cn3nb3&> SjiQunhmb 3(101
03bi03Qi0Q3. jQmhmognpnb Qibipn 9 o0 l533C(T>&nb dohooibg 03g3&b 6:>h3migj,36b
nb, hm0 8036^3360 ^>1155336 o6o36bo3hn bo6:>oieob b h x^g &i0my3636nb
03bidc36Qm6ib, 3.0. 3 *om 9 mj03 g 36 i fihgocobi^j^ 6(336^301^ 3 mj33g 3&>b 30b-
j^3b3bi.
fci'bois 1530^3 0n63hsp3h bi»)33& 60 30*03(136*0^. 3mJ03g3&5 3nhqjm5363n3fn6i<b3 3xnn 30360b 03mb3&oo*:>u bm(n503(x?3&i. b'bm-
,50b g3qjnuoljnbbb 3001^936^ qyno*Q36o U X ^ 3 fi333:>33 205336001, n3
ghmb, hmg3biu b'bmljnb bn^h&3 06333b b36oij3ob o6,j36bn3m6nb 0600363(^3^6
fcibhgib, h ib i(j q)mlimbo6ai3'bnb bôgac 5i0m bi3cob 333150(136:. 0r*bg33b.
50
fob. £ 5 5 >>S33nhn g j m j >m b n 5 o i 3'b n b ] 6 ib b i 9 1

iV y n b 3] 9u3X(n6 ib Smhfib. (A) C4 8)536:4133-
6ft; (B) C3 3u]6ih]]6n; (C) ]ho>6(Vt>6 ] 6n; (D)
b]0nihnexn h]2>nm6]6nb bxvjhmqjruvjhn
0 6]fiJ]6n, 8 ih ie 9 6 3 i6 ] b ]]6n , ^hmSnixn 63090-
0 100 200 300 400 i6n ^y]]6nb b ] 81536*113360 (Evans, 1989;
Total leaf nitrogen [mmol N nr2] Komer, 1989; Sage & Pearcy, 1987)
- 128 -
&68016 03^6 803666330
6ib 9nhbignb ibn9noit}ni q> ibnyyib fbho>3»

gmijrribnboi^'bnb ily ^ m b i rfbhgjbi bbm^nb 930i5330m6nb bhgnb ihmimhyn^-
pig (cgmcnQnb b 39^0(1^3 5ii6jphr>936001) nj^6g3, b i 6 i 9 qjij^mh^n ih 5ibg3bi
0iQn9n^nh363Qn (Bib. 2 .5 5 ). Bygnb &i9my363bnb 3g33ii3hm6nb 9bj,i3big 93nd-
03 &i 5i 9 mnai3iQnib ibm^nb 5 i 9 my363bnb 3<33Jii3hm6i (NUE). i j 3hoi9 i 63 -
cnnb5i6 369^ &i63ibb3i3moi qjfnlj)mbn5cn3'b9 r» i'bm^nb j>i9 my3636nb 3<33jt!)3hrn6i
(NUEPh) gi gi 5 hm33b x n itn^^nb 3<33jii3hm6i 3 hmg3j(!)n x fT'&i9 n (NUEph) [ 16 ].
q3m^mbn5oi3lb9n i'bm^nb &i9my363bnb 3<33ji!Ohmbi 93993&6inhi9 g>i9(Tinbil|)36i:
NUEph = Ph/NA(ji9 mpn •C 0 2 &’2-59-1 / 9 c 9 mc n N ^ ) (2 .10 )
bigiij ihnb qjmox}nb )nb 5i9my363611b 3<?93JtJ0h-
m6nb îSm bia^^pig ghmnb bi65hdpn3n 33hnmpnb 5 i 69 i 3Qm6 i 9 n 9np36xn
mhj)i630n 99hi0n 6n3<jin3h3bnb 6 i 9 i^b (bic^pm g 0(3*3636 'bhgnb ^hnmgnb
bi6{>hdcn3m6ib i6 3hcnn 60nb 3ihjnb q3mcri036nb qj36junm6nh36nb ^hnrngb)
iqjih93636 3 iibn 9 nonh363Q mhj>i6 ri3b9 n ibm^nb 939 (i330fT|îboii6 :
mh5i63cn 90hipn 6n3cnn3h3bnb 6i 9t^)n (j>h 90h. 9fc>l»jJ>-lC)

NUEP = ------------------------------------------------------------------------------ 2 .11 )
Rihoi3Qn ibm^n
C4 9u36ih336n, 9icncn3nb pi9 ibibnioi363^0 9 ipipn qjm^mbnBcr^^hn

3<33;jii3hm6nb 5i0m, ^1130133 <bmf)n3hcnn C3 33h330 n 9^36^3, C 0 2 9 cni5 cnj9 nb
iihj, 6 ^ 3 3 6 3 6 0 3 ^ icp633b 9i9n6i(5, hmî 9 ion ajm a^bSn i'bm^nb 930(533Qm6i
'bm9n3hni. C3 6ipibm363bb i'bm^nb j)i9my3636nb 303(101 gibiQn 3q33j^3hm6i
ibibnicn36an - i 9 nb 3m9336bi(jnib nbnb6n ^ 3 ( ^ 3 6 3 6 n9 nai, hm9 9 iOi ^ ib n ^
lj)369n i 9 3Q3936ljnb 3b3n 9ihig>3&ni. ibm^nb (J30nb 9 ip i0 bnfiJih3bou>6 3hoii9
nbn6n 6ib9nhbi9nb 9 ip i0 6 i 9 i (jbiU i(p63336. 3 3 j i 9u36ih33b9n ibm^n 6 ^ 3 9 3 -
6i jbm3n03b9 n: 9u36ih 33bn, hm930 cni qjmoi03bn 3hcn b3<bm6‘b3 83^ b i 5b uroub-
0mb36, ij>m6b333b36 cni3nb itm^b. i 9 nb 5i9m, 9n3b3gi3ig bmfcr^hcnn
bn6ai3'b3hn ij^n3mbnbi g i gibi0n 3hmg3jlin30 m6 nb, nbn6n n'bhg3bni6 i'bm^nai
Cfihnb 6nigi536'b3(j yi.
S .2 .5 .5 b ih o a n qm bFlhObflb 6 i 3 CD6 i C 0 2 -nb l33C 2)*bD
6363&hn3 3nhmb369n j>ili30m cgij^mh^bn 8(536^3^3 9^1^8393636 hnj,nh(3 qjij-

^mhoii i,m9303jbn, hm90 nb ^ 33^ 30 36 ^ 3 cgm^mbnBcr^t^hn biSib^bm h3ij(jni,
Ijbigni, J,i6bb3i393bi (3i0i3 iC ^ b x hm930 n93 qjij^mhnb 1339^)83936^3 b i 3i-
b3bm h3ij(3nnbi5i6. 9i5i0ncnig 03nd036i 5i9mgj,3b 63v5m-qjm^mbn6oi3'bnb 9i0n9-
n,jnh3630n gi m3vi)n0 i 03 hn (i3933hi^3h36nb q)ih503 bnb b3Cn03bi. 5i6icn36nb
jpbhgnbib (6ib. 2 .56 ). b363bhn3 3nhm63b3n f)i6bi'b(p3h30 qjij^mhoii Jjni0303i-
b3hn 6i3036i, (jbigni, J)i6bb3i393bi 0ibmhilynhn369n 9 np3&xn 9n6iU303bnb-
ij>i6. 9m66i(53a36nb gngn j>i6663311b j>i3 n, g36j(5nm6i03hn au>3nb363h36i6n
3<3hm h3iQnblj)3hig 6ih9fn53ng53bi gni5hi33bbi g i J,hiq3n2)3b9n, 3ngh3 biSîgn
- 129 -
66b3flrt&<^ 0l> 3n b 85(*>3&6 ^ 83rt 6«=J0 6030)0063b0b P 6fn9n d 866
1000 |imoi photons-m'2'S‘1
E
3
10 15 20 25 30 35 40 30 40
Leaf temperature [C°] Shoot temperature [C°l
6 ab. ( 3 6 gjr\)mbn6oio'bnb m3,yi0] 0 nb U3C<XX)&> 3ter«l> hoemunnb 'bhpibcrufi ip30nh0n

(Hall, 1979; Grabherr & Cemusea, 1977)
100
%
(fl
55
| 80
c
>*
l/>
I 60
I
1 40
5<D
cr
20
0 200 400 600 0 800 1600

i— k
irradiance [jim d-m z s ’ ]
t -i -?ji 2
m o l- m * - s '* )
65b. SL57 63^m-q)mlyT>bn6cn3<bnb bo6^cn03‘b 3 pi>6m yip3&X3^ d^^rnhm cra^n-^h

3nhm&3&9n. 6^1^36^ \râv\: ( • ) (nh^bc^dn 9^3, 6m^nm ib^hn gi (0) 9Vi^5n 9(^3, 09hsqo 5>3hn
(Polster & Neuwirth, 1958). 3^13336^ tjhdom: Chamaecyparis obtusa-b 6olyn^9fno>mbn6ori3<bob
bn6>cn(V>b 9*1390 (Negisi, 1966)
bibnoi ijbhnoD6 9 n (6ib. 2.57).

^ “bcnb 01536^380 &56b i‘b(j>3h i336 5 ^ 3 gb 9 ngi 3nhm6 ] 6n gc’j-c fi^ jh n .
b^'bmB^hn g i i 3 n5gtb3 gaJJpiyigs&xn U3pnp3&ioii qjmB'b;). hm£,mh(j B^bn, hmQ^-
- 130 -
&68(J)6 0 3 ^ 6 8 (3 0 6 6 6 0 9 0
Cn3 ] qjij^h o yn 330oi3ob 0 o6 o0300 m pi 6î3DDCn ehmnoi 'b p x ^ b C 0 2-nb

3 on 6oij0 ib. 63636(103 3ohm63&9o 5ih30m gjij^mh36n ©bmgrng 003010119 g i
abmgng bi60mj)03 ghmob 5 i 63 i 30Pi6i 0 oi o3g36ig 630biyh3QO g i n0036ig tph-
5 ig 33cru>6 i 6 ym5 n0 n, hm3 q3(nli|iTibo6oi3<b3 i 3nbp6 nmb 30ipo3bn 96 r>036301^1616.
bb33igfTig, C 0 2-nb 9 on 6 oiJ0 nb 0 iJjo 0 i 03ho gcî^hn 06083630(116360 9 ijbn 3 i 03 hn
q3m^mbn6oi3lbîn ij^03(Ti6ob 0bm0r»g 70-80 %-b 013 0319536b.
di0m6 (101300 6 ih9 m8 m6 ob 0^1303611 63tjrr»-q3rri,jrnbo6ai3'bob 03ig(j’nb 933(13-
bm, (11615 516613301(136001 68ohig 195000 i^ b b3 ©(53611133600, 519163(136061
g i gngn 101(101^03606 6i3ib3brrig, n0 ghmb, (1019361(5 5i6ioi3&30n6i 0 ij 6 o0 i 03 hoi,
6153360 063(1361, qjmoiQnb 3056001 C02-nb ^016(536^(11(501 (Cj) o'bhg3bi
(q3rn^mbn6ai3'bnb ihi&i5o60o3hn n63n6nh3&nb 01633636300) gi q jm ^ b n b ^ ^ i II
q3fTi^mjn9n3hn 1^030161 ^3300^361. 10133 ghmb, qjoioi03&nb 6ygo6 3oi£)6(5oi0o
b3 nhig 315301 (nb. 61b. 2 .50 ). 03igpnb g33h3bni 030g3j,n bli)h3b3&nb 30186061(50-
nb 03g3{,ni: dgo3hn bn6ioi03 (0301,1)01063060(1361), 6y0ob 3ihymcgnam 6i0i6bo
(63(1501(106 gn530i), bnoi63hn bljh3bn gi '601503(101 030016333100 - 0301010360016
ibn0o0i^36ob 0363036300 51019361.
C 0 2_nb 0on6 oij0 ob 0O0<>yih3&i 63636(103 3nhm&3&9n

bri0n3ho lj)3033hi(j3hnb 30(10163600 (hiy gi0ibiboioi3630oi "boi0o3ho
L>ihli)y0nboi3ob) C 0 2 -ob 1600001(5006 0oii3ih 0 i 0 n0 oôh36 x qjij^mhig, 330(133-
03b ym3gobi, bo6ioi0nb 93030(50^0 5333006361, hi(j ph3&0ni6m6noi 16 0”bob
gj,mdnb gi6ipo 33016001 ihob 51801633300 (ybh. 2 . 14 ). brn0O3h 30O0 i^ 3h <bm6 i 9 o
53016 030mg5m0i'b3, 'biSoiihbi g i 5i'biq3b3o'b3 113033(11^3(11 bij)0img gi6igo i
g i 3 nbn 9oij03g36o6 03 g 36 i 9 0 ih 19063163 015361(133600 060036300136^ 315301
6ib0nh6igob 0301306361. 60flb 10 33hnmg9o 0301010100(53360 03hJ6oi6o 0(5361(133-
6n dnhnoiigig 03010101051(5336003601 gi 613036:^ (13150(13636. 16301
Ijbhncn 2.14 3i6n*i*nb ih ib x ^ iy h x n <3*ljPih;)&nb <t>06;,3C06nb 83935^9 C02-nb 330i3nb36nb

e^toncn^&i (%-3n Sjbidc^bQQn 3 ijbn 330n0i 5 ) ’bhgnb Sjhnmgnb J>i69i3pfn6i9n ( a Negjsi, 1966; b
Schulze, 1970; c Schulze & Hall, 1982; d Komer, 1982)
Pinus dernif Fagussylva Prums armeni- Hammock sco- Carex curvula

ajyfcjmbft lorti (sbscfcsth- tica (*fnb oca ( * 0930° t » i 66mb (ax^dcnob
63 3U3bih3)a 3(>36ih3>b h36fim6(i)c C4 % ] 6ih 3)c a\j;j&:*3 )d
6069001 gi/i6
ph^6pn>6m6ncn
- 22 - 38 - 13 - 20 - 39
5i3ni633xn bn6iciQnb
eoqwiisn
gi&ipn 16 0bpipo
- 15 -3 - 7 - 12 - 8
M âhstfjhi
COj-ob bi33i(;m gtpo^hn

Sjoijob^bob 9^^33001361
(%-9n d3bid03&3on - 37 - 43 - 32 - 60 - 47
3ijbn333n
916 )
- 131 -
6 6 b a o rt& 6 m 8 nb&>6 0 to <$6 a a ^ ^ o 603moocnoa>ob P^ândae^
[MPa]
-10 c
M arch Ap«il May June July Aug Sept
6ib . 2.58 3g*&6mb 9t536*h33d9n CC^-fri'bcni U30nb b3'bm63hn pn6 *3 n^* (33hijny>C3hn b*'b3&n).
Hammada scoparia C4 bjfijn* 0(jnh3 h*mg36oi6nb gjmoipjbnoi. qjm,5mbn6o)3'bnb 36*hnb d jr& j 8nbn
863*63 60*9*5nb R*(nh8*3]6363ns 6*6c*j,36xn- Zygophyllum dumosum j3*(pmh(pn*6
biSbh^cnnb 3jb3m'bnun3&'b3 9s b6mh j3n*6 *950036^13 Sm'bshpn C3 3355 d^Rjn*. 99h*p b3'b*n63&3 n
Zygophyllum dumosum-b 1^5303* qjrnonoj&n; (?3hoi3&nb 6yonb hjyidn (>j6J)nhn bi'bn) dpn3h
3d*cghg3&s; 9jjnh936* qjoilynbn6cn3‘bn 9* 1x53036* C02-nb 6*o*6bn. Hammada scoparia-b d*Q3db
60*9*509*6 33n6fn3mb b*j>9*hnbn 6y*Qn oi3no) dpnshn 63*033&nb phmbiy; *9 i\jo>3 9nbn 6ypnb
h3gn8n *h nonhj,3636* 9* (1^36* b*33*cm 90163133; 863*63 C3hm3&n dpn3hn bn39 h*<v>b 3nhm6369n(j
in *5h d x 3* 3® C02-nb *bn9np*^nsb 9* 3nbn COj-nb 6*p*6bn >h nh(j>333> (Lange et al., 1975 ).
*6scmf,n3hn 90)6*1533360 b93C0i*33*'b(j>330tnb 9*h*9963*63 9* qjmo)(vn3>6n b33&nb*oi3nb nb.
Tretiach ( 1993 )
tim6 b3 o j,i00n0 cndnp :>95op363nu y i 3 iC’*C dipn^6 0(50(13 6 ^ 3 0

6b ifoon. Q^fnh] 9bhr>3 , ^hm^njj^bbb Q i b^&^hmSn^&Sn bncnbmb, hma^pny
36 n3363 pm3b6 hmpb ao 3 h3 m&b C 0 2-nb 3aib6oij3 nb pn0 n^nh36 b9 n, 33 ndp3bh
5 ih*,333c n 06n93630m6i 3jm6g3b b3C3:JunnbMJi3nb. bhym330Oi:>mg u6n6npni, hm3
Bypnb 9350(50^0 6ih3mb9g,36b CC^-nb hbo3oQhi5nob 3^0000^0(13630 30603363-
pm3i63b 5 ih 30 m a p j^ m h b . 0 ihig 063 b63 &3fij6ihnb COj-nb 15300b 9pn3hn pn6:>-
0 ojjob 03b6b3pi0 ^6336^, hm0 63j)m-q3m^mbn6cn3'bob gpnb 0hjbo030n l/ 5 -gs6 2 / 3 -
093 6bj)036ni 53 *03obib, 3119(13 63 n0 nh6 b3'bm63 n. C 0 2-ob 03 oi3ob36 ob 0bj33-
ncji3bnb 0^6336363911 03hy3m6b jjih6mjbnonh36nb ^n3 ob (€3,04, CAM), 3 u36 :>hnb
'bhgnb g n h 0 nbi 9$ 3 boio b^yoago^ho 05hd6m63Qm6nb, &*h30m 3nhm&3&nb
(j>b6bi2)3cnh36001 6ycnb hbmg36m6i g i 6nigig,nb ph3 b 033636311 3 nbn 3obh63Qm-
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6i0i6bn. jyi'bou) 15300b o6^36bn3m6ibon6 3*ioiig, 050 gi8riij)Og3630ni 15*130133
q3mlj|fnbn6oi3'b3hig ij^n^hn mh5i6n36nb 0ibnb 83031*^361^3 863601^130 jbm-
300360b 8ibiboii6 (bohÔDhX10 qjiJ^nhn) g i 6cob 5 i 66 i 30ci6 i 3 o iboSoonjo-
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g iii h ^ x 0 6 ib3 nh6 ign.
bi C i 3X n ojnioiQnb C 0 2-nb 15300b 6i0i6bn 1633636b, hm3 q)nioml,o6oi3<bnb
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b36oij3noin 1^00311161. oii6iq3ih0m&i i 3 mh 8ifi3363630b 3 mhob 5i3mobio36i
hmftmhis gmoioob 803*1 6 ib 3 nh6 ignb 5i8my3636ob 3qj3jo3hm6i (CUE):
CUE = Ph/R (n8m0O CO 2a-260-1 / n8m0O C O ^ ^ 1 ) ( 2 .12 )
b30b iyh x 50*1^63680 C3 815361*13011 cgmoi036n noi3nb3636 0iib0m3&ooi 3-5-

53h 830 C 0 2, 3ngh3 i>ih5i336 906080011503*1 3hmi53b363o n3i33 ghmnb
51681301*16180, C4 8i536ih336n 30, hig6i6 8ioi ih ibibnioi36oi q}momb;)6oij3i
gi, 3i3ibigi93, 3 iboii6 gi3i38oh3630o C 0 2-nb 016131*150 - 10-20-33*1 830b. 6yi0-
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bi gi 10-b 3 mhob.
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i 3^rnohr,iq330 (8iib o 8nooh36x ) g i SfWOhmohmqnc (8igobo8o0oh3& x ) 6160036b
3 mhob. gmly ribo5oi3'b3hig 3hmg3jonX n 863163 6i6n036ob 6000 51630*11^16363001
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51*138™ qgij^hon 513036001. 63*1^6016 615361*13011 1*11863163 6om8ibob 8608363-
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30b 63*1^160, 8i5i0ooiig, 8301513b 663601^130 jbri3o036ob 8bm0mg 8%-b [2 11 ].
ibo8o0n5o3ho S^homgnb bi65hd0O3r>6i

5101869330 cgyJomhb C 0 2-ob cgnjbn50obioi3ob 6ih8mg536b ghmob ob 8150*13
6^161333010, hng3bn5 33bid03630Oi C 0 2-ob 830i3ob36ob 6icpi0O bnfijih3. 3b
3316^36300 800(^6331 gpob biioi363o, 600b 08 33homg3o, hmg3bii5 8(5361*13
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53103001. gtpnb 5168130^ 6 i 3 o 3 on6oij8 x n C 0 2-nb bi3hoim 318b 3691
5i8mii0gDb cgnoi0ob 8n3h pi8nb 5168130016180 5i0mymq)O0O C 0 2. 60(^36300
himg36m6i 6ih8piig536b 24 buoiob 5166130(116180 qgriofTibn6oi3'b8o 8 mb8 ih 36 x
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(1) 3 bpi 0 ni gm^mbn6cn3(bnb bnfijbh^;
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Day number
Bib. 2.59 bnSojh bihljy 3o3 n (isbjjgnb 801360, mh3&m6rvb 3 ^|>nb bi6:>inhm tynco) dm'bah^n
Pseudotsuga menziesii-b qjmiynbn6oio'bnl> Bcn^hn Sh^gnb (0 ) dogjh36> 8nbn 3nHJ36(jn3hn 80161(5383-
6nb&i6 836636(10 8mp30Ois6 ( • ) , hm33o3nu ?>i8 m h rn jb x ''i Oapso*1 >J3833h i£ |h 36n b i 8* bn88hscnb
b^Smâ39361 (Emmingham & Waring, 1977). y330ib3 ie h ] X n 8*>6 *3l*i 8<,'e 3Cn
9^16x53636^3 Qiyhp6m6ncn 3j ) 86 noni Stalfelt ( 1937) Snjh
6003(10 bi£i6bn bg3bi b x D<53hm gi jqjhm Qig^bnonn, on] 9uQ6ihp36b biSî-

036b ^301 0ru»(p6nm6 gng i6, ypi3X 933oib333i3n, 9^936001 ge'3-pi03h bic>6bb.
360336x^3^60 6b0i(in b 0n(p36> S jb id o ^ b x n * ‘bmSn^h gp 3-p>i03hn 6iQ i6bnb
9o3cnb333i 0n(j, o rj ibr>0ocibr»nbi0i3nb b x b iy h x ^ Sohnmgn b id in g &hdxf^>.
h ib itf igbnon i j j b onftncn, b36^hfn3r» ix (jhm SnijX iS36oi6 h^^nmBabSn. nj,
b ig iu ibndngiurvjhn ij^n sm b i 33bbd03b3QOi 3bm0mg 8 x ^ 3 6 0 0 1 0ni!)C3 3ahn-
mgnb 5b63i 3Qm6i 0n, 6i 3i^ n Siyihî, d n ^ b x ^ ^ e 3u36ih 30i i SxpxX1 cgpvyv-
bn6oi3<b]hn ij^03m bftb g i 3icn 3b<n 803(1 ibn3nQi636nb &i0my3636nb 0 ip ic n
3q)3J^3hm6nbi.
b3 xD<J>ftb 0(536ih336nbiO)3nb 3hmg3jly ix n ^fonsmftnb bi65hdpn3ri5 i ribhg^-
bi 033g3{,n 0136808933(11*16001 du36ihnb ‘bhgnbi g i ion0i£)hn 3nhrt&3&nb
03bi^93nbig:
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bn6cn3<bnh3b36 001300 6pob 5i 63i 30ai6i 3n ( 3i{,., 03hj6ni6n 01536^3360 i6
- 135 -
eabaofn&ôb a m a ^ o ^ a a r n ^ o Goaainofia&ou E ^ a n d a e ^
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63360);
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600b o8 b3<bm6b, hoig3b*i5 3303010100060 *ho*6 (<301010108153360 ^ynb d330»yD);
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5 * 68 * 30 m6 * 3 o 5hd30 *h*b30b *y h x 33homg36b 3 mhob;
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b303ji5nnb*b. 5 hd30o b30b*yh30 O 33hooigob 9 joi63 h35om6363 n 9 *hu303X n i 3C"
(5311360b 8mb*3*0n 33od036* 5*o“b*hgmb *3 33hnoigob (3(139*9 5*8my3636ooi. *3 ob
3 o(p633 * 33 b*d03630 O* 0 oij,*03 hm j,0 o3 *o3ho 3ohm636 ob 'b3blj*g 33b*oy3obo
<3360100^3(10 6o3*6-oi3ob3636nb 83(^33001, 8 *3 o6 hmg3b*i5 “bhgobioôb b*68mj)-
03 b30 b*yh30 O 33hooignb h35om6363o 8 *h(j303X n iXv5D^36nb b303j(5O* 369*
6*h8*hoimb 8*p*0o <30100^06013133(10 36*hob* g* cgml>|oi*bo3o0io36ob 350160180-
3ho 3(j nOn<t)*ljnob 3o3*hoi3036ooi.
C 0 2-nb 6*Q*6bn 8moi5*3b gjmcnQjdnb dsbnb W L (W *hob 6016*)
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(d) 5*801^036300 0301010360b (R-l ), ycmhli)36ob* (y3*3O036nb* g* 6*ymcg36nb
6*0130001) (Rs ) g* q}3b336ob (Rr ) b36oij3* [ 133 ]:
C 0 2 6*0*6bo = WLS d Phb - W L S R l - WsS Rs - WRS Rr (2 . 13 )
o8nb*oi3nb, hm3 5*6nb*'bp3hmb 3i536*hnb 5*t>oi* 15300b bh30n 6*0*6bo, C 0 2-

ob [530* 5*'bm8o0 369* oj63b 8(536*hnb 8o6ob“b3g* 6*6003660 l5*0-tP0i3- 8*5*-
0001*9, b33&9n: ^3663(10100, 3*hjob 8056001 9* 3nb 0 ^ 3^ 33, (?3hoi363o. 8n6nb'b3-
9* 6*6n03&ob b36oij3* 369* 5*ntm8mb 801390 60nb 5* 68 * 30m6 *3 n gtpobnai g*
(p*3 noi. 5*hg* *8nb*, 6*6b*t>p3h30 369* oj&3b l5*pj)3X oih5*6moi* 8*b36n. h35-
ob^hoh36xo 369* oymb nb3om 5*h30m qj*;)(501(136015, h01501(1015** gmoi036nb
bn*b0m33b 3*3hob 0360*6016*, qjmoi036nb 03933^ 03^*, O D ^ ^ i^ h o b 153000363-
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90)^0660 800 G6 rt 0li G6ba 0A 56^ 0b to ^ G b O
N D J F M A M J J A S O N
Bib. 2.60 ijynb tjg i bi<b(3'3bh'b3 Pinus cembrab ip0m63(5363&nb COj-nb 6pfr]hn 6ici6bn. biSoiihSn
gtpn^hn t»ipi6bn, hmyrihu S^bn, 63^ ^ 03^101, 16 , îjoiQb 338016333130 , 63pnb (Tranquillini,
1959). bb3igibb3i ibiyib 9hi33QO bob C02-nb 6p03hn 6i(?i6bnb 90161(539360 nb. Tselniker(1993)
6n rDhmbs gi <3D^3D^>fib bn3b0m33b, Byonb 83315330^3 Qi> 1*3. 6ib. 2.59-<bQ

0mi539xru> bo 3u363hnb C 0 2-ob Bpn^hn 63C36bnb ^ninjhn 835300010.
Bgn^hn 630i6bo S^ndcQfei jâmnb^mb 351130133 hm5mhu 8010113611 9u36bhnb
6ib9 nh6sgnb 5 3 9 ^ 3 6 3 6 ^ 3q}3j^3hm6> (CUEp[), hmaQQny 0^1536300 03bfib 3htn-
93jl3nnb 3<30 ji!0 hm&nb h3m036 n»6 ho3n 5 i'br'i33ob b3933036ib ndp33^:
CUEP1 = SPhjj +ER/ S R (&h C 02 5b'1 t ) 2 .14)
5336531100363 y g y jftjb s g(p3-(j<383h &30i 6bb g i &i6bibn0330O 33homgob

bi55hdcn3m6ib (t); idn^md ibQOwi 530^101303611 5hd30g 3&3 1138936083 >(3ohogi6
603393. 3(,H53030 3n036(!(9n qjmonvib 6ib3oh6igob 630i6bob5i6 (CUEl) 536b-
633336001, CUEpj 0 o 3b ig i£ 36 i 0(5363hob 636ob0o3h mh536mb 93 38936 39 0 mo-
1533b ghmnb 0m63i33cnb, hm030fii5 530633636b b3033(yn 3ohm636b 93 3h3 80^133-
8300 8r'i836ljnb 3380^333001 bn^^nib. 30O|yn0 CUEp] 63h0r*i3g536b nô^hiooh
83^3363631^ g(nnb3 93 p 30nb, <bhgnb3 g3 0mb33636ob 33hopi036obioi3ob; 050
d30O35 06o3363ori3i6oi 8otooi6o 8(j363tinb (93 3(13 3bm0mg 0obo 863363 636-
n036nb) C0 2-nb 63036bob 33biO)ib360ig.
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83hobj)303) 03cg3hg36i 863363 Jbm3n036ob 8ibibcni6 3 ^ 3 6 3 3 0 ^ 0 ^ ^ 5 3 9
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(3.0. C 0 2-nb 603330°^ 6 ^0 ^ ) , 'bhgnb 0U36ihnb 33 hipn 6n30in3h3&nb Qihifcb gi
03ndp36i j,i 9 my363& x n feb bhgnb 3hm(53b369n i.6 0ihij,nb ii303CH5nnbioi3nb.
C02-nb 6n3j3,jnb 6b0i(jbb g i 09 hign 6n3cnn3h3&nb 6305330^161^ g,igng3&ib
9 mhnb 3nhgi5nhn gi &ibm03ib gij330g3&ih363Qn jji39nhni (6ib. 2 .61 ). 99hipn
6n3cnn3h36nb îbhgnb bnfijih3 9 ii iX rn^)*J C 0 2-nb 03oi3nb3&nb j)ibhgibau>6
Relative growth rate [g-g'1 d'1]
6sb. i 61 C3 6iQibm3i6 9^36^33630 (Bjh^jvj&n, bSmhn bi'bn), Salix aquatica-dn (Bjh^ncjdn,

b6mhn bi'bn) gi liqen Cetraria nivalis-9n (3j 6J^nhn bi'bn) 6^rn-0)ml5mbn6cri3'bnbi (hm93(yn3
îbmSnprii m3lyi0ip3h SnhmbjfeSn) 91 03hscn 9ibnb (qjmoiQnb qjihoim&nb 3h0133006 6ro5i ULR)
9i^36ib 3mhnb imh3Qi(jn3hn gi9myigodx3^;> (Karenlampi et al., 1975; Kernings, 1990). ynhxiisni
■bhgnb bnbBhsoj^bi gi qjrfbnmcmyvjh gi gmhqjmemfcn^h Shm^3636b 9mhnb nb. Walters et al. (1993).
imhxitin^hn gi9m>)ng3d3e3&i qjm^mbnBcij'bbi gi b6hiO) gi 6]pi 9m'bihgn b;)3&nb thgib 9mhnb
nb. Pereira (1994)
3hcnig gi, 9i9ibigi03, j>mh3pnh36b cgm^mbnBcr^bnb i jtjr>3iTi6ibcni6.

C4 9u36ih336nb, 0ij,., 9 ijhnb p3h69 nb, bn9 n5gnb, <3363nb ^ h m g jj^ x ^ ^ i 2-
3-33h 3qjhm 9 i(T>iQni, 3ngh3 C 3 0^36ih336nbi (0 i&., 3 ijhnb îhbgnb, nm63nb g i
oii06 ijrnb).
ibn0ngi(3nnb 3hmg3j^36nb bihjb3 0u36ih3Oii 0ibnb 9 il>)36i (3hng3jynnb
bnfijih3, PR) i63 3hmg3j(jn3(VTi6 i îQmnbi^î 3hmg3junnb 33hnmgnb g>i60 i 3Qm-
6 i 9 n ghmnb 3hoi33c 9 n (g(P3, isnhi) 99 hipn 6n30in3h3&nb 0^36001.
PR = d W/dt (2 . 1 5 )
IPC-bODC0 0 l536ihnbioi3nb 3hmg3j(jnnb bnRjih3 03ndg36i 5 i 0 mbi(!)30 nj63b

hmj,mh(3 bhgnb qjihgmbnotn bnRjih3 (RGR) [2 3 ] i6 hmftmhy cgmcncnb 3hcn33Qnb
- 138 -
80)^0660 80866 <n0 b 6 ^ 30 ^ 66^01 66^66U0
bhgnb bnRjsh] (ULR) [2 4 ], hi^ "63ôi-ibn9n0i(3nnb bnfijshnb" (NAR) [81 ]

bn6m6n0ni. 3b ^3h0n636n g i 915636360 bhgnb Shm^bnb i6i0nbngi6,
bigi^ 9 ibnb i£>36nb 3b 830101gn 5^6010133636^, 36oii3h3big, 8ihu30m3i6n $30-
^3h3&nbi gi 63 0U36ih3Oii 63h536nb 3hmg3j^n30 Oi6 nb 93 bib 6 i 30ig.
qyshgmdncnn bhgnb bnRjshj 5^ 60 16^ 3^ ghrnnb 3hoi33C0n 93 hipn 6n3om-
3h36nb 9 i# ) 6 ib (DM) 6tf36ihnb biBynbn 80 hipn 6m6nb (W) 9 n9 ihcn:
P R i6 3 RGR = dW/dt • 1/W (5h mh5. DM - 5 ^ ' t 1 ] (2 .16 )
qjmcnpnb jhcnjjpnb bnfijshj (ULR) 5i0oinbi^36i hm5mh(3 99hi0n

6n3cnn3h36nb 6 i 9 i^n qgmoignb 801300 n8 b3g^3nhnb (A) 8n8ihoi, (1018300(5
3ib 3bnb05363gni i8 0 i(i)36:^ 3:
PR ^63 ULR=dW/dt- 1/A [5(1 01(15. DM- g0 -2 r ] ] (2 . 17 )
(2 . 17 ) cgmh030>9n n5xnbb836i, (1018 89 hi0 n 6n3oin3h36nb Si^ b n b 5^6-

8 i 30 oi6 i 9 n cgmcnpnb qjihcnn X3CDCn hfi3&i. fi3 3 X 3 ^ ln3 3^ ^ 3 ^ ^hnb. i8n^-
m0 bhgnb b^b^n i6ignbn bi^nhm36b gn6i8nj)3h cgoih8x^, hm03Q0n(j 8b3g33-
0oi6i9n 9 n(p363005 qjoioignb b^giSnhnb 6 ^ 3 6 ^ bhgi. i6iQn^n^hn 8oig3gn bi-
nd033i 653630301 $oi6!j03jb3h 3(10103(101801^83036^ bhgnb 360015363(1
gi 35bm5363h bb3igibb3i qjij^mhb 9 mhnb.
ULR 6ih9mig536b 0i536ih3Oii bib3oi&3&nbi g i bib3bb3ioi&36ob 3(10^3^03-
0m6nb bibm9b 0 ioin n6gn3ng3ig3hn 5i63nou>h36nb 5 i 69 i 30O)6 i 9 n n6 5i6-
bib(j'3hx 5ih30oi 3nhoi6369n, (1018036015 801153630 33hnoig9n ^>(1601636. U LR
5i6bii30ih36n0i S i^ ig n i n6^36bn3hn bhgnb qjibi 9 n g i i 8 nôi9 i8 ghoib 5iboi-
9n0n 3hmg3j^n30m6 nb 9ifi3363&30n S^ndgî 03gihg3b 9 boigoig bhgnb 93bi-
6i8nb ogibiSn 8 n p 36 x 8 ^336360 36^ 3hoig3j (jn30 oi6 nb oi3ipbibhnbnoi 8(53-
6ih30ii 353030b gibibnioi36nbicn3nb 53(500363005 3015010301 ULR-nb 8ijbo8i03ho
86n03630m6i bhgnb 8oii3ihn 33hnoignbioi3nb (n6^36bn3hn bhgnb 33(100190) g i
9 nbn bi93igm 86n93630oi6i ibn0n0i(jnnb 0oi30 n ^hnmgnbioôb. (101501(115 6mb-
i0mg630n nyoi, 6i0iboi3i6 9|536ih336b, 5i6b5j>3cnh36001 C4 ^nSnbib g i qjihoioig
6^1530 3630 C3 9(536ih336b bhgnb 30ip03^n bnfijih3 ibibnioi36b (tfbh. 2 .16 ).
0U36ih3Oti 5i6bb3i33630n „30i6blyilj3i5n3hn ^n336 nb“ Shoig^nnb bnfijih33&n
0^nghmg git)i39nh36xni 5 i 6 b ibp 3h x 5ih38oi 3nhoi6369n ihb3&oi6nb 93 g 36^g
55001836533630 9oihqjoi0oi5n3h g i qj36ji5noi6i03h 0^306363(136360^6. 9i5i0noiig,
b3pbbyh30 56001^3(1 5^ 3801 3nhoi6369n 8i(pi0 3hmg3j(i,n30 m6 ib gngn 86n9363-
0oi6i ij3b h3g 3hi03hn 615365(133606, 3noi63hn b33&nbi g i 65(185^360010 jjOi&-
i3h36unnb 36ihnb 8joi&3 bib3oi&3&nb bShicgn bhgnbi g i 6i3hl53036nbicn3nb.
Snhnjnoi, yn3 i6 99hi0 55(13801 3nhoi6369n i6 nj, bigiu 6nigi5n pihn6ni bij(33-
6n 6n3om3h3636noi, 9u36ih336n bibnicng36i gi6ign, 8i5hi8 h3i03hig 5i6boih-
I5n3036ign bhgnb bnfijihnoi (8ihig863i63 63^03nbi6lji6n 63fij6ihn, 6i0n9nb3&hn
i6 hoib3v!)nb36hn 8u36ih336n, b3iX36o3Z»n), hiu 8icn bi93i036ib id033b 9 nip-
6noi6 6m6ib6oihoi6ib 6 ib 9 nh6 ignb, ihinh5i630n 6n30in3h3636nbi g i Sygnb
6n333ljj9n {„b^hjbnb b ^ h s ^ n s ") [83 ],
3hoiQ3j(3nnb 5i0mbi3icn (Yp) 6ih6oiig536b 6 nm9 ibib i38 x X 5n3h 8i<J36ib gi
93 nd03&i 5i 9 oibili)X nJ63b hm5mhu n61j35hi03hn 0336jyni. 3b ihnb bb3iO*6i
bhgnb 33hnmgnb gibi6ynbbi (t1 ) gi gibibh30b (tg), m h5i6x 89 hi0 6030^3-
h3&nb hioig36m636b 9 mhnb.
- 139 -
6 ab 30 tf& «^ 0b 9 Db 8^ff)3 to 99 (n^ 0 603(J)00ff>0 &0b P 6fn9Hd 966
jjbhnpo £.16 #1^0*11316 61336:^3011 ihmgrjiyixmbnb 3ijbndie3hn pi biSx*11 9sfi3D6]6cD6n (&h

30hxyi 3ibi / 9s qjmoipnl) qjihcnn gcoSn) (9hi3icn *3<b,T'hnb 6m6ns39oii 6063931101)
a>jbna>cîn bi33Hm
au36ih3<n> 353qjn 3hmg3fon3cw6> »OTKlter>X m&;‘
k>3D&3i£>Unm >3hfime3n
C4 9i}36ihoo&n 40 - 80 20 - 30
C 3 9u36ih33&n
6hrt6jn 27 18
1bm9n3hn bih^ycnb 693cmb0ihu3C<Ti3&D6n 10 - SO 5 - 15
mhe3&6ni6n 6iQibm363&n 10 - 25 (50) 5 - 10
6icibm3i6n 3ihimb636n 14 - 18
9(33(1130 61536^33611 5 - 10
^hmSnix0 ei bjbvjhrtinEJcyi 63(1^6016360 3 - 5 1 - 2
qjmcnmc8(j336n qjihoimo}<noie<ri3i6n 63360 3 - 10 1-1.5
6063013160 633611 1 - 5 0.3 - 1
Ericaceae-b 6ih9m9ie5363Co 3331 63^360 1.5 0.5 - 1
CAM 9i536sh33&n 6 - 10 2 - 5
Y P = f a t i p R . dt (2 .18)
C 0 2-nb Bpn^hn ibipi6bnb 0b5i3big, 3hmg]junnb 5i0mbi3ipn 15(130133 Qi0 m-

i ne 3 ^ X ns 9(J36ihnb biibn9npii5nni 36ihbi gi bhgnb ^hnmgnb bi65hd(>13^16^3;
nb 2>i6fi|jQnb ^ 3 3 3 5ih39mb b3pbiyh3pn 0^ ihib3pbiyh3pn giJtbrof^nb b 35^303-
61b. i 9 g 36 ig, 3hmg3junnb 5 i 9 r'ibi3ipn 6 ih9 n ig 536b ihb36ncn himg36m6hn3 9 >-
bibnicn363cb 9u36ihnb qj36;)unm6ip3hn gibibniai36nbib, 151130133 51(136m 3nhm-
63611b, 3im0mj)n3hn e>m6nbdn3636nb 9u36ih3b3 5i3p36nb 93b6i3pnbib 16 bb3i
bibnb j,i 9 rij)3p33ioii fiiîh36nbib.
5ih30m qjij^mh^n 91^^9393636 99 hipn 6n3Cnn3h3611b 3hpig3junib3, C 0 2-nb
I53pibi q i 6 ib 9 nh6bgnb 6ipi6bb3 9mj93g36nb ftbncn. 9bihgn 5i9mbbn336nb
(n6 l|)35 bn3m6 nb bhgi^jbSmbnijnnb 5ibi65hdpn336i) 3nhm6369 n ^hm g^ni 9 ^ 3 *
pm6b; qjm^mbnBo^bnb 0b5i3big, 09 hipn 9ibnb Shmgjjunitf bibnicng3&:> ^3023(1-
i^ h ]0 n m3lin939ncn g i hm^mhu Bypnb 6ijjp36m6i, nb3 9n63hip3hn bij)33&n 6(13
cnn3h3636noi ihng3j3i^3hn 16 ihi5i6m6:>b6nih363pn 0m0ih:>536i ij33noi3&b 0ib.
0138151, 6 nm9 ibnb 5igng3&i 9 bmpmg C 0 2-nb 03cn3nb36ib3 ih ihnb giQm*><19363-
pn; i j 9609363001316 hmpb cni9 i 9 m6 b hm^mh^ 3mh9m636noi 2)m6vj)hmpnh36ign
ibn9npi(536nb f,igi3mdh:>33&i, nb3 bhgnb b33i5nqjn3hn bibnicnn. 9 ihoiipni, i 0
3hnu3b36 b 3i5 5 ih39ri qpj^m l^n 001^9303636, C 0 2-nb I530i b 3 9 mJ93 g 36 nb
9bfti3big, 9i5hi9 ihi ym33poi3nb 05033 bihnbbncn. 9i5ipnoiig, Bypnb g3qjni5n-
(jnb i6 3qjhm 93^>g bij;33&n 6n3tnn3h3636nb 6it>p36m6nbib, qjmcnpnb bhgi 3q)hm
igh3 g i dpn3hig 1)330019361, 3ngh3 C 0 2-nb 9 oii6oij9 i (6ib. 2 .6 2 ). i 9 3nhm6363n
!3hmg3Ji5nnl) 9^331101361 gngig ihnb 5i63nhm6363pn qjmonpnb ihiig3j3i^3hn
bhgncn. 6n3am3h36nb ihmgjjynnbi g i bhgnb 3hnnj3b369n fiihoi3pn bb3igibb3i
Shm^bnb ^3033(11^3(1390 9ibibniOi36p36n 96n0363pm36ig J>i6bb3i3936ni6; 3b
5i6bit>3onh36noi 36361 9 ipn9 n(i)nh363p 96 n9363pm636 b i g i m3ljn 939 nb qjih5p3&b
(6ib. 2 .6 3 ). i 9 n(|)m9 ftibnb U3pibi gi Shmg^nnb bofijih3b 9 mhnb bh^Qn
93oni6b936xfTi2>2> y<Ti330Oi3nb ih ihnb 9mbipmg6 x n-
- 140 -
80) ^ 0^60 8l33G6ft0b 6ab3n (n to m 66^ 66b0
6ib . 2.62 (gmoipnl) 'bhgnb bnb6hiqj3 g i 6;xjpv

(gm^mbn5oi3'bn 6ypnb 3n>J]5(ini(vib 3^Q6nb
05016^3 Sbjb^SbnhbSn, bm nibi g i bn3n6g3 n
( 3ijb n 3iC 3h bnb6hiqjjg 3n c9 & 3(vii 10 0 % )
(Boyer, 1970) Water potential
6*b. 2.63 (i^Si^hi^^hnb ^35*3036^

qjmlynbn5oi3'b^3 $ jn j3C 0036^33680 9ih-
U3C3&nb 5s53neri4h36nb 33hnmg3 n (Wardlaw,
1974; Wardlaw & Passioura, 1976) Temperature [°C]
2.3.3. <3niinbn6cnjbnb 3hmei*K)&nb 6ârnyo636^ q:> ‘bhenb

bflfitthD
6(536^3360 gngn himQ36mbncn 3 3 ^ 3 3 6 6ib3nh6y0;)S>b; ^QiCODb 0tf36iho;)d0n

nbn6n 83 hipn 9 ibnb 60%-'b3 8 3 ^ 93^953636. C 0 2-nb ibnSnQ^nnb 93 g 06ig 6ih-
9 mj86 nQn 6 ib3 nh6 y 03&n 369^ j,b6iBnpg3b 60130 0 u;j6 ihp8 n bnboQ&jhn g i
9mj6n0n 6'bncn l$i0 i 3X n mhîBn^bnb 9mcnbm36n03636nb gibii9iymq)n0360iQ.
i9g36ig, 5i6i6n03&i j)m6(jhm0nhg36i 9mcnbm36n03£>ncn gi 0ij>mmhgn6nh3<!>3Qn
93ji 6 n‘b036 noi, hm0xoii6i6 bm^n q)n,yTi3mh9 m63 &b S^nu^b,
2 . 3 . 3 . i <3 nt!»rnbn6 0 i 3 fenb shmeDjiM&nb 6 ^ a n d h i 3 0 6 i
C 0 2-nb ibn0n0synnb Uhmg^loO&n 6i635y33^pn3 fcigi0mdhi3g3&i 0 icnn 3hm-

g^jynnb igj,n0ngi6 i6^ „6yihrrigi6“ gmombnfScnjb^hig ijjjnoh jbm3n03&0 n, n3
- 141 -
G6b 30 fn 56«?0b anbaartO M G030103r t 3&0b P 6(f)9n d 866
jbm 3fi0p6bi ps oih556oi3&ob336, b5g5(5 nbn6n QmnbQih^bi 56 6595)155019 ^ 5 0 9 3 -

6 i (50*63336 Q^hnbtioQx #^013003600, oi3l>C369o, 65901033600, bbdbh^m jbni3n-
p3 n). 65b9oh6yo36ob 550560^53365 ^jhjgnb 9056001 g i 05om 559559500365
03<boiq)o0og56 03001300b bi(5hob3&3ho 800360b 0156660360 33h3Q36ob 3m93o 3:J-
bnbi36 “6015031101 8(536511330 3hm^m630 n boSSoihôb 93j 56o“69 ooi boih(5n3093?>i>
(35(130166360, 05jhob îhbôn). bb35 b5b3oi6369o - 35603(10 (i)h5663oih'yioi. 65b-
9nh6yc3&nb ^hiBb^mh^n 03od03&5 60111(503093601936 551130133 „6y5hmbi“ Q i
n 9m0b 0bh3£>3cb“ 9oihob 5(^36300 30161536,3(15(50300 5(150036,1)06 9o95hoi3oo6oai.
qjom 30i3n 65b3nh6yo36ob ljhb6b3mhlyi 630:1 8o0go65(i3oi&b bii5hrib6n cgnh-
030,3360b qjn0C>h5i5oob5g9n 05p 5oo 6o655c'0g35oi&ob 5^0m. 803630*359 59n bi.
1150556 63360 R333Q36ho3 d50O56 3016(536,311011363005, 5bo9o0i^36ob 6535pnb
5* g i0 mdhi336i gogo5. 550580^(1533656 b3pb 36ym6b qjooi39ob 659511333360, hm-
830015 93933016001 36(136330901090005 3^0153^0 030156569016:) cgmoiQnb 3hm-
03^,3030016560156 (65b. 2.6 4 ) g i 1135300110365 <30,501301(1801636001, 1101830015556
30600363001356363605: 53^6060, (50,1)0130606360 g5 566(5o“bob 63535.
fob. 2.64 ^honnahcngiQmineaaxD&i qjrncncyib

ojihoibi p i qjemjtJnb qjbhcnb 9mhnb (Lush et
Leal area [cm2] al. cit. Wardlaw, 1990 0nb3p3ncn)
550559500365 60111(50300365 3016(536^115(50300 5650036^001 qjm^mbo6o»3'br»b

3hmg3jôi5 bn6oi3'bob 56 9 ioin 9016000*65(5006 595000056 00 5g5o036obij)36,
b5g5(j 05Oi'b3 001016013600365 gngoi (0o“bng3o6 - s t ft o fo n n b u j 6&hjdn). b i3b Q -
6001 556300151136300 <301010360 563016063636 „g55305ymq)n0r»>6 9019685(1360360“
(301663936^360), hoi0o36n(j 0633336 05jbn0503h 0n*boQ35b: 33113x 0 35h(53Qm3-
6360b y3305<b 3 j 33g5 0301010360 565(1553636 <336300 bob(3305b, 303(101 6395 cgmcn-
0360 5bo0o05£)6noi 505(1553636 yooih^36ob “bhgob “601636b 95 556b5330ih3& nan
9353003665 95 8018600)3606 q)5<b50n 0 yoio) 6iyoiqj36b (656. 2 .65 ). 65056013^6
3511301663600 q)oy5mbn6oi3‘bnb 5bn9o05£)6n 3hoio 03010100956 656009365 bb3^>
gibb35 9oi9b05h363cb (3oi6b303hb) 9 oihob (935300360, 659010)360), g5 3ohnjncn,
- 142 -
80 ) ^ 0^60 9006 ^ 0 b 6 t>b90 rti> a m toc^GbO
after flowering
fob. 2.65 ibn3ncs^36nb 6 ih 3 mJ95 nbs g i &>6au3b3&nb ie&n03&n bmhbenb Sijôncn'b]. buhsorfb]
3^30 gj^hn Snynnoi^bb sbn3 npi1i 3dnb jpBbii^cnhjbncn pnpn mg^Bfn&noi 6ihdnrj36nb sesnoo&b, bmpm
nbhnb bnbj] _ 3ioin nfj^jBbn^hn &igi6ii53C3&nb 3n3ihcnx3&3&b (Stoy, 1965). b] 9(j36ih33&nbiCTi3nb
6^6x533360 nb. Dickson ( 1991) g i Kozlowski ( 1992)
Pisum G lycine
6ib. 2.66 sbn9nfo^36nb {>igi6i6ne3&i

qjmenc36ne>6 3m36nqj3&nb 3hm^3b3 n 9ymqj
6iymq)369n. 3^30 qghnari i(p5n35 x n i 35-
60% ibnSnpi^Tbnb 3D3u 33CC'&nbi (h iy
6sh9mj36npns *4C 6ib 3nh6:>gnb fiihoqncn),
nhndn g idl)hnb3nai - 10-35 %, 33ih 3gn6n
gid^nbsnot - 5-10%. bmpm an^cnhn
qjQhnai - 5%-tg 6:11,0360 (Bartkov &
Zvereva, 1974)
143 -
anbaarnOto a a r t ^ o 6030)0D^ 9 &0b P 6fi 9 r)d96^
U^CiDX0 9 n 3 b9 shp5 x n npo&b 5bn9no5&)6nb 653596 bb359ibb35 030101(10056

(6ib. 2.66). 58535(159, 69365 615365(1390 6030103(1365015 0m05h5536ob 556m65b-
601(1365 95 55(156^0(1365 o0ob5, hoi0 bi 3ihij,fri jboi3o036o 653636001 3^363309-
01030060 0^636056.
80100560 015365(106 35(1559 9565056636300 5563001511360650131-^ 8608363-
00135605 nb, hm9 (550333085 ^^13003685 95 0111556013685 ghmb3 8nn(pr*6 53(5003-
6300 5608005^360. Shnmhn^^bnb (530001, 5b3 30^3501, (10500101606 95(53001, <3y
b>d03 &30 n5 9 >p^Cn 9rrioibm36o03&o6 8joi63 8^5(190 01(155601360 3^5(13633(^010300
nj63b 593)35(33(10 85(155001, 9y36>hnb 86336365(13 656003681 30 ih 8oo("oi6 b>9 >-
(15501 6030*131136360. 6030103(1365015 56301 556560036590 80165600301636 <30,301-
301(10016360. 8(5365(1380 ibo8n05^36nb 5565600360b 85(1353001136300 qjobnmom-
5n3ho 83j56n'b036nb 93»5503hn 1301965 95 556653301(136001 0)36936ob, 65yoiq}36nb:>
05 b585(155(11 01(1556013606 5600005^36001 3'bh36330yoiq35 86o93630m3>6ni 85010
(133(10193^50300 365(106 5650n<bob5oi3nb. 33(1330 05(1(53001363600 03035(19365 Vigil)-
h30O 0(5365(106 89h50n 6030103(136065, (1018300(5 6565605 5(3*36300 8pib53(Yib
85(1(335080 (dmbsjQ nb n6g j j b n ) 83hy30i6b Q55b0oi36ooi 25%-056 (bn9o60nb 65b-
3bb35m&36n, ^3530) 50 %-8 g 3 (6110630, j3hn). 35(1301663600 80165300b 0693^0
03hy3m 6b 955690136001 3O % -056 (bowo) 6O%-0g3 (65(nob 0n6nm). 85(11335080
5bo8n05(536nb 55g5g36nb50i3nb b30b5yh30n 3ohoi&3&ob 93j06 noi bn9n60nb 85h-
(330360b 05bnb 93035(19365 y0mhij)36ob b53(ioim 05b5boi56 03b5d03630O5 55o‘b5h-
0mb 24% -056 47 %-093 , 959 o6 , hoi03 b5(5 6ho63oboi3nb 3b 93035(10365 0*^9365
4 3 % -e 56 57%-003 [280 ].
2 .3.3.2 . qjnoicnb Q ^tohxn e:> ane>D6i
lb h0nb5 05 00033(136(30(13606 3(101(3363606 5600005,436001 3<bh36330yoiq35 5h

‘bp3053b 00h50n 6n3oin3(i3636ob jbm3nQoi5 bl4h3^3h360n 65(10135^ 55(105 oQnbx
hm0 6563nh6y036n 065(13365 b5yh036n 6366,556(50065 95 0(5365ho6 y3305 01(1556-
300 65311010b 65690(1650013560 6016660b 93j065<b3, 06060 3‘bh36330ymqg36
0010160136003636b 03l556m03h 363h505b5 (636oij3ob5 05 006311503(10 6030103(1363-
606 lih56b3mh^nb50i3ob) 95 y3305 9565(16360 6ombo6a»3<bobiOi3nb. 5b50o cgmcn-
0nb 55630015113606501306 65^0(101 956565(130 930(^0365 5580100135001b 603im'bnb
3j 3n35036li,ob 56 65b0oh650ob (56 363(15006) 0^0160136003606 qjmh0 ooi (65b.
2 .6 7 ). b565hd0O350 0(56013(1360 95 55665330^36001 6303(1018(11(103300 0301010360
303I1P1 „d3nh59cvili363060“ 5(1056, 309(13 65680130398(56013(1360 6360560 0301010360,
85010 Q^nphmg 033h30O 301035^3(10 6^(i3^|)3hob5 95 oih55630o 6030*13(13636^
956336000 930536o0(»i6ob 559m, hoi00360n(5 9390b 6001^0803(159 8 3 ^ 0 e>oh363Qn
nb3Dio 6030^^36360, hoi50ihn(555 (300360, 90309360, 005606360 05 8301(1.590
83^560100^360. (5b50O5, 3b b505hoi0o56o5 5(1585(1^01 03010103606, 5(15839 8(5365hnb
6635 656 n036 nb, 556653301(136001 n9 6^13^3(13606 9 n05 hoi, h01830015(5 6 5 ^ 9 3 6 0
q336J(305 5^301. 580,301855, (1018 89(150 113500163680, b5g 5(5 5bn0 O05(5O5 b0 nh^g
03'bc^)030O5, y0oihj)36ob 383^3659 ,^ 3 6 3 ^ 0 5 “ . 9501 5556605 3.6. b iy h -
9360 6593 05 5(i505bo3ho 83(^560 (353^363606 Jbo0305).
03010100b b^h^tajhobi 95 60(301(36006 6565(10)00301606 9090 0h5350535hm 6i,,
(i5(5 R58oiy50o605 330103(3006 056do0 ,b3, b53b36noi 50 y5h36b 9 m65(bh3656, hm 0
- 144 -
90)^0660 9 G0 G6 rt 0U BibaOfn&ÔL 66^66li0
Dry matter Metabolic cost

13-53% Polysaccharides--------- ► 1.22 s Glucose
10-35% Proteins ------------------ ► 2.48 g Glucose
2-6% Lipids --------------------- ► 3.03 g Glucose
4-20% Lignin--------------------- *■ 2.12 g Glucose
5-26% Soluble carbohydrates *■ 1.09 g Glucose
3- 12% Organic acids ► 0.91 g Glucose
Carbon cost
Protein —
Lignin —
Tannin —
Lipids — 300 O
Ok
iii.il
E
Sugar —
Raw fibre —
B L, l_2 C
Fully differentiated leaf
&ib. ZA7 6ib 3nh6ignb gmoicnb qjmhdnhj&nb ShmujbSn (Mooney, 1972; Chapin,
1989). iSicmjnjhn 9n 6*u;)9 ;)dn (jhmSnixn b3 0u36ih336nbioi3nb nb. Reich et al. ( 1991) g i
Solbrado ( 1991)
qjmcncnb 5*6b*'bp3h30n ^nUnb g*9*b*bn*oi3630n b*9*h333&n g i, *b3 30^3*01,

"i6^nbi9ih j336n‘ 5*6fn6*b6mh3630fi* n6gn3iig3*03hn bibnym^bcm q)mh3nb* Q>
bibn^rnijbpn 3nhm&35nb 3rioibri36nc36*ois g*b*33*ymq)n036o*g. 0138(5*, 9*h*g36-
3*63 0301010360 0030*6010 5*63ooi*h36ob*oi3ob b*^rihri3636 *bo3 o0 *^ 36 ob 0 3g >
1136001 gng h*mg36m6*b; 3b b*6ri0mri 3*33n, 3oiqo*6o 8i536*hnb*oi3nb b*b*h»53-
60m*: bSnhn, dôghm 3*h3nb 33b*6*hfi36360*g 6*d3b 3b*<}ohm36* ym33060n3h*g
oi*3obo 6063360b 3*bnb 15%-ob 5*6*6036* [215]. *9ob b*3nhfib3ohpig, 013 qjmov-
0360b og36jj}om6*03ho b*boum(jb0ri ghm 3fnj)03*, b*b3<b3* 5*633fih36>go g*6*b*h-
3360 *b*0o gmoi036nb 5*6300i*h36nb*b (0138(5* ôghn 6*303611, 3ogh3 8*h*g86-
3*63 qjmoi036nb 9380*3333*30), 3*{>h*9 3b g*nd033* *b0*g5*00O0O qjpiai036ob
8*(n*0n q3m(ynbo6oi3b3ho *J»yi3m6noi* g* *(bmôb 93lj*6m0rib9nb 5*dQ03h36ooi
(nb. 6*b. 2.24). ^ 3 8 0 8m'b*hgo 3*h6y30b gsRhgopx ojrioi0363n g*&3h3&ob 3hm-
(53bo 830(^036* 5*b*65hd0O3g3b bn^m^bpob 633300 b*6j,hd0n3m6nb g**b(W)6noi
3horio 33b*83gooi g*, *8*boi*6*33, 8*ol3n 80%-ooi nbhg3&* *bn8n0*(536nb h*mQ3-
6016*15 [108].
qjmoiQob g*6*b*h336n g* 801536* 6xobb3 m6b 3qjhm 8 3^ , 3ogh3 Sbmprig
6*b8oh6*gnb 60333^. qyricnQnb bn^m^bonb b*65hdo03m6* y*gn&g36* 33rnbob-
,1,383680 8*cn 3*h^6nmh36ot*6 8301*6*69016001. 6*Q*bnb8^*330O (5bri3x:)2>ob* g*
3*h*<bn^36nb*5*6 0030b g*b*(5*3*g b*65hdQ03*g 8(jbni3h36 qjf*ioi036b 3b*^nhm-
36* 83^*6033(10 @*8(53X 3^0 &^> 33fih*gn 93l^*6mpnlj36nb gogo 3h*3*053*hfn6* (ob.
01*30 1.1.4.5), h*u 8mooibm3b g*8*^36ooi g*6*b*h336b hoi30o 6nmbo6oi3<b3&ob
5*6b*bmh(5nxD2>C^Q- b*56 mj)0 X 8lsbr*i3h36o, 6360*60, 0138(5* 8*p*0o gm^ribn6-
- 145 -
6 ^ b a o ri& ^ m anb&>ri36 d a a ^ ^ o 603U)03 ft 3 &0b ? 6rtand 36 d
OQ'b^hn iJon3fTînlj O j^D qjtnoiQpbn 6iOibnbcyi0xn ubci33Q3&nb 9 n]h 06036bn-

]h n OAD^Snb 03g3{>ig 6 iig h 3 3 ig buncQD^b g i i9 g 3 6 ig 9m&36i 96 n9 -
3 6 x (T’3 6 ie 0yn h g36 i ( 6 >b. 2 .6 8 ).
6ib. 2.60 gmoriQ^bngsB 3(j36jhnb bb3$ 6i 6np363n

Sijhnb (>|hi6b3mhijnh3&nC> giSniineo&XO^ 9bD-
6ihnb bfujmubonb biSfchdcr^miib] 54 &»dm3nb 33
hnbjj'b] (Harper, 1989) Days after sowing
2 . 3 . 3 . 3 . b ib fib n iib c n qjmhao&n 0 ^ b n a n c ^ D b n b a n b a^ h o b ^
9j336ihnb mh{p6n‘bx5nnb gm63 g i bibnum^bgrn qjmh0i 5i6b ilb(T>3hi3b ibn0n-

0iO32)nb 0mb0ih36nb hi j/bi nj63&i ^00^3636300 ^hmg^nh^bnbi gi 'bhpnbiov
30b, 0nb bfH5mubQnb36ihni6m6ib, ]>m6t)3h36unnb 3 Qbidp^&cmb^&bi g i &ih30mb
bb3igibb3i qjij^mhjbnb ^>30^1^03936^3 bi 3ib]bm hQiJijnib. 830935 3ihb5 h i-
033630 i(p63hnQfii 9(336ihnb 5 i 6 bb3i 33&30n Ju33nb hi3 g 36n03 Qijpcnoin.
3htno rrtvto S m S x ^ n b ‘b h gn lu o gn b : 3Jj3i6 b rr)h n ,y d n

3hoi33h3gni6n qjm^rrîjmohmqQon mhj>i6n'b03&n oi3nown6 3^36330 90 10936
oii3fii6oi oii3b 6ib 3 nh6 iQnoi - 0ioi ihi iJscn bb3i ^ 3 9 3 6 0 , bigi(j 603010-
3h3&iOii gi0ihij>3&ib 33bd03&36. 33h3g9o b3pbiyh30n oii6iq)ihgr'i6ii ibnSnp^Q -
6nb 9 i 3hmg3ijnh3& x 9 i°m 0m0b0ih3&x y Ti03fTi636o36b 9 mhob: 9 i5 „ Jcm -
h30i 9 n j0mhm30ibo36b 3ôhi3cn 3hmom30i'b0nb amyxruftnb oinoijdnb 6ib 33ih n .
013 i0 ib 5i3fKn3i0nb6n636cn, i(j>ih 6i&3ni3nhg36i, hm0 bij>336n 6n30i03h36;)6nCTii
g i bn6iOi0nb iihftig 9m9ihi53&nb 33hnmg0n 6yi00y36ihnb 33h3g3&b 033d0n>oi
- 146 -
30) ^ 0^60 aUDG^fnOb 6 ^b30 A 6 ^«5 nb to«=U6bO
gii6hm3m6 ibn0n0i^3&nb gngn himg36m&i, bBhiqjig 0ni(j>6nm6 QyjbnQiQ^h

bmO^bb g i 9in6ypi6 giymqji. iS j^ ih ig, i3lim(i)hmcgxn 3harjjh3gni6n mhj)i6n(b-
3n nyj6;]6b oii3nb 6ib 3 nhfeignb 6i0i^b n6gn3ngoii hnub3nb &ibhgnbioi3nb i6^
5i0hi3Q36nbiOi3nb. 6ib 9 nh6 ignb 0j>i|jhig gig^bncnn 6 ip i 5bnb 3og3&n 3m3x i l 5n-
nb bn9^nghm3nb bBhiojn b h g ii. 3n6ingi6 q}mlynbn5oi3'bnb bnRjih^bi g i g(?o-
(piOnb 5i 63i 3om6i 9 n giymqjicni hnjjb3b 0mhnb 3nhgi3nh3hn 3mhi5n x n giOmingj-
b x D îi. <3n,i)m30i5j(ifT'6nb bhgnb bnfijih] S^ndcD&i ^ m n b i^ m b hmfcmhu 3m3p-
Qilinnb bn0^nghm3nb bhgncn i6 ghmnb 3hcn33g9n 0m3bgih giym qjioii
himg^Smbncn.
6ib3 nhdignb b&iiojn 601536^: n633blyxyi3hn jytin

0l536ih3O)i n633blyi()n3hn ^nSn bibniong^bi diîgn qjm^mbnSoob^hn ifen-
3m&ncn g i qjmombnti^Vihig ij^n^hn jbr^ngj&nb jbmsngcni biQhcnm 9 ibibcni 6
3i(pi0n 33qjihg36noi (hm^mhy &jbn, 50%). bhgnb qjibiSn By^bihnb dn^h
03oi3nb36xn 6 ib 9 nh6 ign, dnhncnigig, b3ih g j 6 i ib ign qjcncng^bnb 6 ih 3 mj36ib,
hm3p36ni5 3 o3 g 36 3bh ]53xyfT'<33D6 3u36ih30n 6 ib 3 nh6 ignb 6i0i(jnb g,igng3&ib.
y3i3ngm6nb ajibiSn g i i 3 qjibnb 3 o0g 35 ibnSngiôbnb 6iîgn j,igi3hc*336i
h^Shmgjji^nxn mh5i6m3&nbi^36, 3u36ihnb bb3i 6i6ng36nb 3m3ihij>36i y» X 3"
bx^& b n3 gm630g3, hm9g36ntf Sijnh^g ic ^ i^ Q & i 9(J36ihnb ihb3&3gn Sg&roSih}-
mbnb 336ihfi:j636nb 9 moibm36ng3&3&b - b6 n3hn qjfncng36n b3 ] 6i g i j>3g 3&i. ibQ
hm0 , ajmcnp^bnb, p 3h d x n boh^JoD^D^0^ <33b33bnb> g i h35hmg]J^nxn mhjpGm^-
6nb 3hm3mhun36n 0^36ih39n 06 r»336x ,T,36ig n(j3pQ6i 0 nbn bibnymybcm (jn^pnb
5)iB0i 3grn6i 0n ( 6ib. 2 .69 ).
0y36ih3oii n633b(!(ni3n3hn o^^nb biîQOi^bm 9 i&>pncnb 3hcn6gni6n 04536^33-
6n 6ih0mig63636. 3b 0jiQ6ih336n b h xig ^ 3 6 3 6 3 6 ghmnb n3 9nj)g3 ^hnmgb,
hmg3biU &ih30m 3nhm&3&n b3pbiyhxr>i bhgnb, y3i3ngm6nbi g i 6iymq)nb
J,i0m6ibi3nbicn3nb: 9 icn n0g36ig iihîg ^6gi 2>idmny36m6 oii3ni6ain qjmom-
ibnSngiooiin, hm0 3 b h 36 3x y m6 jbm3npcni gngn himg36m6nb 5i63ncr\ih36i 3 ijbn-
3 ip]hig bi60 mi03 ghm9 n. 3hcn60ni6n 3tf36ih336n i 3 ib ib]hb3636 9 i 3 n6 i(j y»,
hrng^bi^ ghmnb bi^ 9 img gngn Sjhnmgn ihib3pbiyh3pni bhgnbicn3nb. biqjb3gnb
Days after sowing Days alter sowing
fob. 2 .69 a,b 3hoi6cfn3i6o 9u36iho6 mhi>i6fn363n 1608001^3606 5161600361. a ibri8n0:><$3&ob 51616-
6631916631 mhjpBmSn 516300^13606 qjibob 3i6dnQ'b3. b 6631916631 mh5i6m6 <J)ihgm6ncnn
'bhgi (93hipo 3i 6o6 81^36 1) (Fischer cit. Sculze, 1982 mixedviT)
- 147 -
66b30fn&«^0b 8nb&>ff.0&6 83fn6C?0 6030)0060601 P6(f)8nd866
6i0ibm3636n, 55(130133 0fb3b30fbnhi g i bb3i 3hcn60ni6n i3CvbD^D^n ndp33fii6 5^6-

bivôih 36001 gng 0mbi3iob. b30biyh30 5ih30(n 3nhm&369n ibnOnpiunnb Shmg^j-
^3600 ib30in n633b^nh3&:> 3<bh36330ymcgb h^m h^ 3b3 “bhgib, ^ 3 3 3 3b3 0mbi3-
pni6m6ib. 03mh3 0bhn3, hmg3bi(3 gniiiQjhn 3 nhm636 (i 6^36630^1/130(11, 5 ^5b>-
^301(1361101 013 Bygnb 0ihi5nb 56 6(iigi59n bi^336n 6n3om3h3636nb 6:>i03dm6ib,
0U36ih3 nd3C3630ns 5i6(i3noiihnib 3jUi36bn3hn 0336300 bnb^30i; 3b bmhunQpg-
36s q3moi036nb gihoiri6nb 330(yih3&nb bih3<b3, hibi^ ogmljmb(i6cn3'b3hn 5i0mb>
30 nb 930 j3nh36 i gi h^Shmg^jijn^Qn 3 m(j 36(jni0 nb 0^33(101361 0mbg33b.
9 m3 ^nh 63 m6 i 3biqjhoibfn36nb 9 n“b 6 noi: yn 6 b 3h 3i^ rrxyi 6 0 333^ 1

9 hi 3i 060 fii6 n 6i0ibm3i6n bib3m636n nb3 iB3bhri53636 ibn0n0i^36nb 6n^-
33i|jb, hm9 ^ 6 3 3 3 6 6 ib 9 nh6 ignb 3q)hm gi6ig ^ 390ihnjj 6 i 3 i^b gi i 0 nb 5i9m
^qghrn 6 3g i n“bhg36ni6, 309(13 n633b(5nijn3hn ^n3nb 0in63 0^36^3360. 63mh3
96(103, nbn6n n(p3336ni6 ihib3gbiyh30 5ih30m 3nhm6363nij - 15(13 gi 99hi0, i5 -
h30i33 96 nhn Smgi^nb 3nhm6369n. 9 ioin 5i63fioiih36i gibi 6 ynb9 n 3hcn60(ii636nb
0b5i3bni. 9i5hi0 3353iiil5nD*ln b^h^j^^h3611b qjmh9nh36nb 930 g 35 nbn5 n ifthm-
33636 bih3‘b3h3m Sihiftb g i 0bmpmg i0nb 930 g 35 0693636 y3i3np3&nb 0301(100(13-
6ib. “bhgnb 3nh33pn b3“bm6nb 6m0mbioi3nb ibn0n0i^36n 0R»>i0n‘b3630ni y0mh-
6369(1 g i 5ih gi i0nbi, 0u36ihnb 0n6nbb3gi 6i6n0369n, hm9036nij 93nd03&i 5 i6 -
3noiihg636 9 ibn3h bi0ihi5ri mh5i6m3&ig. y3i 3(i03 bn 3(icnihg36i 0bni0Pi0 n9 nb
039935, hi(j 9 (536ih336n gi05hm33636 06n03630m3i6 „ i i 3nlî0 b“ , hm030bx5 936-
g 359 n 09363636 bb3igibb3i 9 n“b636 nbioi3(ib.
30(1330 630b 0m 0ihi536xn ibn0n0i^36n 5i9rriny3636i 0301(13 630b y0mh^3-
6nb “bhgnbiOi3nb (6ib. 2 .70 ). 03mh3 630b n6y36i ibi0n 6n30in3h3636nb bShiqjn
bn6oi3“bn, hm030ni} bi0ihi5m 0ibi0nb ihb3&oi6ob 5i0m 5 i“biqgbx‘b3 b i ^ i m g
0i0m 3yig3630ni qgm(jnibn6oi3<b‘b3 0mj03gn q3ijli,mh36obi5i6. 013 y3i3n0oi6obicn-
%
100 •
80 -
60 ■
40 -
20 -
A M J J A S O A M J J A S O
Months Months
6sb. 2.70 ibn3nc>vJ36nb j>i6i6nco6i 3hi3ie6efn3i6 9ua6ih3o&0n (Caltha palustris). 81(1(5636.1 b^h>-
onn: ibndnci&)&nb 9^ SpD&ihjR&i bbsigibbai mh{>i6m3n. 81(133361 b^hioin: bfc>3>-
gibb3i mh&i6mb 93hion Sibnb bnpnpo (Eber, 1991) bbsi 315361(13011013(11) i6icm&n:)hn 3^33636-
(V)6n nb.: Iwaki & Midorikawa (1968), Goiyshina (1969) gi Sale (1974)
- 148 -
90) ^ 0^60 9006 ^ff)0 b b ^^G bO
6ib. 2.71 99hi0n 3ibnb 5i6i6n c36n b

ii 6m6'bn3n3h36i 6n ]haibi gi n3i 33
0t336ih ;)3n gi&>0 (600 3 ) g i 3i£>iQ
( 2600-3200 3) s& b m e;)^ bn3i(rC 0D&<ti3-
aXP^cSennb &il5nC0&nar' 3>j
ibndnQi^j&b ij,hm33i>36 3n6nbj333i
mhj>i6m363n: 93bi& i3nbig Bioin d ib iu
333gih3& cig gngni; cr>30ui mhn33
9] 3oib333i3n ojRonp]6n &i9mny363636
ibn 3nQ i^]6nb jhcnbi g i n3i 33 h im g j-
6m6ib (Komer & Reinhardt, L987)
3nb Qy^Gihnb 9bigymq)6 nb ghmb bi333&n 6n30in3h3&3&n bi>)9 im himp36m&noiii,

bbQihi^m Shm^^b360^6 S^gih^ncn înhb^^bmfti y3i3n03&bi g i 6iymqj3&b hfiQ-
6icn. 930g359n, “bhgnb b3<bm6nb gi9oii3h3&nbi|)36 sbn9npi^36n 5iging363&i 393-
Otfbig 9i536ihnb 9 n6 nb‘b 3g i 6i6n03&nbi>)36, hm003 &n(5 03bi6 i 9 nbi0 0 9 ^ 36 36
6 m6 i 9 n.
9hi3ip60ni6n 9u36ih33&n 6n30in3h3i>icni 6 ih9 mj96 nbiOi3nb 6ih9i^3&ncn
01^363636 bi69 rnj)03 b3Qbiyh3Q ^hnmgb gi 363h5n30ig ibn9 n0 nh3&36 y 3i 3ncD~
6nbi gi 6iymq336nb qjmh9nh3&nb 3'bh36330biymq}ig nb333, hmj,mh^ igh] f)i‘biq3-
b3C“b3 9 my3i 3n03 91336^(133611, hm030oii qgrncnpobnb n9 ghmnbicn3nb ih b 3& x
9 ibib ih 033dpni 3<bh;j6330ymb i 3l5r'CD^3Cn 9ibi0ncn 9(336ih3. 3b 3b3&i, 9 i&i-
pnoiig, J>i'big3b30nb jomcgn^bb, hm9o0cnij)i6 &33hn y3i3ncm6ib nBy36b 9 i 6 i 9 ,
bi 6 i 9 qgrnoi03bn j,in0036i. 9ipi00oinb 9u36ih33&9i ^5 g i giny3i3n0m6 g i 9 mib6 -
hm6 cn3b036nb 9 m96 nq)3&i bi 69 mj)03 biqjb^Qnb 5 i 69 i 30m6 i 9 n; 9 icnn qjmjjm-
bn5 cn3'b3hn 5i 9 mbi3i 0 n ihb36ncnig 033b6i3030ni. g i bmpmb, b^Snb 81536^33-
E>n crii3ni5oin bibn^m^bom (jn^nb gibh303&nbioi3nb ny363636 ghmnb 9m6ij>33cnb
'bi9 cnhnb bn|jn33bi g i 'bi<gb30nb &3i03ib 9 mhnb. y330i 3b 9i536ih3 bi^nhm3&b
bi9 ihij,m mhj,i6m3bb (0^3(1360, vi,363h36n, qj3b33&n g i &m0;)3O&n). fcihgi i 9 nbi,
ib3cnn bib3m&36n b9 nhig n3ncnih3636 3jb^36bn3h cg3b3cni bnb^39ib, 9i(pi08cnnb
bib3m&3&nb 939cnb333i9n, i^mh^bnbi g i y3i3n036nb, i5h3cn33 gmcn0nb 1J3663-
6nb bih3'b3, 0 139 ^ qjmcn03&nb 9ibnb 93<3ihg3&i bi3hcnm 9 ibibcni6 cnncnjSnb
3U303pn hR3&i (6ib. 2 .7 1 ).
9ibnb fcigngQbi 9hi3i 060Oi6m6nb bihjt^: iiD9xMyT3hn ^n5n

b3 9u36ih3 - y330i“b3 9i(Tii05i63noiih3&30n g i gngn qjmh9 i - 6 ib 9 nh6 igncn
cru>3nbn cni3nb 9 m9 ih i 53&ib i63bhri53&b oii3nbn bi6j)hd0n3n bnym^bonb
- 149 -
6 ^ 30 6 6 .^ 0 1 1 9 nb 8 i(f.aa6 9 3 6 . ^ 0 6030)03 (fi350 b P i 6 9 n d 9 G6
6sb. B.72 9oicni6 n 39hi(vi 3>bnb {>i6 i 6 -

nca6 i S^bnb bb^peibb]:. mh{,i6rn3 n
(Remezov & Bykova cit. Mitscherlich, 1970
9nb]e3ncn)
33hnmgou>6 936ym6noi. ibn9n0i&)6nb gng him036m6ib b33&n bih3i336 biyhQQSn

g i bi^hb6 b 3mh^m jbm3n036nb qjml^nhodnbicr^nb. i 9 nb ^3130036^^16^ 5 i 63nhm6 ^-
6300* bnb ihjn^j^hnoi; hxnm6369n, bê^lS biibnSnQi^nm Spn^hn ^hnmgn
bi65hdpn3ns, 3b id033b b33bb 5^9^86933^ im6 i 3h36( j x 33nhil53bm6ib 6ipibm-
3^6 01336^33601^6 03gih36ncn, hm0036n(3 cni6gioii6 nfihgn03 &rii6 9 ioi 0n3h.
bfH3rTi|3b0fib 3nh33p 60360n bnb qjmox^bnb 9ib i bbQhoim 99hi0n 9ibnb 6ib-
33ihb 03i9536b, 9:>5h:>9 bnb bhgibcni6, 9nbn brn036nb j>igng36iboii6 3hcru>g
<33fTion036nb 9ibnbi ^hrnboibB oru>6iojihgm6i 0(330365; qjnoi036nb 9ib i 396n936^-
Onig 9i lj)30m6b 9i 9n6, hmg3biu 9ou>3ihn p3hm g i ,1^ 3 6 0 b x DOjhn g i ^qjhm
0bb3n0g3&i g i 9i^ 30m6b 6m6i 9 n (6ib. 2.72). bhgibh30n bnb qjPicno36nb 9i b i
bi^hoim 9ibnb 0bm0Pig 1-5%-b 93ig& 36b (nb. ybh. 2 . 15). 9iOi }6 g i 0mi0ihij,m 6
9ibi0ncn bnb bb3i 6i6n036n, hm930oii Bm6i 633hig iQ ^ d i^ & i cn3noi qjPKnoo&nb
6m6ib. 8x 36n hmj,mh(5 3hm 93^nxfT'5}nb, nb3 bhgnb bnfijihnb oii6gicni6m6non
930(3nh36i 9n 0gj,m0ih3m6b.
33 hj6 ni 6 81536^300 9 noh bi6&hd0n3n ghmnb &i68 i 30 m6 i 9 n 5 i 9 m939 i 336 x r'
93633 &30 rr,&>6 n 6i0ibn3i6 bib3fn6o6oii6 033n6h8n J,i6m6ib6mh363(vii, brt^s^h
nbn6n 5igib6m6ni6 ^19315 gi6i0n 3hrn93j^ n x fn‘:>ncn:> Pb ibn9n0i^36nb 5i6i6r>03-
6nb 5 ihai30 D^30 n b3hbnoi. 3n6ingi6 bo 9(336ih336n ig3n0ig 33h 3 5 3 3 6 ^ 6 jphQ-
9 mb (33on03636b, j>ih38mb 939 b(3>X 3X n opji!)fT'h36nb 4^30365 9 iO«b3 ocghm
d0nohni, 3ngho 6i0ibri3636'b3. i 9 noi 36gi inbb6ib nb ojij^n, hm9 b^h^bnb
3 riJ93936 noi b33&n 3qjhm 9 i 03 np333&ni5 , 3ngh3 6i0ibm3i6n 81536^3360.
q3mairri00(3336 b33&9n 6ib3nh6y036nb 0ihij>n ^mî 3qjhm igh3 01^3936^.,

3ngho qjmor^nb &i9 pi gin6y36i; ibn9 ri0i^ 36n 9 n39ihcnodi 33b i3nh^36nbij)36,
3qjhm j,3ni6 $n 6mhRn y0mh^36nbij)36 (6ib. 2 .73 ). bi 9 ihi&m 6n3Cin3h3636nb
giib0m36noi 93bi 03gn nbih33&i qjmo^bnb 5i00ib 3, hm90369ii3 b x 3i>C3
aii3nbn 6 x n0n 3^gi 03nj)i6ni6 bb3i qjnioi036nb g i y0mhli)36nb O}mh0nh3&i9n.
9 mj)3ni636 nan ibn9 npi^36 n ^Sn h i^big y3i 3n03 &nbi g i 5i63ncnih36ign 6 iy -
mq)36nbii(36 5igi3gn636i, 030936 i i 96 n30nbi>,36 g i 6m0mb qg3b33&bi g i J3hj0n
- ib 0 ig 6 ih 9 mj96 n0n i)3nhlj36nbi g i bibi03&0nb bi9 ihi£,m biyi3360n. biy3i3nQ3
^n h ^b n b gnqj3h35uniur»i h 3 5 x nh0o6i ihi 9 ih^m 5ih3&i6n g i 9 n6 i& i 6 n
- 150 -
ItdOldiknO
6sb. 2.73 b3]63n bib^fepnb giahm33&nb bj'bmB^n (531036190161. Fagus sylvatica (i536^hi03ho
33hn3i): 3iJjn3iQ3hn gi&hm336i 61^3363601 8130 cgjhncn; goeo himp36fni>i - piS^hobsnoi; 8150(13
himg36m&i - 638,500^3160 giS^nbjncn; oiânhn qj^hn sc6n36i3b, hm8 6161836300 ih ihnb, s6
i(j0mfi36n0ns 0bmpme 8nbn iSX*1- 1 j(i'biojbx'b3 osrnoipjbnb ftiSonb 606; 2 qjmoiQOiwib 51800b
ghmb; 3 “biojbx11^ 931 S^hnmgSn; 4 838mg&m9ooi <gmonc36ob 9103361993; 5 îdonihdn yn633&rvb
ghmb. (Gaumann, 1935). Abies vetchii (oi3m6m): 1 ai’biqjbx'Ô j>3oh,>3&ob jpScnbib; 2 <
biqjbxn'J
6mcmb; 3 'biSouhSo yn63nbsb (Kimura 1969). Oleaeuropaea (jphgob ij&ob 909181*1360): 1 {^biojb-
Xt>3 isnh^jbnb e>i8cobib pi y3i3n(yn6nbib: 2 lbiq)b3Cob 831 33homg8o. 63iC3nbib; 3 3yo63fn
'bsdcnih9n 63090160 3]homgnb 938936 (Larcher & Thomaser-Thin, 1988)
b n 5 6 i0 3 6 n o i, ih i0 3 g i j b ih ^ o x hm Qb ib h x D & o B g ih R ^ S n p n b iO b h ifc m
611601031136360. bhgnb i^hnmgnb Qibibh3Qnbicn3nb nbn6n <y>Qi0mdhi3g3bi 0QhJ-
6nb jbm3np3bnbbj)36 g i fthjnbi^B, p^hmbi g i q)ob336nbi2)36, b ig iy
bi 3 ihi{)mg îging^bi (6ib. 2 .74 ). ^hmin^ban g i 99 hiQ h3j,nm63b0n 6pnb
5 i 53 i 30mbi3 n 0ihij,nb gigjhfT^bnb hi3g;)6n03 b^'bmB^h S^hnmgb j)igni6, 0i{>i-
Qncnig, Q3c>3nb m o i b ib^cn ^hnmgb frignb.
'bmSn^hn bih^yonb 3ihig0B3i63 03hj6ni6n 0(5361(133^ ih 1130011(^36 ibip
ygmh^bb (bi8oihnb 0rib3363bnb S^hnmgnb gi0aii3h3bnboii6i33, bi 6 i 3 33(1 yig33
ij30i 03hR36nQn 6n6i 6gnb 6n63]bn gi <301010360. 013 i0n6gn b^pbiyh^Qni, nbn-
6n 55615(^01636 C02-nb 03oi3nb3&ib (0130151 8(50(13 hioig36oi6noi) b i 0 anh9 n g i
igh3 j)ibi<3 b x 'b 3 - hoig3bn5 5 ibiqjb 3ob 3 i3nh^3&nb 51001 n6y3bi, i0 b6n3hn
mhj,i6m3bnb 0nQh 9oii6cnj0300i 6ib3 nh6 ig 0 i 93ndQ3bi giqjihmb 0moiboi36n-
03bnb b id in g gngn 6iBn0n, gi6ihfi36n $n 03O3b36i (313(1013661 g i qj3b33b0n ih-
^>36300 3ihij)ngi6 . 0 ihig 363 i 53 03hj6ni6n bib3mb3bn cni3nb6oin 6165(^0030
biibn0n0H5nm S^hnmgnb 51001, 6333036(103, goi0o6oh3636 <301010100(5336 bib3Pi63-
bb3, 1119516 &hd30n bi0oiihn 16 03 hi0 n biqjb30o bp3gi3b, i0oij>O3bb bhgnb
- 151 -
S^hnmgb; 9i5-, 001369(1, fihgnpni30inb ^ynb bih^yQQSn i6 h3dnm63&9n, bigi^ 633-
6nb bhgib 53bp3i t>cl3gi3b. 3 bmpmp n9 h3f,nm63&9 n, bigi^ ihib3pb:>yh3pn
b3<bm6n dipni6 9 i>^>|jhn pi biSfchdpr^ni (b3<bihj(jnj>i, iC'9 mbi3P3Cin (jnSbnhn, 6>b-
33hip3gi66m36n), cgmcnmp9^336n 63360 g i 63^36(1 i3p i3 3h3gm8n6i6(j3h6n 693-
6ni6.
^ February j
6sb. 2.74 Acer saccharum-cb

q)3b39n bibici^&onb g:>5hm3j-
6 i 6pnb J,i 60 i 30n 6i 9 n. Jbm-
30^360, bigxi bibiSyiQQni
9ij)hfn3ncn ip 5n96x ni 3i 3n
aj^hnoi (Wargo, 1979)
2.4. ai536^h3Dcnon^6^(bn6ên3bD6nb
6^b9nhbênb b rao w i
2.4.1. 0l5D62)hDDCfl b ^ h flb 3hne3jv!»n3Cn6^
601^3830 q3ihcnrn6'b3 ihb3&3pn 9u36ih33pn biqjihnb 0n3h 6 ih 0 mj96 np 09 hip

60301031136366 9(336ih 3x n oii6ibi‘bm6igm36nb 3nh33pign 3hmg3jutii 3601936;,
(PP). 3nh33pign 3hmg3;Juni 9 non 03ljni, hi(j 3oghm bh xê 9oiin6oiJ036i biibn 9 n-
piljnm 'b3pi3nhnb 9n3h 3jbon6^n3h bnb(j30i9n g>i9i3ipn bn6ioipnb 6ijjign g i
hi(j bi6j)hdpn3ni ghm, hm9 pnb 5 i 69 i 3pm6 i 9 n 9^36ih3b 93ndpQ6i 3jm6p3b 9^93-
6noin jjiboii U3pi (ibn9npi(jri3hn ^hnmgnb bi6j,hdpn3mbi).
9(j36ih33pn biqjihnb 3hmp3;J(jni bi0mnbi^3i>i ghmnb 3hoi33p9n 9mu303p cgih-
cnm6'b3 ihb363pn 9|j36ih33pn biqjihnb 9n3h 6 ih 9 mj9 6 npn 98 hipn mhîfcx11 603-
- 152 -
3 oa 6 ^â~3 g;n m ^ b a s n a ^ n a & a & o b 6abao6&,ôb btroyaeo
crmQhQbnb 6m6ooi (gi ih i qjmoiQnb b3gi3nhnb 3 n3 ihoi, hm^mh^ u>e!>3DCn ÛD"

6ihnb 330cnb333i3n). 3 u ] 6 ih 3x n biqjihnb Shmg^j^nnb bnfijih] (CGR) b^CiQDCn
9u ] 6 ihnb qjmoiQnb QhcnQxnb bofijihnb (ULR) g i 3 n6 nb b3giUnhnb qjihcnmbnb
Qhcnjc'bQ 0 mb^cn q)fncnQ36nb b3gi3nhnb qjihcnmboii (LA I - qjmaignb q}ihoim6nb
nBg^jbn) bngng336nb 5 i8 1^3636001 ni6 j,ihn336 i.
CGR=ULR • LAI (5I1 mhj,. 03h. 3ibi . 9 "2 9n6nb b3gi3ohob qjihcnm&n r 1 ) (2 .19 )
ih ^6gi gig>3i3n6yQ3b, hm9 01(1033 3ihi93ohn 6 i 53)6 y 330 Cn3 nb3C3î 9u36ihnb
5>i63ncnih36nb 3hmi53b9 n.
0hm3ni36bi g i b36ljhm3ni369n C4 6i0ibm3636o ij^nohn bhgnb b;jbm63 n
ibn0n0i(jnnb 9 ijbo 9 i 03 ho boRjih3 50-60 &h 93 h. 3 ib i . fr2- ge^"1 ip633b, 3 i 3 n6
hmg^biij br^njh bm6i8o 303(101 gi6ign 5i6iOi36nb 30(10163680 9ijbn9i0;)hn 96 o8 -
3630nb36ni 20-30 j,h 33 h. 3 ib i ■ &2- geo"1. C4 iXiiD^DC 9 l5D6 ih 30 ib 6ih5i-
0163680 (j)iifi6ni bib^mbib) 3b 8^33636300 15-30 &h 33 h. 3 ib i • 9 "2 - gcp]"1 xp-
633b, cn30|5i bhgnb bh^pn b3bm6ob biBjipm bngng] 6ih9oiig536b i6nb jhcnn
03bi03gngi6 6ib33hi3g3 [ 157 ]. ubh. 2 .17-3 n 6iR33036ni bb3igibb3i
bi g i 6ihj,im6nb 33 hipn 6n3om3h3636nb 3 ijbn3 i 03 hn 96083630016360.
qjmc3i0nb qjihomb o6g3jbob (LAI) bhgiboii6 3hcnig obhg36i qjoiôiboB-
cn3b3hig ij^njhr) b3gi3ohob qjihoimbn g i i 8 oooi9 9mbi0mg630ni, hm9 9i(pi0n
LAI-bib ihmg^nnb bnRjih3 9 i(pi0 n oj63&i. 3b 9 ihoipiu ib 3i LAI-nb gi6i0n
96n33630(Ti6nbicn3nb, 9 ij>hi9 hmg3bib 9u36ih30ii bnb9nh3 8(336^330 biqjih 3 n
nbhg36i, qjmoi036n 633b igj, 003680 3hcn9i63onb Rhgn0i336, iSn^mS d0O3h gifi-
hgno3c igj,n0363n bn6ioi03 i{"ih ihnb bi^Sihnbn C 0 2-nb 3mbnvi)n3hn 6i0i5bnb
9nbi(j>633ig. i8oôi8 801300 9^361(13300 biqjihnb Shng^nnb CG R 2103630016b
(6ib. 2 .75 ).
iShnîg, 3b ihnb qjmoioob cgihcnm6nb oi3ô8 i 03 ho o6g3jbo Shmg^nnbio^nb,
(1018300(3, hoi&mhu 63bn, 9nn(n633i 9i 3 n6 , hmg3bi^ g,i8oibbn336i aii6i6hig 8oii-
ncnj83<bi ^ynb 536mbnb &i6^ni03nbib. 3mhnbm6o i 03 hn 0301010360b 9 Jm63 6i0ib-
61(13680 3b 3nnp533i, hmg3bi|j qjmcrignb qjihoimbnb o6g3;jbn 4 -6 -ni, 9 ihi530Pi3-
63680 - 8 -10. ib0m n6qghi6ncn30n ou>6iq3ihgoi6nb (0 .7 -1.1 119/0 .6 -0.7 |i.9 ) gobîS-
ljn3ho 5ibm03i b i33i036ib od033i j>iobm9 mb qjmcngnb qjihaioi6ob n6g3jbo 8153-
6ib. 2.75 ii30nhn bn8n6gnb Qhain 3^3-

O 6 ihnb ojmcncnb qjihcnnb ohcn^xî (ULR)
0 g i bn8n6gnb 5oi 5îonnb Shmg^J^xmdib
0 0 (CGR) 9 mhnb, qgmoionb qjihomb n6g^J-
0 2 4 6 8 10
bnb fpoi3i|vib6n6o5noi (Williams et al., cit.
Leaf area Index [m^m 2] Baeumer, 1991 8nbog3noi)
- 153 -
6 ab 3 0 6 & ^ 0li anb&HnObi ett 603m00(!>0 &0b P ^ a n d 8 6 6
ybhncn 8.17 39hi(Yi 9ibnb 3hmp;)Jijn> g i 6iOt3b3&3n 3mbi3cnb iQi^bnb nbpjjbn

(6ioi]b3&3n 6mbi3pnb ipj&nb n5pjjbn 33hipn 9ibnb 6(yr)h 6i 3 i^ 9 n ]im5m3nj^hie
9608363pn3iB Bnpb; S - (njbcn; F - 6iyrnqjn; M - Sn&ib'b^pi 9ibi; R - 3 n6nbj3] 3i QiSijhp^j-
6 x *i mhfti6m36n; W - SjlvJi&i; Lieth, 1962; Loomis & Geralds, 1975; Siren & Silvertsson, 1976;
Cooper, 1977 ; Wolverton & McDonald, 1979; Eagles & Wilson, 1982; Osmond et al., 1982; Schulze.
1982; Jarvis & Leverenz, 1983; Nobel, 1988; Nobel et al., 1992a)
(a 6io*3b3&0n 9mbi3(Vib ip3&nb n6gjjb n = [3>,m6m9nj3hn 5idm bi3i(Yi/3hrnejk}nnb J>i3mbi3icn] X
100; b bibjJ^&noii g i yci33CGC'n3t,n 9mh6y3non)
93 hi(?n Bibnb 6ioi 3b369n 3 mbi3£nb

9(536ih 3 9 ijbn9 iC ^n iC^)6nb n6p 3jbm>
( i 5 Sahip 0 &)
C 4 au^Bbhjj&n
Sijhnb p3h6i 3 n 6 - 8 0.85 (M)
bn9 n6gn (Oo&ijhr,>3ni 3&bi g i
^hrOnyj&Sn) 3 - 4 -
bn9n6@n (broSnjh bih^yjcSn) S - 4 0.4 - 0.5 (S)
Panicum-nb &3ihnb (jhm3ni)Xn
6ih 3m3 ie j, 36c 36n 4 - 5 0.4 (3 ); 0.88 (M)
(jbm33coii b i i 3] 6 n j,hm3n i x n
3 [S36ih 336n 3 - 8 0.85 (M)
C 3 8bh(}30m3636n
6hn63n 2 - 5 0.4 - 0.55 (S)
bmhbiQn 1 - 3 0.25 - 0.45 (S)
John “ 1.5 0.32 - 0.52 (S)
d g X 1^ 3^1530^363^1^ 2 - 3 0.7 - 0.8 (M)
<J:>m6nb 0^1330^36360 5 - 10 ’
SihynbS^&n
XjDhbi 3 -
•nm 1 - 3 0.3 - 0.35 (S)
1 363h-6 mpj3ni636 n
Manihot esculenta 3 - 4 0.7 (R)
9 ijhnb ftihbiQO 2 - 3 0.45 - 0.67 (R)
iih^mqjnpn 2 0.82 - 0.86 (R)
2 -
Helianthus tuberosus 2 0.75 (r )
Vjcrmb 3ic 8 i 2 - 3 ■
tjynb b > % j 6ih 33&A
Cryplomeria japonica 5.3 0.65 (W)
Pinus radiata 4.6 0.66 (W)
Pseudotsuga menziesii 2.6 0.71 (W)
Pinus nigra 2.5 0.46 (W)
Picea abies 2.2 0.61 (W)
Fagus sylvatica (60 6cnb) 1.3 0.70 (W)
jjnhnqjn 5 0.6 (W)
3jhb 3nb 3n6hnp^6n 3.5 - 4 0.5 (W)
Eucalyptus grandis 4.1 .
Hevea brasiliensis 2.5 - 3.5 ■

C A M 8ls36ih 33&n
i 6 i 6 ibnb 3Qi6 j)iun;j6n 2 - 3 -
1 - 2.5 (4b) -
0.8 - 1.7 (4 .5b) 0.2 - 0.35 (F)

Bycnb 9iu 6ih 33bn g i BusrflKTfiih^jBn
Eichhomia crassipes IS - 20 -
J>i3gn5ih 3 5yonb 3.5 - 9 -
t ^ n b 6yic 3 |536ih 336n 3 - 5.5
- 154 -
6ib. 2.76 U36j|hic 3hn ^cSD&nbi g i 3o63b ] x nl5 i^hpnQn^Qbnb bb3igibb 3i 9l5j 6ih x )0 bih^yx^n
0U]6ih 3x fI^)n*J ly'Ô^xiL) gmcnonb qphamb n6g]Jbn. nj, b igi^ Soihxn SbîhQ x 0 biqjihn igfc-
npb 3Cn9 m6b ih iS jih x . pni 9(jo6 iho](V,i6ib, qjmcnpnb ojihomb n6gj)bn dQnQh 9unhg]6i
(Vareschi, 1951; 1953)
6^3300 biqjihob 3hi3iQn 33iQhi60Cn i3-bicn3nb.

qjmcnpnb o)5hcnoi6nb nfSgQjtm gi 8(536:1)1300 bnb3 nh] (01536^3390 biqjihnb
bihnbbm6hn3n 0 ^ 33636390) ihiSih^m cn3ncnm6 06(1836301^36^ 00^0393636 3hov
Q3jl5ni'b3, ihi03g 013001 j)i6n^gni6 0nb 5^3036^. jjbh. 2 .18-0 n 0oiy3i6n0oi
bb3igibb3i 0 (536ih 330 n oii6ibi‘bm5igm36nbioi3nb LAJ-nb y33Ci'b3 3qjhm gi-
0ibibnitJi3630n g i 3jbj>h30iC3hn 06033630016360. diorii6 b3 nh, 95631130 ^ 336 311
LAI-nb jp'bhgib 15 0s 0 "2-nb <b33ncn (bp3gi3b bn6icn0ob 6ij)C36m6i. (pni 8153-
6ih33o cni6ibi'bm5i901363600, bigiy 6oigij,o 06 nhn, j30i6n, di0Oi6 00hi0ri, 15030
i6 0 oi03i, LAI 0 (johp3&i 0 o6 o0iO3h bn9 ng30 g 3, 0 ij>hi3 i 0 030 oib333i 0 o,
030l5Oh3&i dohooiigig 8(536^3300 biqjihob bob3 ohob gi(530noi ihob j>i63nhoi63-
£>Xn (6ib. 2 .76 ).
LAI-bi g i 8(536^3300 biqjihnb 3hmg3Jyoib 0 mhob 5(1636300 gi001309363-
0360b 0bj)i3boi 950013003630365 i0 33i6ibj)63obi g i dnBnb (b3gi3ohob 1 02-'b3
0 m bx jomhoiqjnoob him936oi6ib 0 mhob (ob. ybh. 2 .18 ). 6yoob 3 ibob
bo6oi3‘b3hig ij^03hn 3h336nb }h0103)1aiîgicgil^nb bihobbob gibibiboiOi36£ig
5i6bi'bc'3hi336 Byonb <b3gi3ohob 1 3 2-nb ^338 Byonb b33li)3n 3i5bm3h36 30i&
j^mBSn jomhmcgnQnb 038(53300165^ Byonb ^giSnhnb 3h01330^33 JonhnqgnQob
bo333hn3nb j>ib5'b01859 b^3 ;Jtih503hn i6ion(bnb 55301936365 (013 (560160005 ibQoi
0603)156001300/6001300 01565035)190165) 0133563015 3'bih3icbihn qj5hoioi636ob 501^36-
I503hn 3hmg3j(i)o3on 6 nb 5i6b i‘bc>3hnb 03bidQ36(yn6ib ndo33i.
- 155 -
6 ab30 (ng»agm
6 nm33 n (Whittaker & Likens, ^ aarô eoaomoô&oij ff6<f>and86.b
ltj o m o in 0 o to o w O rO 04 p O
fO ol oi r bb3igibb3i
fO U*l p*. r—
ei Jcmhrngncnb
in O o ^ ^ t r - ^ p — qco O O _ 04
<D -O C> — cw- —
r-
n6p3jbn
N « O W Kl r - o Or - o o oOH-
Lfl i K f
1— u i m irt in ki
O Oi r- r~: d d d d d d d d o 0 0 0
<3fT'cnCnb qphoinb
Ul o i
l/1 in
oi U1
tn
I-
in in
— <w r- o o o
^ m _ «■ <>
9 - 0
rn , , oj , . • . . o ■ o . in
m cm
oj .
1 -o «t 1 oj m 0>1 1 001 1 o0 o0 ’ r-; 1 S •** OJ
^ o o o o o o d 0 0 — 0 o o o o
o
«- oj
*0 in
to — O' «0 — OJ OJ OJ
1975; Ajtay et al., 1979; Schulze, 1982))
Ubhnon B.18 6efr)hn 6otim-3nh3x ^ e n
&
/n
o
2
! S £ 4 <8*3 &
- 156 -
2 .4 .2 . eoeânenb a i 3D 6 ^ h D x n 6 f i b 6 3 i i n - 3 f i h 3 O0 en
3 h n e D jiin x n 6 ^
0 ^ 3 6 ^ 3 3 0 0 o iiB ib ib n ifc ip r ^ n b 9n;jh 0 3j96n p ddhspn 6njcnnjhjdnb frsdm-

b*3>cb jjddshntf JnhjjQ sg ^hmgjj^b 36(^3636 g i nb 5i0mnbi,$36i hm^mhy
09 hipo 6n3cnn3h36i (i&), hm03Qnu 9 mgnb 6oipi2>o6 qphaim&nb 1 92-'b3 (56
,j/3i)- 63^m-3nh33QiQn 3hmg3junnb h^nmBxn *6 &pm6i03hn 0 ib 9 ,ji 6 ncn 5^6-
bibfp3hi QC30Q3 b ii 9 im bnd63P3b 6ih0mig2)36b pb bb3igibb3i i3^mhob 6016^3-
036n bi^0bmp 5i6bb3i3Q36oi6 3hoi0i63ainb5i6. 0n3b3gi3ig i0nbi, 3^00136 bi6b
bi^Qim 3hmj,h3bn nJ6i 3 n(p63 X n pnb|ii6i5n3ho 633^jhip3hn s6i0n'bnb &i9 m-
93636001 (6ib. 2 .77 ) 0 n(p363c>n 0^3363603606 ^mOS^^f^po g i 039 i 336 noi.
0i(?ipo 63l>)rr>Onh33Qign ^hm g^ni 0 n0 n^nh36 x n i 2>m6l!in636^36n6 g i m ^ i-
6336nb n0 (132,01^1636001, 6ipiU 0(536^33606^01306 Bypnb, bnoi6mb pi bi$336nb
6n30in3h3636nb b3pbiyh3po 03bi036ii. b03p3cn'b3 i0ib 363936001 ^360^6 ^hm-
30^3680, 6 yip9 n ô bi0bh3Oinbi gi fihgnpm30inb 5i63p3&ob 40 ° pi 60 °-b 9 mhnb
cbm6 i 9 n (6ib. 2 . 78 ). y33Cit>3 3®}hm 3630 3hmg3jô30fn6 i i(pn6n9636i n9 ip^n-
03660, bipiu 66303010 g)igigob 6 yip9 n - <bp3n6 bi 6 i 3nhm6 6yip0ihfib369n, 9 ih-
3606 hoqj36'b3, ^0163680 p i 016090 3i3o6 giîm 636x ^y33&9o (ijbh. 2 .78 ;
2 .79 ). 01609, ^36oi6 h3501^163690 ihrnpÔi pn0nljnhp36i 0o63hi03ho 6030103-
h3&36ob bnSôhocn, di0ni6 b9 oh ^33680 30 - 6n6iOi0n6 6ij)p3&m&noiii5. p3pi0n-
606 (b3gi3ohnb (hm^mhu b0303Oinb, ^ 3 3 3 Bypnb) 303^3b 6iBo00n 9 bmpmp (bm-
0 o3ho 636fTv3nh330 ipn 5 hmp3ji5nnb 5ohm636 n ihb36m6b. b9 xD OTnb 4 1 %-'b3 9 m-
6i30n6 0i0o0n6nh3630 ihoôjjp qjij^mhb 6y0ob 6ii)p36m6i 6ih0mip536b (6ib.
2 .80 ), 8%-'b3 - ihib 30 biyh 30 n ^ 3033hij)3h36 n &i6bi'bp3hi336 pi6ip 3hmp3j 6 n3"
Normalized difference vegetation index
6ib. 2.77 bbjigibbsb ^n3nb duîhxJC0 biqjihnb 6jyn 3nh3x*en ihmg^Jjnnb i^30nhn 9(j36:>h33cm-
6nb eogjhêynx n6g3jlxni6 ( i X 33n'J 9x 360^n eôndBjbxni bi^xyilj)36ngi6 dpn^h 9 ipi(vi
h^bmc^nnb hignm93^hncn Sn pijbx 906^3936133). 1 ,£)6ghi; 2 ^35gh>v>inj)nb 3ir»m)m5n; 3 61*1(13-
iC3^n 6n63n3i6n (jyjjftnb bih^yjpn; 4 t>n9n3hn bih^ypnb 339ng3hn ijy33&n; 5 5ihgi 9 i 3ion bih-
iiy3Cn 6063^13^6 g i qjmcnpm3i 6 (jy33&b 3nhnb; 6 qjnOTcri3i 6 n i5y336n; 7 03b6 ih-q)n^36ihn 8383300
iKQ36n; 8 cgn^35bh36n; 9 6icbb6ih36n; 10 ^ miX ii 3h30n 6^9^3636 nb 9n636n; 11 &3fi;j6ih3&n gi
93Rb3hn (ja336nb qjmh9i(jn36n; 12 3gi&6m36n (Goward et al., cit. Field, 1991 9n b x 3no')
- 157 -
6 ,bb3 0 6 & a m
anbaaftaaa ^
- 158 -
99 <^g?o G030 )03 ft 0 J>0b B ân d ae,)
fbb. 2 .78 a bOxjonib 6(yr)hn 63^ 3nh33Q:>gn ihmgj^nxei^ (bh 99hiCn Sibs cT2 6 jcn 6;>(j3 n) (Lieth, 1972). ^ 3353,
iy>m ^nmpnb 33bibi9nbn 3nh33pipn Jhmp^nxot&nb ae(T,&;>C3hn hj^bn nb. Uchijima & Seino ( 1987); b93C3on
3(j36ih 3x ft bigihnb cgmljmbrt6cn3'bnb i3b>b33pn h^jbn nb. Terjung et al. ( 1976) gi Woodward & Smith ( 1994)
a n a G ^ a ^ o o t tG ^ a n & ^ r m o & o L 6 ab30 r t w e m 5071x060
- 159 -
6 ^ 3 0 6 6 ^ 01* 9 Db96fn066 9 9 6 6 ^ 0 6030)00696011 P 6 ff»9 n d 966
fob. 2.79 93gi9n5nb bbpgibb ^ 6 nn83 n 6tyrjhn 6^yn Snhsxê *1 Sh m e ^ iy ix^ ^ (PPn) P1 qjn^,-
m9ibnb him g^m bi (B) (Odum, 1971; &i9my363&3eni 15830133 b b ji i^mh^&nb 9m6i(j303&nu:
Bazilevitch & Rodin, 1971; Whittaker & Likens, 1975; Ajtay et al., 1979)
0m6sb ("bhgnb b:>60m.j)03 33hnmgo 5 i 63nhm6 ^6 ]onb b111503001; 15030 bitgbxnoi).

^hmSn^Q ‘b(p3360n b:>5336n 6n3Cin;)h;)&;)&nb 6bj)Q36m6bb, Smp^bjbob bn:>b0m33b
- Shmgrjj^nxm&bb b [3139336 ih ibs^hnbn bn6boiQ3 (nb. 63b. 2 .78 ).
9 l536ih 30 ii 53936363 gi iDW»n3M5ni Sihibo^bob bi6o6ii(^0g35m pm6obdn3-
6360136 yn9&n6i(5oncn b30;):>O36:>b 00)0333 6030(3^101 oQi'b] 633)130 303)1m 03p 3cn
3nh33Qi.gn 3)10193^03, 309)13 g 303b3bn3Oi363903 6i î3D X r* ^Q6nCn^ 63536ho3n
015363)13300 ou>6ibi'bm£,igm3636nbioi3nb. 69hi0 n 9 :>b30nb y3303b 3 93(3>30n 3hm-
gudl5ni 0np36xn3 3.6. iX iW Ô ^6^3600 ^53(50303)1 h3'b3h333h369n) 53tbhgn0n
6yi03(536^h336nbs6^6 - giibgmQbnoi 10 ^5 3'2a_1 [ 115 ]. b9303 Oinb 915363)133630
(63360b Rioi30ooi) 6 om9 bbob 6003)10 3hmg3ji50i 93060363 jpn'bbhgrnb (*1)133)1,
fob. 2.80 y>3anhn 60n3h 63^1-

3nh3xê Shme3ijnxm^ ,i Si Bcn^h 6303-
J36b 3 mhnb (Lieth, 1975a) Precipitation [mm]
- 160 -
3006 ^ 0 7 )^ 0 roaeaiasngK^no&o&ob eab ao rt& am &0? )}060
6mh3503h, 636361103 856336360360156 3395(136001, 063010 ^3150503(10 (jim6nbdn3-

6360b b5935Q36no», hm5mho(555 'bhgnb ym330n cg5bob5cn3ob biôhm 3mh6y3i 013
6059550b 5561^03(1365. 8036395359 50nb5, b5bmq30m-65833h63m 330143(13606 65-
9 350m 0mb5350n 33(1-3311016001 116365 3qjhm 956500, 309(13 33od03&5 80(316330
njBob, 39oi53h3b59 qj5hoim6 ob 31)6^36603(10 95 5h5 06^36603110 5580^363606, 63-
636(1030 3hm93jl|)o30m6 ob ^homgob 5h5bh30O 550my3636ob5 95 33ihg,nbn 013b-
0360b cn3b3nb 558m.
6 nm05bnb 9090 55801653500 6583503656 od0335 558m3oy36moi 556653301(13-
6noi 05(j>50n 3hmg3jô 30m6 ob 8jm63 3i3365h336o1 63900 85bnb5 95 6001363115006
039803, b56j)hd0n3 6y5hmg; 0138(55, b9oh59 801300 3(10193(50(136300 85606 55-
80^36365 33(1 b 3hbg365. oi3b036ob 338016333580 5580^36365 653(10101 33 h50o
6030103(13606 (0mb530o56m6ob 0693^0) 30 -60 % , 863563 85bnb5 - 85 %, 0ohj6ob5
- 50-70 % (ob. i5bh. 2 . 17). i 30»S3h x 0l5365h3Oi5 5hmg3Jôxm6ob 55bhgob y3305
3(5030m6ob5b 3695 5356601393^ hm0 o6^36bo3ho 430^035^505 (55836365) 5563b-
h3059 0mooibm3b ti3j6oj)3ho b533503&36ob5 95 36311500b 909 95656511336b. 013
55801300130001 3hm03j(5onb 5hb36ooin 55gn936ob50i3ob 53150036300 956565(133606
3cg3^3hm65b (363h5nob 9565b5h3n 60(3136300 558mb5y3636050 35h5obo 6000856116
363h5onb 3hcn330b3 55955655110336001), 5(3*8(11669365, hm8 <33(163(130 033(163^165-
3 o 5580^3636x11 3ho0 oôx<i 8301019360 50-53h 333013605, 053o6 (1019365(5 01565-
039hm33 65601(3001-65333(16301 8301019360 - 8bm0m0 2,5-33h. 563 hm3 , 35hai50Oi
5335859 6 om85bob 3(10193^(505 <35(10101606 3hoi330b 3 55(50036001 83^05, 309(13
d330 9(10180, 855h53 g565b5h336o 0 ^9365 55(50036001 3qjhm 36o33630m3659 95
66(150359.
2 .4 .3 . 0l5D6ihDOTD 0 1 ^ W <bn6i9n D 6D 6flb 6^b3nh6^9f1b

b rm o ib n
2 .4 . 3 . 1. 6 hD ibn-3 h n e 3 jl 5f l i Q } 9lSD6 ^hOCfD C n ^ b ^ m to e m D & n b

bD601 j 3 i
3(5365h3cri5 oi565b5bm55gm36ob 6(13,401-30(1330590 3(10193^05 (PP),) 33350mg

3 nhg53nh 33b 55obm33&5: 8 o5b0 m36noio 553 moi305 8 mo(553b 6(5365(13015 0156565-
bm55gm36ob 63(401-30(1330590 3(10193)15006 (PPn) 563 30(1330590 3(10193^(5006 55-
8016530065 95 63601)306 ( S R) 556b5(b(3'3h5b:
PPb = PPn + ER (2 .2 0 )
b36oij3nb ,,3089065(13 956565(13366“ 5580165,45336 3(101(536,43690 56 50im6 ooi

6o0363o. 65056m356 8(5365(13015 oi56565bm55gm3636o 636oij35b3 65(135336
3 oi56oij030n 65b3oh65gn6 20 -50 %, ^ 3 3 6 0 95 3335 63^65(1360, hm830oi5(5
<301,4016060^3(159 5j^03h0 35b5 9395(136001 9090 5^301, boi0o3h bm653o - 40 -
6 0 %, 01600 95 ,536056 ,4(1013033680 30 - 955600136001 70 % ((jbh. 2 .19 ).
- 161 -
6 ^ba 0 fn6 ^ ^ 0b 8 P)b8Art35 A 83 <f> ^ 0 6030)0 0 6 3 J>0b ff6rt8nd86i
2A.3.2 63^n-3nh33C^Gn shmeiMiinnb a ^m b ô n
63 OrTV3 nh330 ig n Shmg^Jijni 510010936361 01)1516300 6n3tnn3h36nb 16193636019,

hm0onb 616090 031)15361, hi^ 6906 5160139016180 J06 nb R iO n^boo biq^bb
(L), bmpm 6i6n0n 3(1 518^0936361 301663036^3606 803)1 (5106013361) (G).
91(1636090 63OfTi-5i0f1ibi3i0o 681^9361 6019150b qjihonm6nb 3hoi33Q<b3 1(^36390
qjnofn0 ibnb 5i'bh9ib (600181606 sibh gi = + AB). 3jjn0ri5n3hn 3hoig3jijnn6 516-
0^0361 ip63hb 6ib8nh6ignb 6i0i(Jnb 51616110361b:
PPn = A B + L + G ) ( 2 .21 )
3hrn93junnb 10 516(501936180 y330i bngn93, 3.(1. 600181606 1530(10361, 61)163-

636(1, 8m8b8ih360(ib 8(i3h 51001393636300 6i6(i0n 831^9361 38310019 5i6o6i'b(j’3-
hmb; 0ionn 318(1 6111801195366 bi'bmSb, (1018396115 ^ 3 3 x 0 ^ (13 (19363636 630m-
3fih330 ign 3hmg3jljnxiTi6nb 5i0mbibiî9 (518011651)1083606 815190am nb. ubh.
2 . 19 ). 6363611(13 <3(1^01153601^3680 y330i 30i93m636v3nb 5 i‘bm03i 193110(1 ih ihnb;
ib3, 0933661 91 b0303cnnb bb3i 0hi3i0nih3bm6n 8j536ih3xn Oii6i6i‘boi5igoi36n-
bioi3nb 3hmg3ji5nnb 816336369360 3ihi39noi 3691 830)1636, 013 nbn6n 5 ^ 6 6 1 ^ 3 -
h306n ih ^636016 08133 phmb bb3i hm03 fc>3hbocn.
6903)10 60 n6gi 30)1339190 3hmg3junnb aghijym (6n0n), 6ih0m9536n0n L 91
G Qi6i2)ih5ncji, 3jjmbnbo39nb 8u 36ih 3xm6nb 6ib8nh6i9nb 6i0i6bnb 0131961^-
hobncn 3(1000339 qpj^mhb 6ih8oii9536b (61b. 2 .81 ). 60n3hn 61(16363606 3hm93j-
I501 (930(10006 163 Ri3m6ii5336n 83391(10 0ibnb, hiy J86o6 bigj36b) 3q)hoi 0 ip i-
Ijbhncn 2.19 ^ynb Jhmejj,503^63 gi rnhj^xn 99ti^Cn <kbnb i^h63> (Bpo^hn 63Q36bn)
(y330i 9m6iu38n 0my336nen3 35 a^-V); a Mar-Moller et al., 1954; b Kira et al., 1964; Yoda, 1967 :
eji^C^n 8m6;>u303&n 'bmQnjhn bih^yQfib 6063P133611 (jyjQ&nb &3C36b<b3 nb. Vogt, 1991 )
sshsQJJhri 60 60013360 63080360 ,593

SnqjcSitin ^ 3 îm 3o i 3(vi
@3603 a ..... i*>nC36eo h
LAI S.6 11.4
AB % PPg AB % P Pt,
qjmoic3&nb Bpn^hn 3i£ ) 6i 0 0.003

Cohm^dnb 6003(10 6^363 0.53 0.29
g3b336nb Bcn^hn 83,5363 0.16 0.02
J39n: 0.69 35 0.313 2
b>q336nb 6eo3hn 63^363 q jm o ^b n b b s h jb ] 0.27 1.2
bicj36nb Bprrjhn 03,5363 pjhm ^bnb b s h jb ] 0.1 1.33
b 30Q 6nb 6003)10 0315363 q)3b336ob b3h3'b3 0 .0S 0.02
J 39n: 0.39 80 2.55 20
PPn = AB ( 6nm03b3) + L (b 3qj36n) 1.08 55 2.86 22
gm oipjbnb b^fcoifoi'bQ BQn^hn 936333)12,0 0.46 6.01
C>3hm36nnb b ^ & jijsi'b ] 6003)10 936333)15,0 0.35 3.29
q}3b33&nb bêatjjb'b] Bpo^hn 936333)15(1 0.07 0.59
330n: 0.88 45 9.89 78
PP„ = PPn + AR 1.96 100 12.75 100
C U tp p = (PPn + A R ) / A R S.23 1.29
- 162 -
3 pa 6 aff)a;pg?o m ^ i^ a n a ^ n a & a b o b sabaoffr&agm &033060
Cn> j,hm3nij]Q ^y^jbSn, b:>gx5 a^gj^fjb bxJîgmg 2jj$ &■%’ (0136^ biQbhjoin
ibnnb B3n0nb6 (jy^bSn, n^n S^ndp^b nymb 1,5 i&Qâ'1 ) [ 1, 175 ]. Rbi^^npn
9uo6 bhoxn <3 fT'h9 b(3n36 nb 30 hi 3Q3bm6 s9 n, hmfcrnhni^s 0bj,bpncru>g, ‘bmOnQhn
bih^ypnb i^yQD&n g i 3 X 3&n- 0:>fin3&n, 0i6£)hnb3&hn 0|536ih3xrn6:>, 3D3^> &3Rj6ihn,
Deciduous m ixed lores!
PF* -G*L> - 2
:r: ;G
B.6
S o il o rg a n ic m a ile r 35
M eadow Arctic Tundra
63b. 2.81 6om93bob 838350 93 888390 (*18536390 6n3cnn;jh3&nb 9o9m|yjQ3: i^h^rv+toyrib ^hmSn^p
30833039 6308036 (Odum & Pigeon, 1970), b39bh3on 630)836530106 63839863363 qjmcncm336
0>y33n (Rapp, 1971), 6305006 ®D^3X <30101001336 >jy39n (Duvigneaud & Denaeyer-De Smet, 1970),
30^3690 3353 i>3fiJ5jhnb 0136363^01539^36380 (Larcher, 1977; Schmidt, 1977), 8m8i3038o 60336036
993Qm'b3 (Rychnovska, 1979) gi y>63gob ihijn^ ^3698380 (Bliss, 1975). 93801113016393690: 018536-
390 93h3(xi 6o30i03836n6 9383511 (j£ 8*); g39ljhnb3(vi 38336(1 93 6y33^0QO 63,b36ncn 339m-
b3t>c'3h3cn 83801.1)301639360: 888390 8360b 6003811 80801^393 (1,5 9"® 63eo63eo"1; Ba 9o6ob'b393
6nm93b3; Bu 80606^3393 60018363; Be 1)803^015393606 60019363; Bph ^36ghnb 33638014,393606 6001-
93b3; PPn 6jyn ^08339390 3110193^0300163; PPna 9o6ob'b3p3 63^01 30(1339390 380193^0390163; AB
6om93bob 600380 93^363; L 888390 83bob 600380 438533 93^80^06 b3bocn; La 9>j8rynb 636001
806066393 03bob 93634385360; Lu p3^hoôb b3booi 80606:13383 93bob 93634385360; G 99hico
93bob 69038(1 93634385360 6013363^3
- 163 -
G^bBOfniiÔb 8 P)b9^fnDM ett 3 3 6 . ^ 0 603010a(f.060 b P6(f>8nda66
9013306903(159 65(16363606 656001 yoi3og 43igh5^ x ODijh^ ^ 2 ^ .336 0,6-1,2 45-

b. (135001680, 6595(3 6906 65h433x phoi6 88(1590 ^3(100190 95365. <301010198153360
63360b 65(16360 (15019360165 03595366 0,3-0,645&'2a'1, 6^333630, 65633(153956-
6013665 g i ,5369(13630 40 - O,l-O,5j>50fia-1 . 6nm35bnb Ii50ig360i65, hoi8x n i 5 035(1-
536^ 8ni3b8ih 3690b „6yipn6no»“ , d50Oi6 55666353363905 8(5365(133901606 ^n3nb
0nb393ncn; y 33C^> 3330^333580 956535(15360 633(159 65393605 ^^33630. 3ngh3
6505601356015 o ^ i^ b o ^ g o ^ b iO o .
(1010365(5 8(5365(13015 oi565bi'boi55gr'i3636ob qjn^mSibi 3(53C3Cn (16365 (AB =
0 ), 55^(195 (+AB) 56 9(10136001 838(50(1365 (-AB) 95801309363005 PPn-ob 5580165-
3X?b3 9b 9î)bh53on 6n30in3h36nb hbm936fn6b‘b3. 013 53 836589369016360956
(1013390 (1J6365 5580^3636x0 801(53830 340ibob^38i3o, 556n65t>(T»3h365 doho-
015959 58 33565636300b b5b3m6hn3n 838595360016001, 90658(14001, 5b5j)00i 95
3056001 5(1636390 63636ho3fi 95 5601(1013015363(10 qj5;^4Pih36 nai. 6 ooi35bob y33-
05^)3 303(101 3630 95 3 y5ho 5438305(305 56560501366 ^ 3 3 6 6 .
683930106 8(5365(133606 803b 8ai56oij830o 65680(165906 3/4-(b3 83^0 9585(1553-
63905 03 hj563 o.
2.4.3.3. bb 3^b b3i at336ih33Cn 0 D 6 ^ <bn6»n36nb ShmQOjbfinb

e n 6 ^ a n ii
^ 3 3 6 0 , ^yob6n 5950936(1 95 3335 63RJ6iho 56 8301(1590 55^056360656 83(1j6-

056 8(5365113015 3hoi65ohn 5bi4nb 63360 559056 bhgnb 6 ^ 4 3 33 0 qp'bjtAi. 65658
013365(13360 565955^955, ("3(10136065 95 g3b33&nb 35b5, hoi03gb5(5 65630(1659001
0301019360 535(1553636, 3395h36ooi 8(50(135 95 63l4m-3nh3395go ^hmgoJyob, 836565-
3nb5g, 9090. 3b 6006536, hm3 0^86365 mh5563Q0 6030103(1365015 6548501 65(1550
95 815365(13390 65035110b b53hoioi 6 ooi8 ib 5 yoi33069 o3 h5g b6 h5qjig 35v43cm6 b.
l536m<bob 5b54nb 85(436560156 3hoi5g 3(10193^150390165 0156950156016001 „ 5 <bit>
3bb6ob“ bo06 nqjnb 035^56 [ 170 ]: 58 qjibob g5bi6yob3n A B 33(1 3og33 gig^bn-
0105, 303(101 3 m53n5636ncn 4n 83(^30166 6390156 5bcmb. q}n^m85bnb 55<bhgnb
bofijbhob 3b 95^33001365 55630(101636x1^ 63(301-3(10193^006 838(501136001, 0136(35
813365(13390 65035(106 556300i5h365 5(^399365. h5(5 303(101 06(436603(105 "bhgi, doon
3l5oh35 863563 jbm30C36nb 5(1586356360156 8335119365. 580b 550 m,
bo6oi3'b3hn 6585140 65485(10605 86019019 56590 03010193606 65h8 oi65j 86 x ^ e 9 *
yORh(43&ob5 95 <3363015 606(43806 3 ibnb 06o3363Qm356o 559093606501306. qjm-
010108(5336 ^ 33680 3 ohj6 ob 680695 85,5365 <33(169365, (1019365(5 0301010360b 05 b *
b53hoim 05bnb 1%-b3 6530360 69365.
83(1^6056 8(5365(13015 0130015565693659 oi56ib5,boi55goi36368o, ho^m hrnjii
63036936301, 3C+>56n ^ 336 0 , 88(1590 h35noi636ob 636^65(10 <301(105(50360, 3 3 3 5
63^65(1360, 3(1019(1oix^ e 5(16360166 bb35g5bb35 565306 0690309360. 8(336^330(1
b5q)5hob 5563001511365, 3nbn 33895(1180 95460636001, 69365 0mb5O£)h5g 95
001905659 95 3hoighr'ixi Q [ 189, 196]. 58001 80139 0156565^15590136530 9 nr»c^
6335 5565835936390 6m65b6m hxf' 0961*185(130165: 3hmg3ji305 95 9 5 6 5 ^ 5 0 g ^ 6 i0 -
566363905. 5893659, 3h5350O 690b 5568539016580 A B 6390b (40100 *16365 O h m ^ j-
j&m hjcn b&>eni) [ 170]. 0138155, 6noi85bob 3hoig3j(3nob 6^556^(305 5h 65h0m i^-
536b 6m65b6mho>6ob ih o i^ ^ o ih x n b^gonb 531303C36X 8563363639b.
- 164 -
aoa6a<na?reo m a e a b a sn a ^ rm o a o b GabaotiMqob aoaxaeo
hngQbiU 3 (j35ih p x n biqjihnb bhgi 313jQi 6 x ^ G 3 6 nm3 ibnb 5i683mh36ign

gi6ij)bh536nb &i0 m, hi^ 6 ipibnb3 î 330oii dm336nbi i6 b i 3 oiih3 n a»i30 nb bi-
q3>h3 miCD^>Xn 6i6n036nb 8myn63noi 33nd036i nymb 5i63nhm6363pn (nb. ipS^hn
3 3 ji 6;)fij6ihn 6ib. 2 .81 -‘bo), 6 nm3 ibi hfi3&i bij)3 imp 3x3f»3n 3hi3ipn Bpnb
J,i53b3pm6b3n - 3 ij>hi0 >3 8 3 8 0 * 3 3 3 :^ 3b ihnb 5 ^ 3 qjyJ^mhoii 3mj03036nb
9 x D 6 n-
a e x " 6* W>3* &dc^bm3^6n cii6ibibm5igm3&i6n

3hmp3ii)nxn 33hnmpnb jp 60 i 3prri&i3 n ipj,opo ij3b qgn^mSibnb bBhiqj fcibh-
pib, 3 ij)hi3 n3i33 ghmb ypmh^bnbi pi qj3b33&nb 6i6npn 33P36i pi 0mnb0ih3-
6i 3m6b3336lj)36nb 3 n]h. 6ipibri3i6 oii6ibibm&igm36368n AB 03hy3m6b 63001^6
ibprnb. 33 hip >13^^163680 I530ign 6^03^36001 3hoin3hoi3m3p336m 6p36nb 0i6-
dnpb3 AB 0133036^ bi30img png gig3dnoi g i 3ihymqjnoi 0 ^ 336 36036b Smhnb,
Si^hiB 3hi3i0n 60nb bi33i0m bngnp3 ym33poi3nb 0in6|j giib0m3&ooi 6300b
(!>m0o hf>36i. 6i0ibm3i6 oni6ibi'bm5igm36363n &3636hn3n g>i630oiih36obib bhgnb
b3'bm6nb 6mpmbioi3nb 030101036n y3noi0p36i pi bS^di, ypfnh^36nb pi q)3b336nb
6i6opn j)0 buo0p36i 3u36ih3b. b^33368n 3b pi6iiihj,36n &33hip 33^ni, 3nph3
9cn30n 600b 2>i63 i 3pm6 i 8 n O}mh3nh3630n qjn^mSibnb 6ib33ihn, 3pi66mb
3(336ih 3x cni5ibi'bn5ipm36363n yi, hm3036o(jj dnhooiipip 3qj303h x n bib3m&3-
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3b 6 i 3 i^n 3 m3 b8 ih3630 oii 8 oi30 n 53600b g>i3 f*i3336ib 33bib3h36i; G 8i(?ipni,
bi83i0m 0 mb0 ih 36i 63 ^m-3nh330ipn ihmp^jijonb 2 /3-b x^nb. gi6i3ihj)36n 6ih-
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036001. fiidoh3nb boRjih3 gi8m3Og3630ni mhj>i 6 n'b9 nb 'bm0i'b3, cgr>ih3 ib i Qi 6m6-
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ih m p ^ n xn bhgnbi g i 3hrn^3j^mh3pn qgi'b36nb Oii60n0g33hxn &i30i (03igi-
h3oi oii63 n3g 33hri6 ib b8303Oinb 33mbnbli)30368n). iji(3, biOin6n 0g30fi36nbi pi
bidm3h36nb 0b5i3bip, pi6ij>ih536nb pnpn 6npn ndm336i<b3“ 0mpnb. i0j>3ihip,
6ypnb 33rribnb^30368n G ^ L, bmpm 0 ioin 3i0o 03(jni AB-b3.
- 165 -
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3cghm 3g33jii3hn ^hmy^bni. qjm^mbn6cn3‘bnb 3hmi53b3o 9 bnb bbn33hn 363(150^
5i9my363&nb 3133^3^61, u6m6n0ni hrr^mhu higoujnob 5i9my3636ob 3<33j^3h-
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<30^0*136390 jnQr^hn 3133011360b bibooi.
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163 qjm^rr^&ob himp36m6i, hm93Qn(j biôhmi 3hcnn 9 m0333Oi 02-nb j,b3 mbiy-
mqjig, 3ghnb 0-b; 3hcnn 9 fTiC3i 3Ci bCDi^nb 6ih9mbij0630iQ biôhmi 48 gjm-
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kJ-0116 ( i 9 363*150^ 33111513b 400-700 69 -ob î0(j>ob bn^hdnb 9jm63 <3Pi,jm636nb
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33 bidp 3&30 n 3<33j63hm6i. 0^0190*136390 jQmhm30ib^36nb, 16 33)139301 b^bS^B-
<bri36nb 519^ 36360 0 1 (013^09193*10 3nhm&36n; C 0 2-ncn 5153(1361) 90036390 3jb-
33*10936^390 916336369360 33ig536b giib0m36nci 30 %-b, h19 516 363*1500^ 31*15-
31 nb3cn 3*101(^36^3, *101501*10(51.1, 3nh330 hn53n, bnoi63hn gnbnSium, 51*1913-
319m . 9i6mhi^mhni3fi, 3hcn6inh 013^09193*1 3nhm6363n C4 9(5361113011 qjmcn-
Ijbhncn 2.20 36]h^nnb i4hj,34 (% 9oipns6 n 5pm6iC3hn h4gn4iying46) 8(5364113014

64b9nh&ignb 4bn8n04(jnnb ghmb (Beadle & Long, 1985)
jS ^ n n b ^ 5 3 5 a i& ^ x ic n . qj4hgn6 ncnn
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higniynnb ih3i3pi g i 5466334 5 - 10
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83Pihign 3hm(j3b36n 19 - 22
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C 0 2-nb 3 i 0 n9 nonh36 x n 830 l53X m&nb 6 *9 ™. 8301(13x0 3(101(536360 000000)1363-
poi gm^mbn5cn3'bnb 0bhogi6 . 6h3^m-gmooibo6o3'bnbai3nb RUE Q^hy^mbb 8-10
%-b Smhnb. b^5cnj3i'b3 gi6i3i)i53&nb 5100130360b 839935, 63ooi-goiooibn61i)3'bob
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9101616010136300 60830360b 3i0oihn03ohnnoi. 6 ii 5ih 0 mi3nQ3& x n 03 hi0 n 6030103-
ho6i 83015136 15-35 kJ 6h-l, 0136151 3ghoi 69 nhip 3b 61633636300 3hcn 5(110
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01011001 60301031136130 6 ib 3 oh&ignb 330(33300161, 0 ooi 03^01 0 obo 363h5ni; 0^53-
6ihnbioi3ob 306033630013163b 6030103(136366 61(1001195366 6ib8oh6i9ob diQoi6
0ipiQn 330(33X'T>2>ob 0^0163 00309360, 0056060 91 ho5n 03fnhigo 03oi&m0o'b0o6
3hoig3Jo3&nbi, (101501)10511, 0i5i0ocmg, o'boi3h36oing36o gi 0icno 61)10013630360
(ubh. 2.21). i063ihig, b ig iy ih 3691 oyoib 0013100^)36300 3b 6o3cno3h3636n, 0>
51900119, bj)03hoi0oihg30 16 gobnb 33013330 goioi0368o, 0O56o6o'boh363Q 01(151-
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53630033)119 51630(101636390 jo0o3hn 60816-0130636360. 03hj6oi6o 815361)13360
3631150001 3ghoi 3go9ih6o ihoi6, 309)13 610160136360. h01501)115 6360, 6yQnb 0(53-
6ih336o 830(51336 613036 363h50ib b0303oiob 015361(13360116 33gih36ocn; 3hov
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,5360; 606301316011 go^m0ibi, R 33X D ^n3> 0301513b 3ghoi 330 363h50ib, 309(13
Ijbhnon ££1 363h&nnb 333u3xoi& i du36ihnb bb3igibb3i b] 6b^>6 (yii3 n (Paine, 1971; Lieth,
1975b)
pfejhj/tnb
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g i ]hcnp3&6n*6n (M) 9u36ih33&nb bb3*pibb3i 6i 6 nc3 n (Pipp & Larcher, 1987). s6scmfcn3hn
3m6j(j3036n bb3sgabb3i mjibnb 6ah3m9iga363c°ii oi^bco&nbi p i 6iymoj36nbion3nb nb. Jordano
( 1992)
qj*h30Oi3b0m3*6 bp 0(536^301* qjn^m9 ibi; 3hoi0366n*636n pqghm (j>*hn&6n ihni6

363^150001, 3 0 ^ 3 mh0366n*6n 93 hj6 n*6 n 0(536*h336n. hmj,mh|5 656b,
5*6 300 1^36* 0n0gn6*h3m6b 363h5nnb 3im6m0n3h*g n633b,jnh36nb 0 n3 *hcn3 O3-
6001 063, hm0 99 hiQ Sibibp 3mb;)0n pBph^nnb bn0i3hn33 pqjhm g*6:>0n* 5*6-
3nc7iih363 c q)mh0363 n, 3ngh3 3(1000^030 mh5 *6 n'b0363 n.
5ih30m qjij^mhpbnb 5*3036* 01536*11390 363hj>nnb 3306330 ^ 6 ^ 3. dnhncn*Q*g,
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nb 33015330^ 6 ^ 3 $000*^06 303*om 0nij03936oo>. 5*63gob 5(1*9036^360 pqjhm
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bi03hfnqjnc3hn 9(536*1133606 9*p*0O 6000. 6hgo0m3ovi6 5*63gnb 50 -75 ° -b 3 rnhnb
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6*60(501(56001 cgoih9n6 0(536*h3363n, 1101501(1(5 6*6b, (56n9nb gngn 0*h*5ncn 3 6 9 *
nymb 5*63nhoi636xn.
- 168 -
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2 .5 . 3 ai 5 3 6 ih D 0 n i OD 6 ^ <bft16^ G n 3 6 3 6 n b 3 nh 3 3 0 iQD
ah P ierJjiW lX FI& n b 3 6 3 h 6 3 i!>n]>3 Cn 3 <3 3 jiW hm &:)
3 nh330ign 3hmp3ji y ix rn^n'jcn3n^ 9 bnb higoi^nnb &i3my3636ob DgjDJ^hmbi

RUEpp 3395364 0oi0ni6n 3nh330ipn 3hmQ3j(>,nxrT'6 nb 363hj)nnb &i0 mbi30 nbi g i
n0i33 ghmSn Boigiôb 3(101330^ 0O3h 3crji6 cnj0 x n higrii^nnbiîS. 363h5nob 5 >
3my363bnb 3<33Jii3hm6ib, 6333036(103, j>i3mbi^)i336 hm^mh^ Eicrgh 6h3^m-3hmg3j-
I50i80 ^^3930^^3630 363h&nib 3hmi536lj 363 n bhxn q3mvjmbn6cn3'b3hig ij^n^hn
higni|5nnbij>i6 (PhAR).
hiii 333b3&i 6 h3^m-3nh330ig Sh m p jj^n xîb , i& h m jx ^ n b 30(10163680 3 m-
tiihgn C4 3i536ih336n oii3fii6cnn 3hmp3j^nx'Ti6nb 3nj>b i(p63336 3 % -gi 5 (bn0 n5 -
gn) 6%-693 (033^30) 0 yjbn0 iCDhn 3qj3j^3hm6nbib; C 3 0i536ih336nbiOi3nb 3b 9 ifi-
33636300 03hy3nbb l, 5 -2 % -gi 6 (3ihj)mbi6 oii 303(j3bm6i) 2-4%-3q3 (6i0ibm3-
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bi g i bo3h(533o (530*9 3nhm6363o 3u36ihob 803(1 363h5nnb 5i0my3636ob 3033}-
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b3obiyh30 3ohm6368 o(5 in, 3 i&., 63n3 oi6 ^ 336 8 0 , ^36046 4p5 n0368 n 3 0 3 3 ^ ^ -
m6i 2%-'b3 gi6i0fii; ^y336ob 30hs303bm6nbi g i 6iOibm3i6 oii6ibi'bmg)igm3-
63680 ijo - l% -<b3 gi6ioo. bxîQmg 3(10^3^0300 015364(13x1 oiiBibibmî-
gm36i6o (^33360 , 6^33(^939^6601360, j)36ghi) i(p63336 0 ,2-0 ,5 % i o?Tl.SThm6ib.
05033 ooiJ3 ob mj)3i63oni 303ii3bn 6i6n0nbiOi3nb. 0i("i0n ohigoiôob ^hm3nj)3o
(13501^163630 (j>ohfibn6nbi g i cnbnb hjnb ^hni3o i x n (135001636^ boibom33b) 6i-
03^360 3\5nh3i gi, i6ojjm0, 3hmgj)^n3crri6i gi6iQni. 03tt>3ho 81536^330*1 biqjihn
bhx^g 3Dh ny363&b 3 m b x higoiyoib g i 3q33j^3hm6i 0boi0mg 0 ,05 %-b, i6 ô-
64b. S.BS ]6]h5nnb 339^3300164 (mh^4& x n 64b0nh&4Qrib 04h4j,n) 606301346(1 (C). 0ihp366ft46n
(D) 94 3)10103660460 (M) 9u364h33&nb bb34e4bbj4 6460060 (Pipp & Larcher, 1987). 464C<Tij(rr)hn
9 0 1 6 4 ^ 3 9 3 6 0 b b ^ 4 ^ ^ 6 3 4 m j 4 b n b 6 4 ( 1 9 0 1 0 4 9 5 3 6 3 0 0 1 4 o i 3 b o 3 & o b 4 9 4 64y m q j 36 n b 4CTi3 n b nb. Jordano
( 1992)
- 169 -
6 ^b3 0 ffi56 c? 0 U 9nb 3 .bf!S0 W ^ 9 3 6 6 ^ 0 6030)0 3 6 3 &0 b P^6 9 nd 966
933 3ghm 6ij)036b ih 1P 301O361 (6ib. 2 .83 ).

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36001 05033 0mu30m6nbn, hi(3 J3i6ib3nhnb, 6363611030 51'bnbi 91 6i3oim6nb
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03'bob 3hmi53bnb 03933016001 01(1159361 cnooijOob 10-33(1 03^0 363h5oi, 309(13
9P3obicri3ob 0nnb9ih36i 80130 0bmqj0nri3n. 0138^1, qjn^rn0ibnb1 hm^mh^ 363(150-
ob dohooiign 6yihmb, 5i9my3636i 031^0361 0bm0mg 516^3301(13630 3ohm6369n
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31115360b 5300ib0noin 5i6b3ob 030935. ^ 3336 0 , t>ihbo30n 0^30033611 gi 3916-
61^1360 03ib!J3DCD^ bi3336336ob 0i6dn0<b3 qjn^mOibnb 3jb^36bn3hn 0mb0ih3dnb
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oj636i ODmombnEkrg'bnb qjm^mjnQr^hn 3hmu3b36nb 51005361 363h5nnb 9 m0b9 i(i3-
6300 03mhi0n 3hrn53b36nbi5i6, hm03Qcm bi6m0mm 0303501 6nm0ibnb 6ih0rrij0-
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6030103(136*01* b3Cnbiai3nb, 3hr*ig3jl5o30f*i6 ob* g* (bhgnb*oi3ob. *9 *boi*6 , 96(103-
630013^60^ 9 *oio ih* 9brn0mg bi^nhm (1*01936016*, *(1*839 bb^nhm ShrOmhun*^
3 .1 6 n ^ 6 fl hnbnhb W i)3 3 6 n 6n 30n n 3h 3636n b

ey^hn
3 . 1.1 8 n6 3 h X D h n 6 n 3 0 1 fD h 3 & 0 6 n 6 f l ^ 6 9 fl
0063(1*03(10 $33606 303836^360 60*9*530 *hn*6 666*g g* 69 x ojmh93b9n.

6n*g*5nb 6y*09n 5*b66n0n* 96 m0 f*ig 3960936301*1 h*mg36<*i6* - 901300 9*h*5n6
ihi 383^36 0 .2 %-b*. 6001303836^3606 oinaij8ob 98 % 5363936* 1*1115*630
6*hR36369n, 3303680, d630*g 666*g *h*rnh5*6x 6*3(1013690 g* *51130133 8390*6
9n63h*03bnb 939*95360016*00. 3b shnb 6*^3360 6n30in3h3b36nb 113^31130, (101830-
nb ^383606 0n63h*0n'b*unnb* g* 5*9mj*h3nb 3(101(3363606 b3g35*g d*0n*6 6300
^393ncn 0m6n0n'bg36*. 9*11636000 2% 303036^3606* *gbmh6nh3630n* 60*9*506
i)m0mnp3&<b3.
- 171 -
306366^36 6030)0060560)6 300130b366
3. 1.2 6 n ^ 2>an nn636nb êbnhbisn^ qs 153w
oiobob 301001093(10 6b6o0b336o 6* ^30063)10 6030103)136360 aib30b6oio 'bQgiSn-

(1300 3CDJii^Xn &Dboob 6yipoi6noi o‘bogi336 00163661 g i gn^mQ^h 9010333036b
gb 6 bh0 mj06 ob6 33^(533*0 &03&b (nb. (jbh. 1.2). i 3 oooi0 6nigij,nb 30100109360
©01^0393636 (101501(1(5 00160(3360. 9 ioin 0 o0 oi(530ob 361I10 9100130936x01 ijonDh
'b^gi^nhb^, (1010390(5, 6151000119, oio66ih 9o63hio 9m6(39oihnooi60(jb ij3b 600 -
800 Q2^!"1 , broom 330063b 6030103(136^ - 700 O^Si'1 gb ©300. oiob6ih 0063(11-
Q36bb gb 330063(1 6o3cno3h3636b 511660101 îh6o 3bhymogocno 836(30. i0oom0
obo6n, 33nh3XDb yoi30obi, o'bogi336 3101001636b. b0bboib6 3(1009, bhob 15(130133
9193600119 9b03bij30O 366360(3, bigi(5 33bid03 &30 Oi b6om636nb 0o3hoi36b. 013&-
15b, bgbmh6i50b 93060019 6936b 3101001636061. (101501(1(5 63bn, 333013619 0no{bng3-
6b 9bC?bp3b036,yib6n 0016360. 9i5i0oong, Ca2+ 9nn'bng36i 33301, 309(13 K+; 013
3(1010 gb 05033 31036,301606 00163601, 9 i 3 o6 33301 9oo‘bog36ni6 61303619 Soghi-
00(136x0 0016360 b63 obn6o, (101030011(5 Bypob 3(3(101 016390 5bhbn 13(11301. b0
601(16(50300 60360b 6yb0oi6ooi, om636o 9090 (1101936016001 5(10130361 6y0noi 51-
30(1333630 60661(1 9b 330063(1 6030103(13636^3.
00163606 0(13 3(101531(10 „03b0b3b0Ob“ 0ybh 6obgb53h cgi'bibi gb 6mgi5nb
bb6bhb 9 oihob. 013 6obgb5ob bb6bhb 301(3361 b6 1309361 hm030O03 om6o, 0 i 5 o6
b95O0O bj3b 0 n0 mi530ooi 3hoi(53bb. b0b33 ghmb, 3qjhm 0 ybhbg ^01(160(136300 oov
6360 3oi0mognb 0O3h 030*6 0oobog3&b 9b 3 obo ‘639130(10916 5b0mo933&36b
61303619 bmh6nh36b90 0016360. 310100163690 bmh6oh3&ob 60 nb d*0* 3036x016b
8309350 0o0bhoi3036ooi: A P + - Ca2+ - Mg2+ - NH4+ - K + - Na+; bm0m 1600163630:
PO43- - s o 42- - n o 3- - ci-. 0do03 ©30*03606 0016360(5 bgbmh6ohg36ob6, 0136(53
©bmQmg 03Obg 0(5oh3 (1101936016001.
0016360b bmh6(30X 60 bb ij3b hb0936o03 919360010 0bbh3: 5i0mjih3obi eb
3303bob 918906 3 x 36*9 6*0moib3nb3q)03630n bb333&o 6030103(136360 bh 031(153-
6b 9b g*l330Ob 5*0mh3i5b3ob*5*6; 6obgb5nb bb6bhob 3016(5360(11(501 (16361 gibbon
91 5b6m6bb6mh36xo 9b b0 oom9 , 033633661 gb 6obgb5ob mh5i6o'6036b bh bfocn
mb0oib3hn gb03nhoi336o, 0130(31 bbôhm3&obb036h 0016360 qj3b336obioi3ob 1930-
019 b300obi63gm0ni.
60191506 3010010936^ bb6ihbi 9 1 60191580 0063(1103)1 6030103(1361011 0ihi5b
8mhob ihb36m6b bi30im0 hoi30o 0mdhi3O 6m6ib6mhm6nb 31380(10, hm030bii5,
o0 i 39 hmx*Q. igi3o*l5nDô 5ihgij06ob 361(10(5 511R601 g i 0m6mg3630oi 0i53&ihnb
0063(1103(10 6030103(13636001 36y330o 0m0ihi536obi336.
3.2 an63h^CDhn 6D30inDhD636nb 33013flb36:)
Bypob 0(5361(13360 bi333& 6030103(136366 0^3636 001300 0113060 t63gb3ohncn,

0i8o6 hm(5i 663030106 015361)133680 3b 601)1(50309361 0 ^ 3 0 1 1 bobo30nb 83833m-
6001, hiy 0336306 3hoi-3hoio 38003(1360 0336^011 (b^&bohioto gi0i5h36iboii5
3hoiig). 9 o63hi03h 6n30i03h36b0ib 9bm0 mg db0Ob6 015OI13 6b6o0o 6 3 9 36 1
0(536bh33n (*>3(1016 t^giôhob 5b30ooi.
- 172 -
00636^ 7)60 6O30)O36O53J>Ob 9a0)30b056
3 .2 .1 Wi)3D6n 6n30nn3h36D6flb 301^6009^ 6 f l ^ 6 n e i 6
cgpbpn 6nipij,npi6 npp&b bij^pb {jnpomphpbpbb bbsigsbbpi bphbnon. pb

93 j} 6 n<b0 p&n 6iR3p6pbn:> 6ib. 3 .1-“bp:
1. nm6pbnb 9 cni6 cnj0 } 6nigi5nb bb6ihngi6. pb nm6pbn ig3ngig bppQnbiSp-
gm9 ni, 0}j>hi0 3n6ingi6 bnipi^nb bb6ihnb j,m6i5p6^h:>uni 9 p^>g gibicni,
nbpotn nm6p6nb ap0u3pemb:>, hmfcmhrnjii S 0 42", Ca2+, Mg2+ ©jjnhpi - 2 -
5 mM, NO3' -nbi - 5-10 mM, K+ - 1-2 mM g i P0 4 3 ' - 4 |i.M-6gp;
2. bmhbnhpbppn nmbpbnb dndm|53Qncrin 9 ou>6 cnj9 i. bpboifcnb Spppfcip qjpbpn
j>i9rriyniqgb H+ g* H C 0 3“ nm6p6b, hiy bpeb pBymbb nm6pbnb 8 m0 mu30 ib
qjpbpbi g i oinb6ih g i 3p9pbm3i6 biBnpi^pbb 9 mhnb. i 9 pj(i6ibj)6pcou>-
5^6 qjpbpn H+ g i HC 0 3" nm6pbnb 6i^30ig nppbb bi^nhm 6n3<nnphp-
bpbb (Bib. 3 .2 );
3. jn3 nphig bBppn bi^pbn 6n3omphpbpbnb 0 m6 npn'bi(jf>i. pb bnth(jnpQpp6i
H+ nnbpbnbi g i mhj)i6p0n &ji3pbnb 5i0mymqjnb biSpiepbncn. 3 n6 phiOp-
hn i33&nb jn9 nphig 60 ppn XD^D^pbn p6n6ihpbig 9 dn0 p apîppbrti,
hm9 op6 nij ai0moii3nbpq9cgpbni6 g i £,igigni6 bpQi^ph 6 iphonpb9 n (6ib.
3.3 pi 3 .4 ). 0pî0mbpOi|jp6n bi6ynb 0 gj,m3 ihpfn6 ib ih pbhp6Qpbr»i6. nbn-
6n igpriQig 3 <nin6oij8 pbni6 gpbppbnb 0 nph. H+ nmbpbnbi g i 0gi3pbnb
5i0myfTiq3i gi0pii)ngp6pcni bp6oij3nb n6(jp6bn3mbi<bp g i 3 pbi6 i 0 nbig,
gpb33bnb gi65&ignoi amQihi^pbibp, 6 ib 9 nh6 ig <b3 gi ^pQiphi^phi'bp.
Root hair
6sb. 3.1 6nigi59n 9n6ohaCDh 60301(1^36^ o H+ and HCOa

Qi 3ioin a30)3nb3&> a x p y k m Mineral nutrients
3r>oh (Finck, 1% 9 , 0men<3 ntjnh3&3cn Gibibn) ^ Released organic compounds
- 173 -
8 0 6 3 6 6 ^ 3 6 6030)036066016 3301301356
Ebb. 3.2 nm63&nb eD<2JnlSf\y*> &>3Cfes

yi^hnb 0^33609^ H^-nb BnsesfcSn
5jpib30nb fp'bhpi'b]. <33^300 &sh93dm
j> 5iO>36x n 8030001360 pH-nb 4 .5-
3g3 46 3qjhm ^3331101 9^1339^3
(Kramer, 1983)
fob. 33 6n3cnn3h3&iou> 6n9rnj3Qi 0)363368(1

(M arsch ner, 1985, 6 s® sNbn33&X n ^jo&O
OH
OH .0
2 Fe3*
+ 2 Fe^ + 2 H*
HC = HC—COOH HC = HC—COOH
C affeic acid Caffecyl-o-quinone
c N (CHaCOOH)s
N (CHj COOH)j
HC— NHCH.
I
COOH
6sb. 3.4 hy>6ob 6:>3ho<36nb 339unh36i
Cb3Qi 6ibiVi) gi br6oi3|bob
S ii^ C n o ijb n ( J3 3 9 1 6 > b iV i) (Iseim an n ,
EDTA EDDHA 1980; O lsen e t a l., 1981)
- 174 -
8 0 6 36 6 ^ 7 )6 0 6 0 3 0 )0 3 6 3 f> 3 f> 0 b a 0 m 3 0 l» 3 J> 6
3 .2 .2 nn636nb 90b 30i 3XhD93fl Q} Q^6hn336i
3 n 6b n p i 6 0016360 B y o n b 5 ih b o o i i h o i 6 J,b h 330 m h ^ y 930n, 9bcnn 83630! b m l 5b -

*C 33 h 3g 3 n 5 ^ 63036300:1. 6n o i 8336h i 6369o 3 i c n n ^ h b 6b 3m h^nh 36b n m 63hn ,538-
6013606 b i 935036ocn 6011115030936;) (ijhnjhjpn bnb^jds). 938935 o b o 6o 3j h 3g 9 o
c g n jb o h 36x S m 'b o ^ o ib 0^33636 {nmbjhn shbjdn gs ^nhidnb (jnpjdn) g i
5 h m 3936o i 6. i h b 36m 6b 0016360b 3 m 3 m [5 3 0 o b b i 8o 93b i d 0m 5^ ! : 1 ) 3o 6i o g i 6
0016360 3 C 3 J b h 3 0 i 0 g i 83b o 30n ! , nm 63h o i h b 36ocn 3 ic n o 3 3 3 6 h i 6 i 3 o 83630^
93b i d 03630O i S b o io m g 08 933cnb 333i 9o f 013 o 8! 39h o i 3p i p 33 h 3g n g i 6 558013:)
b i 6o 6i i ( j ' 8935r i g 95836^390 om 6n {tjm-^hbbb^mh^n ±6j bnd^mh^n)-, 2 ) o m 6o
3j h 3g 9 o 5iO [ 5303& i 08533 836,50b 8jm 63 b b 3i o m 6'b 3 (b^nhnb^nhm ^h^Bb^rnh^n
s6y bb^nhmh^n)-, 3 ) 01350 3333 i h b 36m 6b 3 C D j6 ^ 3 0 3rn ^ 36^ n i 0c n i b b 3i o i 6i 838-
6h i 6n b 5i h 3 g i 9 o g i <b 3g i 3n h 36b 9 m h o b (Ssbnjhn ^h^bbimh^n) ( 6i b . 3 . 5 ) .
53h o 3h 30 b o b li)33 i 9n i g 5 o 036o , 6595^ b g 36i 0016360b g i 8i 5h 36i i h n b b 33ijn -
q g n j^ h n , ym 330 838016333580, 36 59509360 b b 3i g i b b 3i i 56001636065 g i 550100163-
6nbiOi3nb.
33^13980 00163606 i j ^ n 3h o ^ h i 663m h o n h 36i g i ^ i 39o h 3630o i 303Jii)h m jo 8o 3h o
6h i g 036ljn b 5 i 9i 0i b 3i b c n i 6 g i , i 8g 36i g , 01601390166 363115006 55)133330 g i 6i b i h -
336b , hi( 5, o i ! 3n b 8b h o 3, b 36c n j 3i ‘63i g i 8o i 3o g 3630o . 580,5018 653336 603010311365-
P lasm alem m a
120 mV
PD C atio o uoiport
PD Anion uniport
'.'a a:-,- 10-30 mV

AD P+P
1 Citrate Vacuole
p H — 5 .5
Anion-
cotransport
Cation-
exchan ge
C yto so l PH
A p o p la st -pH 5 .5
6ib. 3.5 91 nmG^bnb ^hi6b3mh^nb Smgxn (Pitman & Luttge, 1983;

Kaiser et al., 1988; Martinoia, 1992)
- 175 -
80606^36 6030)0 3 6 3 5 d0 to 330 )30 L3 to
cn> ij^n^hn 8oii6oij0i gi8myig3&3poi n9 3501901503(1 opijfnhQb'bx. hmOcQftnu

3ohgi3oh g i oho6ig 001^0393636 6360^306 n6£)6bo3fn&i,b3. 1560160901, hm9
b36cnj3nb 06^36^30161 n'bhg36i 0016360b 1^03)10 ^hi6b>mh^nb ghmb (*9ihnpnb
b36oij3i"). 6nm0306hi6360n ATP-Snghmpn'b^hn 1509360 (83861116390 ATP-i'bi)
qj36junm6nh36b h01501)115 5htn^m6 x n ^306m g i i8yih36b XDJ«il1fnjn0n3h &hign-
36,5b 0306hi6nb 51663)103. 1^503)10 8o8mu3oob 03ji6o‘b036o 8oij0x3&36 Shoi^m-
^giW ob 00139b 51113 bitJCîM ^ (30i‘b0ig30i8n g i (5m6rOgibti8n) g i 6nb
0056001 - 5039516, b igiy 50 69361 0016360b ^hi6 b3mh^n 501031)1,5036^365
0mhob.
nm636nb ^hi6b3mh^nb sfon jh n p i 3*bnjhn 83^160^83606 3)101016900161 5 16 1-
30(1016366 bij>336n 811109360b 9on6oij0ob 030 g35 g iS ib ib o io ^ b x oii3ob3-
6 3)136 36 ^
i|O03OilSn3hn oi3nb3&i. 815361)1339 33)1396 ij3b 36iho 5016(536^)11150300 5)1190-
36150b b i 3 ohob3nhmg 0oii6ovnb g i 3*33J(J3hig 9115(10130^ 0016360, 516615301-
(136001 315301900. 3b 01306361 83^19 06o8363C<T»3i6oi 6ygob 615361)1301101305,
3n6iogi6 0itn 51)1380180 i X nCD&D0O 3930366360b 0bmQfng ©(50)13 5016(536,5111(5011.
03hK33fXJim6i. 0(536ih3x 3jh3gb 03b633b 361)10 0on6oijoib 66019019 nb 00163-
6n, (1016936015 0ib 516612)301(136001 361^01101361. 6151900119, 160016360116 6 3 9 ^ 3 -
6non, nb 33ohi,53boi6ib 160^366 5101001636b, 601901 51010016360916 50 *601503)10*1
5(10139361 51(50936001 63^0 (1101936016001. 3b 01306361 0(5361)106 qjn'bomQP^rQh
50166,50,53(503(1 ôiftî gi0ri5og3639o. i0 0bho3, bb3igibb3i bib3fibi 3)1018163-
cnnb5i6 853301(119 5160116331.
03hR33no¥n6nb 9080^1. 83hR330oim6ib 5116 6 0 1 6 ih 5 3 3 X n t>p3iho, hoi8c*nb
njncniu 33)13911 33(j<ih ibmhun3Q36b i0 <336^016 g i 0ib0o 5)1013936016 obtain
0016360(5, (10189360(5 ih bôhg36i i 6 6 15936 :^ 5i0moy363&b. 3b 5166153011136ncn
8 3 6 18 6 6 330 1 9109183639 ^160,51,53680, bigib 9sho036nb 5ih 3 5016(536^(11(501
n 0 g36ig 8 ip i9 0 i, (1018 35)13980 5hm3936i Na+, Cl* g i bb3. 0016360 8 3 ^ 9 gn gn
(1101936016001.
3 .2.3 q}Db3nb ana^hi636^ nn6D6ncn
6 15 336 0 6030103)1363606 6(5361)1380 836390b 606^1(13 91601509363901 <3336301^

ih30n 6063)1193(1 6030103)1361011 5016(536^)1l y o i ^ , 51)1533390 0016360b goqj3<bn n b
6o6jih3-
6 ^ 5 6 3 9 6 0 , 916190 go q o V ^h o 5013050(5036,506 516^1, 0mdhioi636 5H5O036ncn
6 391.
0)36306 803b 6153360 6o30i03h3&36nb 63013063606 3<33j<53)ioi6i g i 51)1,533:300
ooi636obioi3ob 330(11^3601606 6060^361 8)536ihnb 5363*50531119 5i6 3o h m 6 36 30 n
oi3ob36ii. bij)336o 6o3om3h3636nb 9oii6oij8nb bofijih3 gi8oiyig3630oi nm6oii>
83hR33noim6ob 3hn6i5n3b3. 0 336 336 0 ^ g i dicno 63b3b36ob bnb8nh3‘b3, 6 o i g i 5 3 n
im6i536^hii5onb 5hi9036^,‘b 3 g i bb3- (6ib. 3 .6 ).
- 176 -
g d ^ & o tin an tfo w .33360 G030H)3 fn3 &36 ni>
6ib. 3.6 b>(jhi6b3mt\yTi 3jh39j6n Helianthus annuus-rb hn'bmg]h3j 3 n hyi6nb Q^qjnun^b 3nhm&3&8 n
(Kramer et al., 1980)
90606^ 7)6 6030)0960560)6 300 )00 b056
3 .2 .4 fin636flb tMrtBbSFlhiMl <3Db39n
6nigi&nb bb6shngi6 bi»>33&n Srôin^h^&^&nb 01^16360 63936(1^6 g^bsnb jjh-

jnb 3ih36jn3i3n 6yi0Oii6 3hoiig gnqj3(bnnb 338331^61101. nj nm636n 5hm^m3pib-
(jnb 5*h3 bi'b[p3!htb3 ihb36;30n 33b(j36nb bi83i03&ncn &igigni6 6630 3 g&m3sh;|-
m6 i 9 n. 3b ihnb 63n6 pi îbn^hn 3hmi53bn, hm03cni5 &i63nhm&363cni ynB^b^hi-
Ijnnbi g* 3C3Jii^30n 93bli36nb j,hign36ljnoi 6nigij,nb bb6ihbi e* qj3b3nb 3ngi
ih^b Smhnb. qj3b3nb <y>h;j 'bmBib, bigii} bg3&i 5 ^ 3 ih3boii6 gnqj^bnnb bi03>
03&noi im6i535(!ihiiin36nb &ioii6i6h36i, 36 1^ 36 ! "bnpjgn oii3nb3qjicn ih3“. 593-
6ncn gi9oj)n|3363cni1 hrn0 i 3 lbm6i8n bg3&i nm636nb gngn himg36m6nb 303^ 3-
gn 60 i (5 ib. 3 .7 ).
8oii6ov|0xn nm63&n (1693636 oii3nb ftbib 053b3nb "bnp^g oi!3nb3q}!0 ^380".
nj nm636n b3g3&ni6 3ih36jn930n 3jh3936nb unof,ri3ci'b 3 i 3 n gi 5>igi3gn636ni5
33hpgnb B3360n. 3jh3gngi6 ^jh39811 nm636nb ^hiBbimh^n broh(jn30g3<!>! SQb'b-
3 mg3b33 &nb bi83!03&noi, hm0036n|j ij)i33nh36b 3 m3 n36 i 33 33h3g36nb ^n^m^Qi'b-
9 ib. 3ij)3m036n ibh3Q36]6 3 bmgmg nm636nb gij,hm336nb qj36j|jnib gi ihi6inhig
0 m6 i 5 ng3m636 0iOi 0mj)C3 gnbîBijni^ gpgiSm dh^îSn. 3i i 3mQ363n S^b^gn
bii33&n 0ihng36n nj33 hfi36ni6 3i6i0g3, bi6i0 ih gin Byî 3ioin i j ^ h n
6igi6ii53036i (jntjjmSc^îSn. i0hn&ig, dm^pj gnb^b^n^bj ^hibbimh^n bnhyn-
3063 &i 3ij)3ri036nb 533hgnb i3gncn qj3b3nb bndScsbtfnb [ 163 ] biS^^&noi, i6^
limybicn 3hrnljm30ibli 36nb □By33ljn ji^ n b 33333^6(101. bn330 ib^3hn 5 ^ 36gn>
g3h0nb 5*30001 3n30ihoi36i u^^hip^hn (jncn6ghnb*i36. nj 3ih36jn3xn 35h3e 3‘
6n im6(jij(inh3636 6i0îh 3g3036o36oii6. qjgnvjQnb D3li3g36n nm636b 1^ 36 36
bnOSgiboDhn ynBoijonb bi83ig36noi. 690001 833b3& x £ W C 0&9 n 6* ohij3nP363 n
nm636n *063336 hen^mhu Sibn^hig, t,m6 (536oh!ijnnb 6hiQn36^nb biqj3d33c'b3,
ibQ33 iji^n^hn ^hi6 b3mhonb ^bnoiiy.
---► Apoplastlc transport Q Carrier system

— ► Symplastlc transport
fob. 3.7 qjQbjnb (yien6ph3 n bb6ihngi6 nm63&nb frSQ&myyjbs {>hd x

SiSdnctD frigid (!>i6n bnb(j]anb 93333016001. V 34jomen; C un^m So*^; ER ^ m S e i 'b a ^ ^ n
h>jniXD 9 n (Luttge, 1973; Lauchi, 1976)
- 178 -
8 0 6 3 6 ^ 7 ) 6 0 6030)0060606011 300)30l>0&6
3 .2 .5 a i5 D 6 ^ h D 3 n 0 f l6 D h ^ C D 6 n b 6hd3C a ^ d l C t t ) ib h :> 6 b 5 m h i!> n
jbncp9 i 9 n l»i>33bn 6030x13113636(1 3 np9 >h01360^6 i(r<9 >3iQn ft'bncn 0 ^ 3 0 9 * 6

P3hm36nbi36 tjhi6 b3nhiunxn 6h^>pnb 93933(^6001 g i 6i6npg360^6 80130 0yp-
6ihp9 n. ^3h^pm3i6n b(ib(t>39 nb bi6m0fnm 3p6j(i)369n 001636(1 jp 9^phn $(n6p6nb 3>-
hp6 ;jn0 i 9 n gnqjp6gnhp6p6 pjhpgnb iPeCD6ngi6 Shm^mîb^pbnb 'bpgiSnh'bp, bi-
QÛ nbn6n ij^nphip J,i9mnymq3ni6. 838935 nb33 bn03 Qibg}hn fcVioi b(ntnjnp0g-
36b 9 mj)03 gnbî6|jni'b3 ^hi6 b3nh^n g i bij)33& 6n3omphp6M7o &ih>>33xn 6i6n-
0(i nb33 gi8ihi&g36i 3si)3[Ti0p68(i. bi$33& 6n3oin3h36ioii ^hi6 b3mh^nb j i ^ S n
9oii3ihn 8i0n8n^nh3630n cgij^mhni qj3b39 n bnOîb^nb 5^30001 nm636nb 5 >
giip5(i036i; hm8 ih i pb 9i0n9nlyih3630n qjij^mhn, ^hi6b3nhiunxn 6i|>ign 5 >
150036001 63^ bi.5336 0ihfi0b j>ipinî6pi.
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q)0(Ti30nb hm0fi. &hd3Q gnbî6uni(b3 ^hi6 b3mh^nb 3b mhn ^ b i - jbn030i gi
cg0m30i - 0u36ih30n 9hi3i0j)‘bnb 9 mpnb 3hoi9i63oioii6 im6 îj^ 9 n (5i6bij)3on-
hpbncn qjpbpan g i pphmb 03bp369n) gi 3hoi9i6poioii6 gij>i39nh3630ni. 0mb0ih3-
6nb ig5fi0369n ibn0n0i^36oii6 3hoiig 8(i30ihoip6i q)pb3(igi6 836^13(10(1 8n63hi-
pphn 6(i3onn3h3636nu. bb3igibb3i 6(1(^1303836^(1 3hoi6inhig ig3dgig ih j>ip>-
9ndhi3g36i (6ib. 3 .8 ; ybh. 3 .4 , 3 .5 ): 3b 303836^36(1, h(n00p6fH5 030g3j,9n mhj)i6-
X qjfrih9i 9 n 5ihginj06 p6 i (0 i&., C, P g i S) y-hfcig ^hi6b3mh^nhg36i, nbp33
hrn^Rhu ^ n b (101636(1 (K+ > Mg2+ > Ca2+ = Mn2+) g i Cl", upgig j>igi0rid-
hi3g36ni6 0dn0p 03vj)i0p6n g i ii0ynp0n, hm0p0ni3 i0n^rn0 pb3ig 0ihij>g36i
gmoi0369n (fchdx gnbî6uni,b3 jbn0303hn j/bnoi ^hi6 b3rih^|nb 6m0rri Bph^noSn).
Bib. 3.6 Lupinus albus-nb (3363^16 C’3hm3g3 K +, Mgz+ p i Ca2+ ^hi6b3mh^n qs iej,np9m6i(}3C3m-
&:>. (jnqjho&n 33(1368(1: nro63&nb i i 39x n*1D&i W ipnb Sjhnmpan; W>3s (jnqjhjijn: jjsSmymqjnb bnRîh]
(ji 0mcn nm6n 6*1"1 6360° 5m6s bor1); Xy^hs6b3mh^n jbnQj&ib bi33sc3&nai; Ph li>hs6b3mh<jnh3&:>
qjcm33nb bs33so36noi PX 133b39*1 <3(^1330916 J?nc33i3n ^sîbx^ (Jeschke et al., 1985)
- 179 -
30 G0 rf»^c v'3 6 603010063660)6 330130U366
3 .3 a i5 3 6 ^ h o g n a n 6 0 h^cD h 6 n 3 c n n 3 h 3 b ^
9OD6Olj0:> QydMbObS
3 .3 .1 93h 2)C a^bân 6 ^bhnb a o a i 53 D c n 6 ^ e:> abD 6 i h 3 3 c n

6 i l 5h n b 9 D aê 6 3 6 C r i 6 i
QncnmbcgQhmSn 33843490 y3304 jn9 n]hn 303836^0 8mo3m3364 8)5364(1330

6*154(180. ubh. 3 . 1-3 n 9ri(53830O4 3(5364h30i4 bb34g4bb34 353030b Si^hnb 838(533-
0m6nb 3o3mbnQ34. 64(54(180 8063(1493^1 6030103(1364014 bi3340m 333(53391^64 3 ^ -
1538300* (5bh. 3 .2-8 o. nb30io 393836^360, (11*1501(10(544 N, K, Ca 9* ‘bmj^hcn 8(53-
64(1380 - Si, 3i536ahnb 64(54(180 91190 (1401936016001 80103013364 ( 10-50 5(1 j ^ '1
88hi0n 60160b); Mg, P 9 ! S-nb 838(53390164 944690136001 10 5(1 i 6 ' 1-04, Fe-nbi
- 0.2 5(1 jjj,"1, bb3* 'bn5)n3hcnnbi $o sh 4(^384^364 h43936o33 85 36° ■ bXMOOOn
6nm303836(i)36ob 11401936016(1030 oi464q34hgm64 8^3301)149 g484b4bo4cn3630n 6n-
845-oi3nb3644 (101501(1(5 b^CiDXf1 b4b30i6nb4 94 oi34bob4oi3ob, 4b333 3(5364hnb
bb3494bb34 01(15460^40130^ 4(^6086390 0146*0341190164 0(539364 3(5364hob ibi^nb
83b4648ob4g. b336o 94 63^360, hm^mh^ 63bo, 830(54336 83^ 4‘bm^b, 309(13 3490-
38b (3b 546bb3433636o 546542,301(136001 oi3408nb4i538o4 b364hj(i)oi,39 b4b3m&3-
68o); 6404bri346 0i5364h3368o, 38363b 0380^333480, 3490380 b£>h6oi6b 4tbm^b.
546543301(136001 g404b4bo4ai3630o4 01464034(190164 Ca : K. Cariophyllacea-9 n,
Primulaceae-d n 94 Solanaceae-8o ©3^04 3490380, bm0m Fabaceae-Q n,
Crassulaceae-d n 94 Brassicaceae -30 - 349(50380. 84(1(53901363680, ob0368n, 34Q-
83680 94 830(33680 Si 030(04, 309(13 Ca. 8401 64(541180 Si-nb 830(5339m648 83nd-
9364 801390 04bnb 2 /3 -b 8o4(p6omb. 9046018311 6y498(5364li33&8o, (101839014 343-
8460 iohob 83434b 8301543b, Si-nb 838(53390164 90 %-b 4^4(1636^
8(5364h30i4 30363bm64 3031101 836 gmbcgrahb 8301543b, 309(13 50150(19^ 0138(54
4hnb b43nhnb3ohm ©45*9001360(5 (B ra ssica cea e-rfo oi34bob 6*h0oi84g536936n).
(!>hm3ri3x ^>3014 0301019360 830(54336 546b4330ih36001 9090 (1401936016001 493906b.
946019016, 4hb36m6b 9i5364h336o, 110189360(5 5481^0^33046 646(103806 34(ph3b.4Q
9090 83815339^6001 (R333936ho3, 815364(133680 64li)ho38nb 838(53390164 d491146
8(501134 94 3b 393036^0 838(5339fTi6nb 063(10380 86019019 803(10139383663606 6n-
644 64694536x0)- 4630136b 803330136364 9 4 0 9 4 8 3 6 x 60494536*63 Sm'bihgo 8^53-
64113360 (945., C h e n o p o d ia c e a e-b , 45(130133 B ra s s ic a c e a e - b 94 U m b e llife ra e - b 6:>h-
8018*95369360).
64(5hob y3394“b3 89094(10 838(53xi,i6oai 549010(16334 qjoioirxo, 601901 8153-
64 hob 5433 hj63& xo 64609360b 64(54(10 - y 3394<b3 e>4ho6 o4. <3010101980 5(101393-
64 Ca, M g 94 S, br»9m 84(1(53901363680 94 ijhrôô 633680 - Si; 934309360 94
6490103360, 30oi43h3b49, 45(10133636 K, P 94 S; 63014 £>31101360 8394h36ooi 83,30
(1*01936016001 830(54336 Ca.
64(5hob 83049536om64,b3 (54(194 013001 0(5364h3Oi4 633(5003033(10 01306363-
6 ob4) 909 5439364b 46936b 45h30i33 Ib&o^^Sn 8 o63 h493h 6030103(1364014 64(1.4-
50. d49046 I3>4hn6 6o4g453&‘b3 (54664330^36001 83433 6o4g4536'63) Sm'bihg
8(53641133680 6415(106 838(53x^64 33h49 84b48o 83^49 8(50(134 94 1-3 %-b ih
4(^384^364. 05033 ncnjSob 330030^36^13. 3ohojooi, 89483 60494536133 801^4(19 8(53-
- 180 -
803666330 806066 ^ 7)6 6030)0060560)6 30 )660)d86 ^ 6 ^686606056
Ubhnpo 3.1 8(5361113011 bb3igibb3i 36oqj9n pi 8oihmoih{(i6o'b9369n 6ib9nhnb 01831001 838(53300161

(%) (8(113100 i3iynhnb 806393001; g^ yiC D ^ nb- Pipp & Larcher, 1987)
5>b3nhnb 338(j33et'>6i % .
oiiemgnaj&n
61^3(10360 8 - 10
boi401360 7 -8
3fioi6Rboi 3pi6iiim63hn 6y>p9i536ih33dn ~ 5
goi(joid36o SO-8g3
bi3b3&n 2 - 4
&!Qib(n3>6n imh8mqjn^3&n
8g30oibi gi jynb 6x?ibm363611 6- 10
81(1(5300136360 gi nbQQbn 7 - 10
53mqjn^36nb 80606^30:1 6160036(1 5 - 10
53mq)o^36nb 80606)3381 6i6oo36n ~6
L>3iX36vl36nb <301010360 10 - 20
^3iX36tf36ob 1^3(101360 10 - 15
310010)0,5360 10 - 20
(|)36ghob 610160136360 - 5
îm6ob 8(5361(13360 5 - 15
6yoob 8(5361113360 5 - 12
3^0030^360 ~ 3
83(1^60160 8(5361*13360
<301010108(53360 63360b! gi 63^^36nb <301010360 4 -6
81(119863163 b336obi gi 63ft;i360b gmoxxj&o 5 - 9
<301010013160 b33&obi gi &3fij36ob imh^jbn
gi 51(13 J3hjn 5 - 8
<301010013160 b33&obi gi bjRj^&nb 83(1^160 0.5 - 3
6063013160 b33&ob 6063360 - 4
6063013160 b33&ob yTihlj3jbo gi J3hJn 3 - 4
6063013160 b33&ob 83hji6o 0.5 - 0.7
0016360b <301010360 ~ 8
6ih3369n 99hiQn 9 ibnb gisbcm^bncn 25 % 6!(j!ho 83!pj,36b. 0 icn 6!(5!h9o j>o

fti'bhgnpro Na, Mg, Cl g ! S-nb 939l}3Dcm6i. h3g3h!p3ho 8(536!h336o, hmSco&nu
o'bhp36n!6 b!>)336n 6n3cno3h3&36ncn 9 gngbh 6n!g!j)36'b3, £>!9mnhR330!6 Na-nb
3 b(pbpn 938(3330016001, bmpm 8:>cnn ^jh^gnb 63360 9 gng!hn! 6n(jh!(i36oai.
9n3b39i3!p nQnbi, hm8 9^36ih3b ifob 36ihn 93hR33ncnip 9300i3nbmb 3b! 013
nb 303936^0, 93hfi33noim6nb 0n9n(|jnb <y>9 m, bb3! ihb3&30n 303036li36nb b h x n
nj,6mhnh36s 9 in6 (j 33b b3hbg3b> gi s9 n(jni9 0U36sh33n bb3!g!bb3! gmbnan 8!o6(3
9mn3m336i y330s nb 303836^0, hpi03pn(5 pi9ib>bnson3630ni j)m6 ih 3li 3Cn 3!60(j>
(5nbcn3nb ((jbh. 3 .3 ). bb3sgsbb3s {,31^0803!™ 9 ^36^3300 353030 g!b!bn!O i36xn!
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036phnbs Q! ^n^nb), !b 333 606! ibnnb 3pi66m369n 8mb!hg b!b3R&368o; *>!0-
13039001 9gng!hn! 83 h!0 n b3&(!)hm3n i x n 6^ »jhf',' 3fVMOn h3j,nm636ob 6n!g!536n
gb, 93bi6i9obig, nj 6 ! 3hl3303^ X n 9l336ih33&n, * ^ 3 0 13 3 ])nhj3n!636<b3 8m‘bihgn
8(536^3360; b9303cnnb 8(536:^33690 bo0o(5o38ob 9 i(pi0 O 938(33301^6! p!8ibsbn>
0136300! 3i3!(jfnhxn bm6ob 8i(536!h336nboi3ob (ohmSoixn i!>y33&n> ^3^6360).
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- 181 -
a06 0 fn^c5 ’3 fn 60 d0)03 (n32>60)6 30 U)30baiW>
(jbhncn 3 £ BmpifcSn p i b03O3<nnb 0|j36ih 33&i}n 9n6yu>C3hn bi93iem 9o0b3X fr* i

(jh i^-l 09hion 6m6i) p i 9n63hi(v)bnl>i00n 3mcrtbm36nc3&i (Epstein, 1972; 1994; Altman & Dittmer,
1972; Baumeister & Ernst, 1978; Bowen, 1979; Lieth & Markett, 1988)
OSe?.‘j9o£>C>0 b*r3*1ji.e;>r>(o 3c>ir>t>p>;ib<>^;)2w>

fe> 5oa^‘3o % 3&><<>33&'3o
Si 330 0.2 -1 0
Al 70 0.04 - 0.5
Fe 40 0.002 - 0.7 - 0.1
Ca 15 0 .4 - 1 5 3-5
K 14 1-70 5-10
Mg 5 0 .7 - 9 1-3
Na 5 0 .0 2 - 1.5
N 2 12-75 15-25
Mn 1 0.003 - 1 0.03 - 0.05
P 0.8 0.1 -1 0 1.5 -3
S 0.7 0 .6 - 9 2 -3
Sr 0.25 0.003 - 0.4
F 0.2 0.02 -3g 3
Rb 0.15 0 .05-3 p3
Cl < 0.1 0.2 - 10 > 0.1
Zn 0.09 0.001 - 0.4 0.01 - 0.05
N i .05 0.005-3e 3
Cu 0.03 0.004 - 0.02 0.005 - 0.01
Pb 0.03 0 .02-903
B 0.02 0.008 - 0.2 0.01 - 0.04
Co 0.008 o.oo5-ae 3
Mo 0.003 0.001-003 < 0.0002
Sib. 3.9a-e 0cni3ihn 3n63hi03hn XO&jfaijbrk

(9 i^hmx 39]^ 63^<b) dnhncnipn 3hm3mh[yi]6 n
gmonripSn. a ,jhm3ni3pn 63(161116(1 (jyj b
'bmSn^hn bih^ycnb 9]b 6 ihn; c 'brndnjhn
bih^ycyib ,j36n;>6n Sibnj^Qbob &ipitm3i6n
9(536ih33&n; d SifyTiqjn^jbn: e cni6 i 3 nhns6
6nigij><b3 Smtshpn 0(536^336(1. Na pi S 6ih-
9(ne636ncoi shcnbi p i 06133 C3hdb3. 9m6 n j 3 0 3 -
6n dmy3i6neni g ifc"1 93hip 6m6i'b3 (Boukhris
& Loissaint, 1975; Klinge, 1976)
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bmqjgnb 033h63m6!8o - 33bid0363po! 03nhfi3b bib3job 3b! 013 ob 3!hoi6l!)fi. 33-
03h!g 0m(b!hgo 9(j36;>h336n (6!ijhob !6!0o(bob 83g35ig) 83bbd03630O!
IJ363630 nj63b hmj,mhi3 0!g6nb b!6!gm36ob 3hcnj,3!hn 0600^0^360.
3 .3 . 2 a n 6 3 h^C3 hf1 J)3 3 6 flb 3 0 3 9 3 6 6 3 6 ^ 3 9 nCJlbn 3 6 nC3 6 ^
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0!bob b6h!ogn g!5hoi336!. y330!“b3 96o83630oi3!6o 3038360360 b300ob!63gm6n
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h3&ob 3hm(j3b9o 9o63h!03h 6n3oio3h36!Oi! 03oi3ob36! g ! 0!oio bb3!g!bb3i>
oih5!6m8o fi!hoi3! 606 3b6h36b 6ooi0!bob 6!0!oob 6 !h 0 oij9 6 !b. b3 0u36!h336n
N, P, K g ! bb3! 3030360360b g!5hri336! bg3&! qjfrioi036ob 5!80!0g3- 8 3 6 9 3 5
6ib . 3.10 bb 3ig ib b 3i

qjmbqôbnon 9m9bhi{,36nb &i3C36:> 09hi(yi
Phosphate concentration in external medium 3ibrrb Shmg^Jjsni'b] (Rorison, 1969)
- 184 -
aooB^fnoan 8060 ^ ^ 3 ^ 603ai0 Drtaw(jto aowGmdaa ^ a a fo a a o w
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6iou aii6iq}ihgfn6i 0(539361 oihj,i6xnb bibihftpbgng. 00hi0 0ibi‘b3 519116-
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nb. 61b. 3 .11 ). ibi^nb 01,536160116 3hoiig, qjmoim00n n‘bhQ36i Ca g i S-nb 636-
I5330m6i, i&h30i33 091,436b 612103619 9mdhi3n 303036,5360b (Fe, Mn, B ) himgp-
6m6i, 9 i 3 n6 , (101(51 9oidhi3n 303936^3606 (N, P g i 36o6ih3big K) 2^6(536,5(11-
(5m ibioîbhgi 03010103680 93^01; 9ioio ibij>ob 9m9b^36nb $310 ^ 16 ^ 3 i 9 303-
936^011 (11019360161 oxo&b. i 9 n^m9 oii6iqjihgni6i: C a / K qjmoim03n 600b 516-
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oiih36nbicri3ob ih ihob biîhobo 9bm0mg 013691(5 1J3301 303936^001 933<bp3-
9130 3m3ihi536i. ihi6ij>036o 060936301*161 ijpb dn6jhsQjhn j c j d j 6^jdnb dsQ-
s 6bb, 163 9 ioi 9 mhob 6i *1i3D X n Shoiôihynnb ihb36m6ib.
9 (536ih336ob 0moibm36n03&i 9 n63hi03ho 213360b 303036,536061990 5i6bb3i33-
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îh6o hioig36o)6ooi gi ihi 601390 2)fT'03c 3jbo. 63636180 163010 hi0 bg3&i gi 3-
0133&X 6oi9ij,36'b3, oii6i 3ohob 936(5330 6oigi536cb3, hx3hi03h ^5003660 gi,
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9mhnb .y.33nhnb bjadivjohn {^ m b i^ x O ^ - Y j, T2 ps Y 3 SnS^hspjinb Sunh], bi93scm pa 9i(?i(yi
939|j33Cm6i; Mmin P6 ^*max cOh^c0 9ibnb 9n&n9ip3hn g i 9i;|bn9ip3hn Shmp^jijnb (Steenbjerg,
19 51; Steenbjerg & Jakobsen, 1963; Drosdoff; Prevol & Ollagnier ciL Smith (1962) g i Bates (19 7 1)
9nbx3ncn)
bnibpm33b (bbgbij gngn hbmg36m6ocnbb Zn, Pb, Cr, Ni, Co, Cu gb bb3-).
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6nb: ^bhdbg Qymgjn 3030360360b ©bmgmg 9tfnh3 mg^mbnon 930 130^6!; 93m-
3nb3&nb 930g 35 ^bh6n 3030360360b 5b63nohb036b 9 bcnn @b03;)3nb b6 yn 93 bho-
83603690 gs 3jh3gnb 633600 obn^btfoob 5^1001. b0bb 5bhgb, bhob bbb3m636n,
hm0036nu b0g(pb363636 bbm^h 0g5hbgm6bb 0 dn93 03oX?O&nb QnQn i>Pi6u36ohb-
Iin3&obbg0n (otTijboimcgoo3&n). b0o^m0 5b6nhfi33b 0do03 03OX?36ob 036(53300
6obgb53&nb obgo^bomho ^63016360: oy3nr>b 939(3330 6nbgb536'b3 - Festucaovina
gb Agrostis tenuis-, 150630b 939(5330 6obgb53&‘b3 - Viola calaminaria, Thlaspi
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6obgb536'b3 (Cr, Ni gb Mg-ob gogo hbmg36m6ob 930(53300 6obgb53dn) -
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b5h3cn33 0n3j)3oi3636ob6 Silene vulgaris, G ypsophila patrini, G ladiolus- ob
bb3bgbbb3b bbb3m6b, <bm503hcno 0 bh(530ri3b6 o gb bb3bo.
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- 191 -
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3nb pnifchift) (Frey-Wyssling, 1949)
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m9 gmanc^bnb (j3Qi. 3 hi3 ! 060 m3i 6 n 0(536ih 336 nb;>CTi3nb □db^h^^m-
h^cn ^hmijpbnb. 3 n63 hi03 h 6n3cm3h36iOn! 3(30(13 6i6nen 9 u 36 ihnb |jbm33o3 m-
j0393&nb 33hnmg5n33 5i3mnymq)b bpjh^& nb, oJbjjhQ^bnb g i t^^hj^^bnb bb-
bncn (6ib. 3 .12 ). gh3n-3nbcn6ftnb ipibnagnîunnb 3nb393noi, bj^hj^jdn ihnb
ibn0npiynnb jpSmymgipn ihmgpj^p&n (bbSign 6ib3nh6yc36n, 3 i{>., 63j(5ih3n;
jph6m630n 89^3360 g i i3n6m9gi336n qj3b3au> 5i3m6iymcg363n); jp ih j^ jb n ihnb
Spmhign U3pnb ihmg^j^bn, i6 jjiîbmonbQnb bibmpmm 3 hmg3jtJ36n; bmgm
hpjjhQ^Qbn ihnb 0:>hn03&n, hm3036n(3 (jnpcnnphQbion! (33Qi3n ih 093636 fiihonx^
g i 9(336ihngi6 5i3mny(rigni6 ihim hjyôn gmh3 ncn.
hp^hpitfnnb ghmb 3y36ihnb 6013^0 <b3gi3nhngi6 îSmnymagi 3 ihn03 &n. 63n-
3 nb 6ycn g i 6nbgn b^pb 36ym6b 3 icn jjiQmh^bpib. bm^nphonn nm6n K“ ,
N a+, M g2+ g i Mn2+) a p g ih p & n o n i g 3 n g i g . S p i S p i g j , n p 3 6 3 n g i b i b -
03630 3(536ih336b b3 nhig ifooi SihnofciOmSymqjn jnhjtfQpbn; jpi^bnib (3 i-
bncghiî) bib3m636n C a ^ O j^ -b fciSmymqyB 3ngi^r'ig36nb bi33i036noi.
bpjh^nnbi gi ^jb^hp^nnb pynomfcnphn 86n83630m6nb (iC 3 om3bcnni, i6^n6n-
tnbn, bib3m6ioi3mhnbn g i 3ngibib3m&hn3n iF'&iD^Buni) 03bib36 nb. 3ihi5-
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3 ib 6 ib 3 nh6 ignb, 3i65&ignb g i Byigbignb 8 3 6 9 3 5 hifng36ri6hn3ig 83010163 ig5-
ngn 3>i3ni. 3(336ihnb yomh^bSn 3 nbn biSpiom 338(5330016! 2 -4 %-ni, qjmanrnQ-
8(33360 bppbnb gmcnopbSn - 1 .5-3 %, bj^hm qjnoph q jm o i036b! g i 6n 63363n -
1-2 % , qj3b3363 n - 0 .5-1 %, So iSj^ n ^ h Byi03 (536ih 3363 n - 5-0 % , bm0m bm îgig
3(536ihnb (3n0r\3i6 6 0 30 ^ 3(136:^ - 15-19%.
3(S35ih30n 6 ib 3 nh&ignb (330! 3 ^nghmgi! g!»>!39 nh3&30 n iboi^nb (330!boii6.
363h5nnbi g i 8iTi03i303hn b^&b^B^np&nb B ih Sn ^ ib , hi(j ibm^nb Rihoi3ibcni6ii
- 192 -
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g!j)! 33 oh3bxn, 3'bh36330ymcgb 6ib8nh6:>gnb 1530!; Qgmhg 8bhn3, 3(336 ihnb bn6-
cn0 ii3 h n ( jb m 33 06m J 9 3 g 3 6 i ^ b ib iS n b ig , m h 5 i6 3 Q n 6 n 3 0 in 3 h 3 6 3 6 n b 9 ^ 0 1 3 3 6 ^ )
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3 i^n ihuoi3 n03nioiig cnSn^nhgd^gni i'bmonb hing 35 m6 noi. ibm^nb 5 i 3 mbi^-
390 3i3ihnbm6nbib 6(336^3360 bnSxpggSn 33b ntbhg36ni6I 3 iom jbm3nQ36nb 33-
h3g3&n 8(50(13 'bmSnbii, bmpm ^hggnb i3gC3?>n 36360 (ibm^nb 3i3ihnbm6not
j>i3m6333pn b^jhm'bn, i6 3 33n6m3mhqjm'bn). 839 36 ^0 ' 1133(^03^50300 i;)on3m&!
nh(j>333i, 3noiihg3&i igh 3x n gi& 3 h36i.
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6ig-i3omlj)hmqjxni, ihi 9 x i'bmo-iS^m^hmqjxniS- ibn^n 6nigi5ngi6 8oiin6ov
j333i Bn^jhi^nb i6 iSmBn^Snb nm6nb bibnoi. ibm^nb 386o83630m3i63bo Byihmi
nm6n NO3", hm33pn(3 ho’bmbqfthmSn 83^0 himg36m6noi 3mo3m336i, 3ngh3 NH4+ .
hm^mhis y33g! bb3i nm63bnbioi3nbI (33b380 8 3 b x n ibmonb îgibiîbigi^ bi^n-
hmi 363(150! (83bi6i3nbig, 3b ihm^bn^ gi3m2jng36xni b36oiJ3i‘b3). iSn^mSii,
hm6 (303 6nigi536'b3 g i I539n !3hi(5oob 3nhm&368n Sm'bihgn 3(5361(13360 5160(5-
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6ih33n b3g3&i ihi 3 bmpmg j,ih3gi6, ihi83g, !5h3ai33 ibn8n0nhg36i. 8oii6-
cnj3 x n ibmon i3n6m353qj36nb bibnoi 3hoi336i 6 ib 3 ohbignb 6i3hoi368n g i 83g3-
5ig, 6 ih3 mnJ3636i i 3 o6 pi03! 336o gi i 3 ng36 n (6ib. 3 .13 ).
I'bmôb ibnSogi^nob 3nh33g 3vj)i3'b3 Bn^hion 10 0 5 36 ! 6nohnoi8g3- i 9 3hmu3-
bnb j)i l!)i0n'bvi)fTih36ni qj3h836o36nbi gi irnqjijont^nb 801300 31^30, hro830Oii
Smhnb Scruîhni 6nohioh3g3Joi,b i (NR). 3b ihnb ynom’bmgSn ihb3&xn 3 6 ^ 8 3 -
6nb im33p3jbn (ynajij^mhgbn FAD, ^nomjhmSn b g i Mo-g^3hngn6n). bi^nhm-
360b 830aib333i8n N R bBhiqjig bn6oi3<bnhg36i, bmgm o6!j,5n3!(5n!b 5i6n(5gnb
bibmgmm 3hmg3jonb (NH3) o63n6nh36noi. 3o6ingi6 NR-b ibibnioi3bb d19016
bBhiqjn 6h363i, i 3 nom3 ibmôb (530180 050 ib h x 36 b 03i!>!0 86n83630Pi3i6
hmgb. Bnohivjhxîi^nb ijonsmbi bb3igibb3i 3ibnoio8 n 6 :^ 3 3 6 3 ^ 1 (5bh.3.7-8n.
6inh6i0ib368n 3 nbn 93301^3 gogn ij^n3fn6i 83(1608636! 03^0193680; b3 8(5361-
h3368o - q)3b3368n, i6 y0mho36bi gi gmoi0368o, 0138(3! 3b 8in6(3 giSm^ng^-
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0m3booi 83ig(nnb 838g3&.
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3!, hm93cbi(3 fcsgii^b cru>3nbn NH2-363q}n 6Cni ,T,0n'bnb 3hm(53b3o gi 2>h36bob
IjOicSn 6 ih9 mJ96 np bb3! a- 336m9 gi33&b3 ((jhi6bi9n6nh36i). h9 3nh33pigo i8o-
6m93*336n9>6 bo6oi3bohp36i bb3i :>9n6m9g:>336n, hm030ai! 6}b3 ohi>igob fim66bo
qjmh9ohp36i 6 ib 0 oh6 igob 13300b 93^C3g3hn 3hmg3j 63&ng;«6 , 9mji 0 mhnb, 3 ^ 3 0
60b 150300b ^D^DPDho 3hmp3j(i360956 pi ^^mbmainibaji^ho i5O30Opi>6. >3 n-
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6obiO»3ob.
Ijopob bo6oi]bn. 3b 3hmij3bn d(y»3h g i9ming36x n i ^ ^ ^ i ^ D h i ^ . i 9ib o ii6 ,
9 y 36ih336b 5 ^ 6 0 5 0 1 639^3hi63h3cn i g i ^ b ^ n b 35ihn. (jopnb bn6oi3b 0n 9n 6 -
i6nQ3 i6 9nbn 6o69b6h36o (ibm^nb i^jo^hn 9oii5oij9i g i 9nbn ^hi^bSmh^n,
i9o6m 9gi336ob bo6oi3‘bo, 6hi6b|)hn3i3ni g i ^^^bpm jjiôi) y330i 03lji6fTiC3hn
3hmi53bo aii3nbcni3ig 05601300363005 63033^ 3 ^ ^ 3 g i bibnicig3di b b 3ig ib b 3i
63033hi63h3cn 3013030150366001. i0 i 3 3 ghmb, 9ionn bofijih33&o 53150036019 3 6 9 !
3cni6b036mg3b 3hoi0i63oib. b6pih3g 5006^10, i0 3hnn53b36ob 3hoin6popi6ob 633-
^3hi63h x n fn>6f»030n> h r^ ^ h ^ 63bn, b i^ im g 3n6hrii. 013015!, obny i(pbi6 o3 -
6 i30 i, hm0 ynpnb bo6oi3“bo bibnicng^bi diQOi6 bShicgn g i 3pib6033hn 9 3 5 3 3 6 1
6ib. 3.13 J(vnh36jn0nb :J3h30D69n :>‘bmljnb ibnancnyinb 546^ ^ 0336x 0 bjo9i. i6:>emyr)hn
9m6iU3ao6n <3 ^ 31* ! e» cybrnqjricnb jjhoQj&nbcr^nb nb. Stulen (1986), bro<v»i Sihynbi&i
6ii36ih336nbcn3nb - Streit & Feller (1982)
- 194 -
68r)60 b aoad& nôsao
ISbhnpo 3.7 bb3*e*bfcr3* 3*6n^j3o 6o^h*^h3Q3ij*'bob b*33*cm siy>3m6:> (|idm(vi NC^'&h"1)

(Gebauer et al., 1988; Stadler & Gebauer, 1992; Downs et al., 1993)
3^36 ihQ gs 5s6n£HSn gjmomc0 3nSnbj339s hrphmeatonxn

mh5*6m3bn mhfc*8nr)6n
h^gQhicjhn 9(j36ih33&o 14 1 4.5
i'bmlyKn 3pnp*hn 3*60(^360 7 - 13 0.6 - 1 1.6 - 4
6*e*bm3636n j*cnb ( jy jjib i
g* 6y>Qb^3336nb $*h330m 9 0.6 1.6
b*dm3*ho apxm jbn 3 - 4 0.3 - 0.5 0.5 - 0.7
5*0^61*136360 9cnnb bidmshjbV) 1 0.3 0.9
6^^(5^36360 pihf>6 60^ ^ 536^3
(l}*h|5n*6o 60*9*5360) 0.6 0.1 0.2
6;^\£>m36360 ipsho6 60*9*536^3
(bnpnis1j3hn 60*0*5360) 0.2 0.1 0.3
Ericaceae-nb 335* 63^^61(1360 0.06 - 0.1 0.04 - 0.07 0.0
<30^36*11360 0.4 1.6
Acer mbrum 1.5 0.7
Fraxinus excelsior (10-15 600b) 1.4 0.1
gm bncn (6sb. 3 . 14 ), hi(j 3 3 ^ 9 96n33630rn3i6n 5 n6 i 3nhm6 i i 0(536sh3cn> 9mQ3j)3-

C^hn, g36Ji5n3hn gs 0mhcgmcmj)n3hn b g i S ^ r ^ n b s . b^hpbSn 3 ymqjn 9y36ihnb
y n p n b bn6cn3“bn n63n&nh3&3oni. unpfib Qp^mcn'banb gbhp333nb j>i>8rn30n6:)6:>:»
i 0 n6 n 9gi 33i)nb ShmSmhunp&nb 9^(7333!, oi3!Q3 nbi(336nb 3 hmprv
6 n b jjrn6(33Gvi)hi>^5nnb d Q f^ h n 3 m 3 b li|3 6 i.
Acclimation
temperature
o 4°C
A 10°C
□ 20°C
• 303C
■ 36°C
6*b. 3.14 (jncnb bn6oi3<bnb Qbdmyip3&3C36;>

(Weidner & Ziemens, 1975) Ambient temperature [°C1
- 195 -
806066^7)6 6030)0060660)6 300)30U0&6
3 .5.3 ybmnb 6ânyn<3^
DQuPOObn 815361)13 ihimfi5i630 ibmiiS3615300 6030103(13636b 3oii6oiJi3b

8mgi6, 8151118 nb 83,519 063010119 91 8^(13 (11019360^01 5ihgiJ86nb 6101 ih^>-
m h 5 i6 x qjm(161893 (NH3). 6(536^0916 I’bmô dnhnongig, fcidmnynqji 01(1516-
jQn qgoih8noi: <3363360 5i0myoiqj36 1606018313366, j,C30i 9n6nb 91 ib 3ih ^ ^ n b
831336b, 503^18066, ib 3ihij,n 6b (boifcr^mo mjibnb 6ih8m0agj)36e36o - L5niir
h3cn6b g i iciB^mnjjnb 031336b) 91 bb3i oih 5i6x ^01001316 6i3hoi36b; I'bm-
,506 gngn hiotg36m6i 051I15361 qjoicnpo&nb 91(5336061 91 6iymg36n6 61801(533606
636935; ojoioioiqo 6301513b ibm^b 3hoi(joi3oi‘b8i 8n g i 15)130133, 35)13906 63368 0
6nli)hn(j36nb bsbnoi, 6oiqoi 01360360 15(10133636, 16333, baQshayn (500366. 6 1b .
3.15 6163363601 ibm^nb 519161(53036061 g i 30OQobi(5no6 6^381, boiQoi 616. 3.16
0^15383001 iboi^nb ^hiBbimh^nbi g i 1608001(5006 5181)1,50336300 801930360.
01(1516x11 gmh8ogi6 1(1101(15163080 ibm^nb 5igiy3i6ib 1601)1(50303636 I'bm^-
33iS3hfril!|hmq}3on oih5i6nb836o (1561*13x360, 601501360 91 61^3)10360). 8boiQmQ
nbn6n i6h303636 oQupCD^n 8(5361)106 iboiôb (5050b, (1016300(5, 18 531)119 ,
9190150936x01 8n5hmrnh5i6ob836o6 ibnoob 15X ^ 0-
fob . 3.15 s tc v ja ja ttto c i y T>93iT'6D61j36f*b

^hi6beniji(5ni pi jyie n ’bitfni 31(1153001363690.
pvii qj^hnb oblho: I<bni(j3 3 3 lj3 x n
6nb ,jhs6U3oih,yih3&i JbnQ3dnb biSjscsdooi
(61110150916 qj3b33&ob 513c™1 3uo6ihnb W>3i
6i 6no3&3 n; Qjjn cjjhnb nbshn: ^hi6b3fnhj,nh 3 6 ^
P3hn3&ngi6 qjooi33nb 5130001 953^33630
(Nikolas et al., 1985)
196 -
;> 8 n m 83666 n ^ 08 an
Xanthium Pisum M a lu s
t ft
NO j
<N > N itrote accum ulation Protein synth e sis L o n g -d is ta n c e tra n s p o rt of

N itrate re d u c tio n Protein breakdow n - - - Inorganic N com pounds
0 N j- fix a tio n by sym bionts ^ P hotosynthesis — O rganic N com p ou nd s
[a ] A m m o acid - a m id e pool N -free carbon co m p o u n d s
6ib. 3.16 i'bmjinb p i ibnQnensni bb3igibb3i duîhjSn (Thomas, 1927; Gaumann,

1935; Wallace & Pate, 1967; Pate, 1976; Beevers, 1976)
3.5.4 a n ih n n h 6 ^ 6 f r b a 3 6 n b a n o h q jf ij w is n ^
bb3igibb3i 8m3hmrrih6i6nb336b 8330)011101 1^8(^6033^811 1^3630*1 ibmôb

qjnjbnh^bi; nbnSn 5(^63336 N-^^riLjhriqjnnb 3j)fn0Pi6O3hi9 396(193630^131636 grn-
63b. 363601 3hm2)ihnm636n - 61^3111136(1, uni5m6ij63hn36n 91 ij6o6rTi8ni5363&o,
hm9036nu 6i6o0m6ho3 011306393019 91 6i6nQrn6hn3 bn96 nfT>b3 hig 1^36(11636
6 nigi 69 n.
0113116303019 9ubm3h36 I^8m6q33h30n ibmôb 9iqjnjbnh363Q 6ij^3hn36b
9 mhnb 3nh330ip 91953600 nJ636 Clostridium pasteurianum, Azzotobacter
chroococcum 91 A. agilis. ihb36m6b 633hn bb3i 61^3(101, hm930 n(3 iqjnjbnh36b
9 m03i 303h ibm^b, 6^6611)301(136(101 q3n^m^hmq330n 6 ij(i(3hn36o g i <bm6n3hoin
6 y i 09 n 51311(5303630(1 N2-9imjb(ignh3630n 61^3(101. 9hi3i0n (5ni6m6ijlj3hni (9 i 6 -.
Synechococcus, Pleurocapsa, Oscillatoria, Nostoc, Anabaena, Calothrix 91
Mastigocladus) mh6i6m636xn 303936^361101 9m9ihi636i8n b h xn ig i3i!pMibXni
- nbn6n hm^mhu 6 ib 9 nh6 ig-, nb3 i'bm(i)i 3iimiihf,1<31DCO^r' ihm6. ibm^nb 9 >
030^11(1363011 (5ni6m6i;j(33(1(13611 63^3g36i Byi09n; 63D3Cn63&ni6 hm6mhy 3nm63h3-
6n (9 fi9330 b36b^hi6b3 gibib036nbib) 6013661 g i ihj^niô; 01311930 6yihm3&-
bi gi bb3i 3 ^ ^ 3 6 10 3 (1 195003680. 3363(irTi(5o6l336116 8161(136300 61^3(103611
0(1^36368(1 61(18^11953636 63bi63 3ih66 omhb (bRjjm3i6n yn83m6366nb 3 -6 %).
0nj3636nb oii03bnb bmynb 9h39n 9330ib3 b9 nhig 63^39361 Nostoc.
011306303019 8^1*13(1360 ibm^nb 81030^0(136300 6^63(10360 516^3301(13600)
3hmp3jlin x36oi 01600 g i 83g8o3ig 636016 3i6n6i6368n: 6110630b 30i66H5H369o
- 197 -
906 a r n ^ 3 (iS 6(13(J100rt 0 &.MJto 93 m3 nbOW
I5nb5 m6 bj^3hn36n bqjrtjbnh^ft^ 50-70 2)6 N2 3y iy i [77 ]. ’bmOr^hn bih^ypnb bib-

6*3 9n63&0n bib^j^nb 9 ;x£>6 nb 936935 0icn 3n3h 830013^36! 0bmpmg hi8g36-
n03 N , 3i ‘V , bmpm ,jhm3n2)36bi gi b3&(jhm3n2)369n 03bidp3&3pni 100 55-
0Q3 N 2 3i^ i"1 1 03cn3fib36i. i'bm^nb 3iogn;jbnh3&3pn 6^3(11136(1 )jbm3hm6;)6 i5-
(130133 33n<3nli,3hiQ ^hrOr^xn b3 3u36ih336nb ogmcnp^'b^, bigi(j 0ian 033dpniOi
03 rtcji3 nbm6 5 2)5 N2 S y 'i" 1 [3 1 ], 1339^9 giSpign b36b^hil5nb &jm63 6nigi50n
(0i5-, 63gp3303bni6n 6nigi5n) gi (303 h35nm63&3n ibm^nb cgnjbijjni bijjOimg
gibipni; b36ihj(!)n2)i9n g i ihj^rîSn 1150, ig5npb:>0ymqj3pnb 0nb393ncn, i ^ ^ b
0 .1-2 2* N 2 3i ‘1i-1 [33 . 8 0 ],
bn96 nm‘b3hn N2-3iO)n;Jbn(i363pn gi 6ib9 nh6 ignb 0nb3g3ncn ^Qhri^hmcgxn
6ijii3hn3&n 6>b3nh5yip6ig36noi 3‘bh3633pymq)nb 3hm6p33ib 61^33^36 n0 ncn, hm3
bibp936ni6 ^^m^hmcgx0 3l536ih336nb 33h3g369n i6 n2)33636ni6 hn'bmbqjhm-
g i6 5i3mymcgnpn 3jbxîJ3£>ncn. i 0 n(jm0 0 ioi 033dpnicn iÔmbqj^hmb itim^nb
&33hig 3qjhm 3q33jii3hig qjnjbnh36i, 3ngh3 cni3nb3qjpig 0^013(136 6i;j^;)hn;)6b
(bbh. 3 .8 ).
306n33530m3i63bn bn06nm‘b3hn i'bm^rtb 0itgnjbnh363pn 6ij^3hn35n 32)3ai36ni6
Rhizobium-nbi gi Bradyi'hizobium-nb 53ih3&b, hm03pcni (3;>Ci3X n bibQrn&Qdn gi
0hi3ipn qjn'bnrnprn^r^n hibi (5bni3hm6b 3ih2)f,ibi6cni qj3b336nb yngh363n. 3b mhn
53ihn 5i6bb3i3936i 3hcn0i63ombi5i6 0iam 0nph32)fip36noi bb3igibb3i 0ib3n6d3pn
0(j36ihnbig3n: Rhizobium-nb bib3m63&n 2)fT|Dh36b j36 ni6 dnhncnigig Trifolium,
Lotus, M elilotus, Vicia, Pisum g i Phaseolus gi b&h30133 bmnib 3X 3^ qjmh3i 9 n
{G lycine so ja ); Bradyrhizobium 2>fT'm33hnh3fi>b Glycine 0 ib, Vigna-b bib3m636oii6
g i bb3i b36l5hm3n2)X g* i2>hm3n2)X 3ih2)mb636oii6 . 3ih2)fTib63&nb 6ih0m0ig5363pn
S esb an ia rostrata, hpi03pni5 33hnmgx^g gic>6mhnp ig5npbh3ymq}Q3&9n 6n6ighm6b,
n3ncriih36b 2)^3^36b (P3hm3&nb fthjSn. N2-0iO}njbnh363pn bn36 nm^36ngi6 u6m&np-
r»i i5h30i33 3^36(1 53ih Frankia-Q$5 , hm3p3&ni5 J36 ni6 yngh36b Ainus.
M yrica, Hippophae, Elaeagnus, C asuarina, Ceanothus gi 'bmj/i^hcn bb3i b3
3l536ih3bcni6 gi 63fijcni6 [21 , 77 ], ysx ^ 3b b3 01536^3 013 636jn 3nrn63hni.
nbn6n (jbm3hm636 i'bm^ncn (jv>hn& 6nigi536<b3. nbn6n ^061*130153^36006 bod-
6nm(bnb djdjjmbnm ym33pge>03hig iq}ojbnh3&36 50-150 2)6 N2 S y V 1.
3(}36ih3bcni6 303(1™ b^b^ bn36 nmb3 n (ibmijniijr^hn bn96 nm‘bn) o3ymo336ni6
ibm^nb 3icgnjbnh363pn 6^3(10360, hm3p36nu bibpp3&0i6 0 ib 3n5dpnb q)3b336b
3 mhnb g i b33&ob 0n2)nhnbi9n. ibmijni^n^hn bn06 nm(bn 3i6ol>)hoqjoi5oh363p 6yJ-
(jbhn(yi 33 N2 bn9dnm‘b;)hn qjnjbi^ni (y , N2 ) 3ihimb5363n (Wemer, 1992)
anenaân / bs93JCm
Lens 50-150 80
Trifolium 45-670 250
Pisum 50-500 150
Medicago 90-340 250
Lupinus 140-200 150
Vicia 100-300 200
Glycine 60-300 100
Aiachis 50-150 100
Sesbania 600-800 700
N2 - 0K3nJbnh]i>xn 80-500 150
- 198 -
6 8 0 6 0 b aa&6 &ncio 890
0)3hfi]6bi ga £hr>dni)3C 9ah(j30fn36366 9 rohnb jVhrfijyyym qjii 5-30 i 5 N j Sa'^

qjnJbnhjftab, bmnm Anabaena azollae-6a ga Sypnb 53n0ha Azolla-b 9 mhnb - 60 -
120 35 N2 3a** a^ [2 3 6 ]. ac>0mba3Q3oi a'boa9 n, 6aga(j Azolla mgnoi5a633 963563
bab^Jag 5a0mny3636mga, a9 ab gngn 86093630^6! aj3b 6hn63ob 3oa6vjau036ob
abm^noi 3<6h3633pypiqjobaoi3n6.
bn96 nmIbo 3ah301663606 033633690

hn'bmbriQbn ahna6 0i&p05a63hn a3hoi636n ga (56013(101636 bi3h0KJj0(j3pag n9
h3j,nm636n6 6oaga5369n, 6aga(j 9aoio 9 a63 o6d3po 9(5365(13! 6a3hi53p363po. a6ap-
5abhga 0!b3o6d3po 9u36aho6 o6ojouoh3&a ga 6oigxa(5na (2)013(13606 9 ] j 8 &a)
6go&a 6aJo3hnnb 53606 9 n3h h353pnh36ago 6o56ap3ho ja ^ n b 63933016001 [2 0 7 ].
hftig3ba(j 9mbg36a 936a6a9n6n 9 ab3o6 dpob a0oi|56oi&a, (9a5„ QuoSahnbo^nb p3j-
0n6nb 03633016001) ga 9ii36ah3-bn96nm6onb 6aj(jl3hna'b3 lbh363ab 06-
y3&b y33Ca‘b3 abpm 0yoiqjo qj3b3nb 63b3bn. 6363606 i3gpnb ag5npm6hn3ag 5ab6-
6n6 03933016001 6ajo3hna6 5‘ba 3dp33a 0936306 Johjnb 3ah36jn9a9g3. aj 6aj#)hn!
5a9mymgb bn56ap3h 6n30in3h3636b, hoi9p3&ou 0633336 33(^ 36 0 6 03 hobv!i30x
gaymogab.
301911360b 5a63oo»ah3&a o6g3ijohg36a n6goip3bo5ob 9ga336ob, un,jmyi6o6ob!
3o63hxn636ob j)m6(j36liha(jnob 0m0a,i)36noi. 03ba6a0obn 53606 03833016001
0 ab3n6dpn6 33h3g3&9o o6y3&a jjoighnbaoôb 63315003031™ 6o30io3h3636n6 (ynpo-
6n, 5p3oa0n6-6o6oi3oa‘ba, 3hoâ6a, PEP-3ah6oi;jbop!'ba ga p35-330m5pm6n6o) 606-
013*60. a9a33 ghoi6, 6aj(j3ho36o, hm9p36oi5 n6q)ni5nho6n6 a0 6oagna‘b3 9 oipoa6 ag
9 a63 n6dpnb 6ah3<b3 0^33636na6, o6y3636 5a0ha3P36a6. 0015305636001, 6a6330363-
pn goihfrib 8^0163 ijapj)33po 6aj(j3hn3&o 5ahganj0636na6 3qghm gogo lbm0ob 6aj-
vj3hn36ag, hoi0p36o(5 bbg3&oa6 9 ab3n6dpn6 33h3gn6 33hn6a;)oohoinpx 0306ha-
6ab. 3.17 Viciafaba-b (g jjjrl) 516030 ^tinpo. B 6i i s 3hmnen; SS bna&nm'bjhn 1(13 q i bnQ&nmVjhn
9j96hi6i; PM Jjhn&iiyjhrrxyi 8386(1161 (303tohm6xn Snihm^hiojoi D. Weiner; 20 OOOX)
- 199 -
aOBOff^Tlff. 6030)00 ^ 3660)6 930)30b066
6580. 05b3 o6d0 ob 33h3pob I5o6o)305b0oo* 55h838 oi(i6 y8 x n <336^50016503(10 3(101-

3 DC3^n ("bn06 nmbm036 n") ^ 3 3 3 550oioyoiqjo56 3(10105630^5556, (101501(115 33(13-
pnb bb35 65600360 (65b. 3 . 17 ) (201 ). 30mdh5mp 95(163609 6i;)63hmnp360n bg3&i>
ibm^nb qjojbnh3&ob 536360b p3(i33h3bo(i365, h5i5 06333b 6o6hoi5365bob bnb^^-
0 nb 50mj03p365b p5 50ooi N2-ob 93cn3ob3&ob p56y365b. 0 ib 3o6d30o 013365(106
p5y353o036 ob 838935 31*13(1360 63(1936056 p5 035-3300150016060 000365. 50 phmb
6ij6Dhmnp36n 55(19511;)96 365 o6qjoi3oh3&nb 365ho56 65^3^03659.
5l50fT'bqij3hxn ibm^nb 03cn3ob365 0503^636x^5 N2-ob 0 fT>03i X ri*J h x 3 jl5nDC
5^03635^13. 3b d0O3h 36p3h5m630O h35^05 35650obohp365 6o(jhm5365Ibnb bnb-
6300 b 0 O3h. 6 o6 hm5365,bob bob6305 5hob 01(13(10163060560 3oi8303jbo, hm03 o 9 n(j
03pob 3hoio Fe-3hm63 o6 o p5 311010 Mo-Fe-3hm,33o6n. 3&3h50obi 95 (1393^500^01-
30b b5ôhm 303j6hm636b od0335 b36oij3ob 3hm(j3bo (65b. 3 .18). 85b3n6 d 3 C n
815365(13 3<bh36330ymq3b &5j63 ho36b 65b8 oh65gob 03653hoi36ooi; 50oi6 o30ob ib n -
0 o05(joob5oi3ob b5ôhm 36^0036001; 035-3300150 m6 o6 ob5 p5 3Pih3nb 03360b 6 3 6 -
33016001 01353b 6y)63ho36b 35656590b popo 301615366(15(5036065556; j 06 ob 6 5 ^ 3 -
ho3&ob50i3ob b30b5y h x 33h056366X b6 636056 301101636b.
3(10169056 35(130^3680 ( 65hp5, 656^30105, boios) 33536^ n3ho 55630015(1361-^
qj5b 50 o 05 y353opm6ob pi5y36ob 606 03010103600 ^6013^0(136300 65b9oh6ybQ-
659360b l/3-p56 2/3-093 6^6^01(160(19365 03 b 33&b5 95 3013(13600; 0501556 9 .5 .5 b -
00136001 -1/3 0 6 5(1336 5 ^>3601^35^3, I / 5-093 - 3<T»ghnb bhgib3, bm0rn 9565(1^360
45 -50 %, qjoJjoh 36x 5boi60i56 3(10159, 3356 36(1369365 8tf365hob 03010193&bi p i
bb 35 6560936b [62, 268 ]. y3X ^ 3 P*1? 0 bo6oi3b3ho 5j603m6ob 33(1001980, 50 n6 m-
03533&ob5 95 5009360b bibooi ihb 36x 1 5(1 5^016^3 065(13365 65b0nh6y5c6ê3-
6nb b5booi 5hb3630o 4 5h 65b0oh65go [ 176].
bo06ooi6636ob ©03(1 ibm^nb qjojbstfoob 06636^30165, 33nh3X3b5p, giOmô-
9363005 65b0oh&59ob 5bo0o05636ob 0o6m9365(b3. 05b3o6d30 o 013365110b qjm^m-
bo6oi3“bob 956590 066366030165 06333b 5<bm6ob qjojbôob 036x365b, bmooi
q3m6fnbo6oi3bob o6636bo30)6ob 55bhpobib (055., C 0 2-nb 0016019360b 8n0563&5),
65b. 3.18 bnSftnrnt^hn 0n]h I'bm^nb gnjbijjni ps 9n9m(j3pob 3hrrn53b3&nb 5*0^^)03363-

pn 3 me3cn (Evans & Barber cit. Marschner (1986) pi Wemer (1992b) 9nb3931101)
- 200 -
aOGOfoi^Tltf 6030)0063660)6 03^01 0)630b36'3606660 bb36^6bb36 366066630
Bib. 3.19 bb3iesW>3> 9^3-

6ihnb oj3b3nb b6ioip3&9n
5nljhm636i'bnb ijiingmdnb
giflmjingjbxD^1
hi'b] i'bmîojnJbnhjbxn
bn96nm^36nbi 91 iyu>5mdsj-
(iOhn^bnb cns6bp3dncn
(Granhall & Lid-Torsvik,
1975; Waughman, 1977)
mhj)i630ig 69 x n ibm^nb J,i9mbi3ipn 93ndp3&i biSjoh gjirfoihgmb. 5^)1363 ojij-

0mh36nQi6 y 330i ‘bo en9n 06n0363pm6i ij3b Bnigi^nb (J36ni6m6ibi g i
iÔhib. 6nohni536i<bnb ijijn3m6nb ii:)a33hili:)h;)pn igiS^nh^bigni. n&n
39016333^ 9 i335bhnb &i3hi53po6nb ig&np’b ] 5^ ^ 1116 36 30 d)3033hili)3hop h^gnSb
(6ib. 3 .19). 6n1i)hm536b'bnb bnb^Snb bn(5n3nbig3n 05hd6m6nihm6nb fciSm, N2-
nb qjnjbnlî n63n6nhg36i 3333 gi6ipn gig;)6ncnn t|)3933hi(Ejh36nb 3nhmb369n.
3.6 0fl63h3C3h 6n3CnnDh36XID I330nb Cn^3flb363h36^6n

bb3}G}bb32) m iiW iban
<33b336nb br^hunb (hn'bmbgohpib) jndn^hn 5 i 6 i 3nhm636 b 0n63hip3h

6(13011131136^01^ n9 0ihij,b, hnb biOfcôp'bQU ^6gi 0mi6ymb 0(536ih]9 oii3nbn
03Oi6mQn'b9n. 6nigi5nb 0n63hip3&nb 3 o0 igdo 6 ppi&ib, 6nigi&nb hayJunib g i
nm636nb 9 bhij,b doo&iQig 5i6i3nhm636b dnhnoiign ji6nb ftompm^nohn ,yi3n.
6nigi5nb 3n6 hm-bn3h(5nb3X I53i036igm6ib 0U36ih3 36D3&i bb3igibb3i fcbnoi:
Bnigi^Sn b3pbiyh3pn 3mhnbni6 li 36nb 0md366noi g i ihib3pbiyh3pnb 533hgnb
i3pncn; g3b336nb ‘bhgnb j30mohfnq3X^>e 3m'bnljnohn i6 Boî^n^hn hoijunncn;
03jji6mp3hn Spib^nôhmbncn; j 30mo.bnlyi 3o6 nb, hib36nb gi gn'boBJun^hn &i3h-
(33P3&nb 3Jm63 snîh^pn bibQm&obnb 5i63noiih36ncn. ij3gi6 5i0m0gn6ih3, 0n6-
3hip3hn 61130 1^ 36 36 1^ (53pnb Sibnoi^'bo gi0minp36x36nb q335j^nm6ip3hn i6 >
prVbn 6 ih 0 m ig 5 3 6 b 9 3 g i h 3&notn ojjm qjn'bnfnpm fcnnb 5 i0 m b ij) 3 p o 3 ohcrKjhcn
96 n3363pm3i 6 cnÔib.
Bnigi&bo, hm03pn(5 anB^hipohn 6ii3ain3h;)6;)6nb himg36m6<ibi gi 0361533-
pm&nb 3nb3g3ftOT Jjihfcig 5 i 9 mbi03pn oii3nb36;)h3636noi 5 i 9 mnhR33i, 53b3g3&i
9l536ih3oii bibQm636nb gi9 ibibniOi363pn bSQj^n; ij j,3b3p36ni6 "b33unipnbo3-
- 201 -
806066^7)6 6030)0063660)6 330)301356
6n", hm9936b*(j *b*bo*oi36oi oi*3ob363ho 93^*6m9ob9o: 9g*336ob*g9o ^ 9 3 ^ 1 * 6 -

(i»3 Cn bib3m6]6n; 3*ho93&ob gopo yn6(336li>hi(5036ob*p9n 9g5h*go bib^mb^&n,
9*5., 1*9(11030^360; 9dn3 Q 93,^*036001 9gng*h 60*9*590 9m6o6*gh36o - 93Qp0mqj-
0^360; !5h3Qi33 b*631*16360, (11*199360(3 0^*636 9*ho93&ob 9*6*9 im6(336^h>ijn36b.
9(1*3*90 3*110*6,509*6 5*6306091*101 9bmpmg bfnj,n3honn 8*5*90010.
3.6.1 392)30 Q} bybW lh Mbbibh^'bD aPTb^hQfl 0l3D6ihD36n
9(336*hnb 5*h39m9o 6y*9&*gob om636ob j)ri6(336(ih*{3n* 6*h9m*p536b j)336nb*

p i &!3hi53936nb 96o9363gm3*6 9*h35 390(13639 qjj^mhb. 6o*Q*5ob (13*^0* p * 3 -
y*h3&390* 6n*p*5ob bb6 *h3 n 6y*96*gob om636ob i'T>6(336(t|h*(30*b3 (ij^3x?Dhn
9g*30*6m6*) 9! 9o9m9(3393936ob 3n3h H+ nm6Q6nb 9oi*6oij9*b3 (3m^36i3njhn
9g*3n*6m6*). ,5360*6 (1350(1163690 6o*g*536ob 393,j3bm6*b *j3b b3b^n 9g*33 (p H
= 5 - 6 .5 ) 56 630,5(1*93)10 (pH = 6 .5 -7 ) h3*j(30*, 9*^*0163639 6o*g*5b *b*bn*oi32>b
donah 9g*33 hgijuo* (pH < 4 ). 9g*336ob 939^963911 9o63h*93dob (955-, 3nhn^,n)
5*3936001 60*9*56 93nd93&! 3jm5g3b gmi*93h*g 9*6*911 pH. *(109390 ( 1 3 5 1 ^ 6 3 -
6nb 60*9*5366, ,53,53 9*hn93&nb gs 5*h6m6*,536nb 9*5(11*133606 b*h3<b3, *j3an
dnhooi*go h3*ji3n*; 630,5(1*93(10 9*ho9n*6o 60*9*5360 5*9r*i0hfi330*6 bm0n3hn
(13*^(30001 (pH = 8- 9 ), bmgo*6 60*9*5366 9903(10 ,530)3 (13*^130* *b*bn*cn36oi (p H
= 10 g* 93^n).
60*9*506 9*83*30*636* 9h*3*g6*nh*g bg36*: 60*9*506 5*81*1030,53001; 9n0m(33-
p*gn 2>*cnnm636ob 5*9m,5*6ooi; 9(336sli336ob q33b33dob* 9* 9o2>hr*)mh5*6o‘b03&nb
3n3h 5*3mymq3090 mh5*63on 9g*336ob 93933m6non; 3393bob 033609*6 330n6nb-
9* <3393m- 9g*33bnb 5*991*1*950936001 g* b36oij3nb* 9* 93^0906 83935*9 6^b -
9 rth9 g*336 nb gobr*n5o*uonb 93933 m6 noi. *9 *b 39*^36* 9 g*33&nb 939 m,5i 6 s 6*93^3-
6noi g* 9g*3nb 6*h9m9j963go 5*9*^3^0*636390 6*3(1013606 (5*6b*2)3oih36ncn
SO2) 9*93^3*. 6n*g*5nb h3*j(30* 0(33936* 690b 5*69*39^6*90 (5*66*5301)13&ncn
9*91*12)093639(1* 6*93j36ob 6903b 5*6*60936*^3); *j, 5*66*5301(136001, 6o*9*£>nb
bb3*g*bb3* 3mho'bm6^9 o, 6*h9moj9636* 9(5oh3bo3h(5m3*6o 5*6bb3*336*6o. * 9 n^-
m3 , 3*6 o^*^nb g*b*bo*oi36ob ghmb 6o*g*5ob h>3*;J(50* 369* 5*obm8mb 690b 5 * 6 -
9*39m6*9o g* 9ob y339* 3mhobm6 ^9 o.
93hj6o*6n 9(536*h336ob 333,536(116*6, 6*36366* 9* 9oj3636b 933990*01 *hb3-
6m6* 6o*g*5ob h3*j(300b b*^9*mg cg*hcnm go*3*bm69o (63b ,50 9g*33g*6 -
iK3ib0 h3*j(30*9g3), obo6n *8030(51*193(1*6,539360 *ho*6 pH-ob 03*11593690; 3.5-8 . 5 .
13* 9^339 b*b3m636b, 9*oio 5*66*21301(136390 qj0bnm9m500b 5*9m, 5**R6o*oi *hb3-
6m6ob b*2)9*mg 3o6hm go*3*bm6o. Avenella flezuosa-b (9g*33 6o*g*5ob*oi3ob Qi>-
9*6*60*0136390 9(336*h3) 5*6300i*h36nb m3ô939o d33b pH 4 9* pH 5 9mhnb.
*(30g0q3093h360* *5h3cn33 bm503hoio Ericaceae (9*5., Calluna, rododendrortis
633I10 b*b3m6*) 9* b*b3m636o, hm9935o(3 obhg3&o*6 6*213360 6n3Oi03h36^dncn
p>*(in6 *g5n9359o (^*m6ob 0(536*113360, 63360b 9g*33 J3hJb3 9m6o6*gh3 911^3-
6360).
3o2(hmmh5*6ob9oii 9 mhob *hb3&m6b 633ho 6*1^53(10*, hm9390(3 9moaibr*i3b
6n*9*5nb 630^h*93h-b3b(j( ^3^3 h3*j|jo*b, 9 * 9 o6 , hmys bm503hai0 bm^m - 630 ^-
h*93h-b3b^ 9g*33 5*h39mb. *hb3&m636 b*b3m&36o (9*5-, Aspergillus-nb,
Penicillium-nb g* Fusarium-nb b*b3m636o), hm9936o(3 353360*6 pH-ob cg*hcnm
- 202 -
8 0 6 0 6 6 ^ 6 6030)0060660)6 (33^01 0)630b 0b'3fn36660 Ub36t? 6bb36 365066690
Chemical weathering
u n t [ Humification
Biotic activity
H+ and OH" toxicity
Al-Fe displacement
N and S mobilization
Pand B
6ib. 3.20 6mpi5nb pH-ob 513036* xrrrx
6mg35,nb gmhSntî^, 806311103(10 Ca and Mg
6n3om3h3636nl> 838(533crndibi pi n ix x x z c x
K
9u36ih30u>cn3nb 8o63pm8ip(’i6i'b3 pi,
brîpip, 60191580 Ixujmyboob 3nhm&3- Cu and Zn
6^ 3. gni5hi936ob gihcnm 616000 ^ r = n = s = j= r x i Fe and Mn
803010013^ 90631110360b 63^
Mo
93d(j33C(,i6ib3 pi biboiyn^bpm > 1^ 363- T " 1 1 r
6nb n6lj36bn3m6i'b3 (Schroeder, 1969) pH 3 4 5 6 7 8 9
gni3i'bm6b (pH = 2 - 10); 18 133 ghmb, ihni6 dpn^h 63133 16 dpn^h ii3^D 6 ^ 3 -
9 mbip3 n igii^nl^& xn "3n6hm b ^niQ nb^bn'1. 93133 01311630 6 yihm360 n, pH =
1-3 3nhm&3&9n, 6n6ighm636 ^m^nhgbi^hn^nb 53111360: T riob acillu s gi
Sulfolobus g i Acontium-nb, Cephalosporium-nb g i Trichosporon-nb j>3ihob bm ^r^
60. 0i6nijhnqjnunh363cn g i ‘bmjjn^hcnn 18016(10^1501136300 61^3110360 ihni6 6i'b-
nljm03hi6vj)30n (pH = 1 1-693). 6 yiQ0(j361(133661(5 3330)00101 ihb3&m6i hm^mhu
d0n3h 83133 (Cyanidium caldarium , pH = 2-3 m3^n838non gi D un aliella acid iph ila,
pH = 1-2 (^3^0636non), nb3 d(Vi3h ^3^3 {D u n aliella salirta, pH = 11 m3,yi933noi 91
Plectonem a nostocorum , pH = 13-693 01^0838001) 5 y03680 [ 193 ].
5ih36m9 n ihb3630o 69106190b om636ob 2)016(536^1111501 363*0019 801^639360
8(536ihob 35h393&nb 6n3Oi03h36boii (530^3. 6omjo8o3ho 3hm(53b36ob gngn 616-
61b. 3.21 pH-nb 5130361 Ap+ bbBipmbi'b]

oinb6ih 6mpi5an pi Ca2+ 8061^13301^3
5iC'ihnd36x 6nipi53n (Bannister, 1976;
Lathwell & Peech cit. Mengel & Klrkby
(1982) 0ob3P3oai) Soil reaction [pH]
- 203 -
806066^7)6 6030)0060560)6 800)301)056
npn mSonSiCDhiQ 3n0gn6ih3n>6b pH-nb oghnig 3n6hm gni3ibm63n (pH = 6 - 7 ) .

b(i>>6n(vjhn jjh^gSngi 5ih30mb 9 o6shR;)6o&nb:>oi3nb bi^nhmi 3hm,yT»636nb B-j q -
9n3n IJ3C* 6 ^ 3 P^> 3nj>i bn3h(j33&b 9mhnb, hi(5 363(15™^ bihj36oii6 ;>hnL>
9>2(>39nh36xn- ‘bm^nQhm 033o»b333i9n pH-nb 5 ) 1^ 3 6 ^ 4 biqj3b3hi0g3 ip 6 3 3 b .
3.0. 0 nbn caifiigihgmbi ihnb 1:104 [78].
6nigi5nb hoijjjni 9(536^336^3 ibg;j6b 5 iX 3 6 ib n0g36ig, hi3g36igbi5 ng>n
3mj6 3936b b ii336n 6n3onn3h3636nb b3O0nbi63gm0mbib3 ( 5i 9rngno3>, S ^Q jbnh^)-
6i, bsj»335 n 6n3om3h363bnb 0m6nonb3bi, nm636nb 0n0 fn(530i). ib3ain 3 6 6 0 9 3 -
6 30 ^ 3*6 3^ 1 3hcnn3hcngi0 m3ng36x D 6 nb 0^5^000*3611 6^63363&ni 6ib. 3 .20 . d p n ^ h
09*33 6nig:>53n Al-nb, Fe-nb g i Mn-nb nro636n 5i3fnaii3nb3ogc9D6ni6, b m p n
Ca2+, Mg2+, K+, P043' g i M0O2' no3fobni6 (5X *0 03o>3nb36ign 6i3hoi36nb b^bnon
(6ib. 3 .21 ). b0 nhig N H ^ /N O j" o>i6i(3ihgm6i n(5X06i i0m6n30nb b ib ih 536 o m e -
tIIWO 6nigi50n qjrnbcgi^n, Fe, Mn g i bm5n3ho>n ©nyimxoSDbjjn (5i6bij)3orfi3fi>ncn
Zn) 5ig ig n i6 d63gig bb6ie qjmh0 i 9 n g i 6(536^3360 5i6n(5gni6 i 0 XD636,»>3-
bnb 6ij)036m6ib. 5ihgi i0nbi, ODOD 6nigi50n 6mhi036n 0m06i0oi3ig 0 m J 0 3 -
93636.
3 .6 .2 . m sn in cD h n qs wconqjDbDhn ai536^hD36n
cbm5n3hoin 8(536^3 bibog3&i 8bmpmg jjnhni6 g i 2>ih6m6i,iX 6nigi5b3, b m -

Om bb33&b 3hfi336nioi ^nhnoi pihn6n bnpn2 ^ 36 0 g i J3n9m3i6n 6nigi536n. 6 ^ 5 3 -
6ihnb 0noh SibfH^nyib a0mM!33nb 006393001 Sondp^bi 5i0fn3ihRnmo) 2>iC‘b r|-
ifnp^bn (j,nhni6n 6nigi5nb 0my3ihx36n) g i 2>XH5nqj3b3bn (^nhnibn f i n i p ^ n b
0 mdxD 36 n). ijbh. 3.9 fii6b, hm0 ibom 0 m6 iOi3b i 33 bib3m&36b bb3igibb3i ih»3-
ipn ibibnioioboi (3njjihnxn bib3m636n); 3j)ml5n336n 03ndo36i 6 ih3 mnJ96 ib 3 ) 1 -
nn bib3mbnb <3^503600(5. 0(536ih3Oii 3i6nli>i0Or'i6 gi0ming3bx36nb 0 n b 3b 3& n b
Ijbhnpn 3.9 33hminb SoijbSn onjphnx bib^mbiaii Qnjh Sjhfiax0 3s6n(j|i(j 3&nb 0ij)bOr,cr>D* n
(Landolt, 19 7 1; Ellenbeig, 1986)
63^hn jiCUnl fr'C3*1n bibyn6360 i 2>C0,1^D63tln bibom6o6n
Achillea atrata moschata

Carex firma curvula
Doronicum grandijlorum clusii
Gentiana clusii kochiana
Hutchinsia alpina brevicaulis
Pedicularis rostratocapitata kemeri
Primula auricula hirsuta
Pulsatilla alpina sulphurea
Ranunculus alpestris glacialis
Rhododendron hirsutum ferrugineum
Saxifraga moschata exarata
Soldanella alpina pusilla
- 204 -
3063(^^7)6 6030)0060660)6 ^ O b 016301)367)605660 bb36«?6bb36 366066630
9*9536* iyj'bie^tvgimcm&rvphn J>3C33nlj 3hcn-3hoin DdsxQbn 9* âoi^hQbn *b33-

ifeo*.
^nhncn 8gog*ho 60*9*50 (t,oh6*ho) jjnhncn p*hobnb5*6 5*6bb3*393&* 8389350
CTi^nbQb^h^b^bncn: n5n 3qjhm 6 y*05 *3 ^*hn*, * 8 0 ^ 8 3qjhm 88h*go g i oibo0o*,
309(13 bnon^^hn 60*9*50. ioh6*hn 933Cit >3 Dgjhm 08001 5*9mohfi33*, hm8 8 *b9 o
Ca2+ 9* HC0 3" 2)11161536^*150* dpn^h 8(118*^36300*. *8nb 5*8m, jjoh6*hob h3*j-
(30* Sjhy^mfeb 6QO^h*Q3hb* 9* b^b^ 8mhnb. i'bmjjfi jjoh6*h8o 3qjhm b6h*-
03*9 9n63hiQn‘boh93b*; P, Fe, Mn 9* 9do83 33^bQ3&nb 3 8 3^ 1*16 * 3qjhm d63p*g
b3p3nL)^63gm0n* 8i536*hob*oi3ob, 3ngh3 89*33 60*9*580. inh6*ho, *8*33 ghmb,
i*h&m6*i53C 6o*9*5b*u 6*h8m*g536b. *hob *5h30i33 HCO3' 8gog*hn 60*9*5360
(8*5., MgC0 3-ob 838153300 gmQmSo^ho j*63do), hm0036b*i5 bm503hoio jjoh6*-
hob 9my3*h3cn 81536*113 5*3h6ob (8*5., Helianthemum nummularium, Plantago
media), bmpm ioh6*hob 8mdx3f>n (8*5., Anthyllis montana, Saxifraga longifolia
30113633630) io 8*01^3 b*bQg3bo*6.
boQni)*ii)3ho 60*9*50 83*33 9* oinbnb 8*(?*0O 338 |Ji33Cm6 obib 8j>3ho30, ^360-
*6n 9*, *j3g*6 5*8m99o6*h3, 3qjhm 15030*, 309(13 qj*9 *ho 60*9*50. 8*b8o hj,o6*
9* 8*65*6380 9090 h*pi936m&ooi** 9* *030o6ob 6*3(101360 *9300*9 3j 333g 3&*h3-
6o*6 5*bb6*b. 6o*g*5ob y330* 3b 9*b*bo*oi3&30 o 3mj83g36b 3(j36*hob 6030103-
h3bboi* I530obi 9* bhgnb 3hmu3b'b3.
8(536*h3cn* h3*Jjn* *h*b30b*yh3pn b36b^h*,jnb*g3o 8h*3*053*ho*. ^Q bnjn -
Qjhn 8(536^3360 360* oy3636 83^33300 2>oh6*hog*6 cgmbqjmhnb* g* 8o])hm303-
836^3606 d630*g 83Oi30b3&*b. 633b 8y36*h3b (8*5., Fabaceae, Brassicaceae,
Solanaceae, Cucurbitaceae m5*b3&ob bb3*g*bb3* b*b3m&3&b) 5**660*01 3.6. hyi-
60b 3033^11, hm8gob *hbo 8g5m8*(i3mbb 08*80, hm8 q)3b33&ob *h33o 3hm(jm63-
60b 6:>i*gob 5*thg* 06333b hj>o6ob 83mh*g b*ljh*6b 3 mh^m qjmh8*g (1303150113-
6*b 9*, *9*33 ghmb, q}3L>3ob 5 * 8 ^ 0 3jh3g3bob 5*8(1*3036*^ h*i5 630b 36ym&b
83oi3ob36nb 3hmi53bob 5*d0O3h36*b (6*b. 3 .22 , ob. *51130133 6*b. 3 .2 0 ). 013
Improved Fe Iranslocafon
to the shoot
6sb. 3.22 hyi6nb e^qjmjn^nb 5*3036^ mh(vj65n*6

91336^ 301! qjjbsjb’b] (Lucas, 1987)
- 205 -
8 0 6 0 6 ^ 7 )6 6030)0060660)6 300)30b0&6
(jbhncn 3.10 9(j36ihooto 9mhnb yih-J(vnhn'bnbs gi 3p39n6nb (ynjbnjjhmdnb ck&soncrvj&n (G rim e &
Hodgson, 1969)
inh-jcPthm'bMjsedn Snghjinpn y*6iht>3 iC39n6ftb anîjyicn
dm'fcohgil SSCU(iqj35în 8(336^ 33^ Sm'bihgn JXH5ni rr’C3hn
Avenella flexuosa Hordeum vulgare

Holcus mollis Agrostis stolonifera
Paspalum dilatum Festuca pratensis
Lathyrus linifolius Beta vulgaris
Galium saxatile Medicago sativa
Eucalyptus dalrympliana Asperula cynanchica
Eucalyptus gomphocephala Scabiosa columbaria
Eucalyptus gunnii Lactuca sativa
j>sClSn<3DdDh 9b36ih336b inh6 ih 9 n fcig^h&^m, 0153b ^ 3 6 3 6 ojmbcgmhnb 6i2>CD-

6m6nbi g i h^nBnb 6:>xC)6m&nb 6(196360 ("inh-jpmhm'bn"; ^jbh. 3 . 10). i 9 0b hn 3
y33Ci'b3 3<3hm 9j>hd6m&nih3 i»XiiD^DCu63 J)mîiinDCn 81336^3360 ihni6, 9^ 5-,
Citrus-nb bib3m636n g i Ericaceae-b 633hn 6 ih9 r»0 >gj)36x n (hmgmg36ghm6nb
bib3m636n). ^nh-jpmhm'bn ihnb 6n3onn3h36icni U3pnb 4m9303jb3hn pih fp333^,
hm930 ni5 3b3&i Bn^hi^nb, qjmbqjnhnbi gi 9gi33&nb b30ib. inh6ihnb dpn^h
9 mdxD 9li36ih336n HCO3" g i Ca2+ 9n9 ihoi b30j,hd6m&n:>h3 ihni6. ^mhqjnb
bi3b3&n g i ^nhBihnb 9 m d x 0 0^1330(^36360, hr^mhnui!! 9 ig>., Avenella fle x u o s a .
HCO3" gngn im6i536(j)hi(jn36nbib q)3b33&9n !5hm33636 gngn himg36m6ncn
9i0i^b, hiy n6333b bhgnb 3hm|j3bnb 8 3 ^ 3 6 ^ gi, ^ 3 0 1 3 3 q)3b336nb pi'bn>-
636ib.
inhnoi pihn6 9gi33 6 nigi{,9 n ihb3&30n hin6nb, 9iB5iB39nbi Qi £>s6b>-
3aih3bnoi !030n6ob nm63&nb gng9 i 9 ihij,9 i inh6ihnb 9 my3i h x 9bD 6ih3369n
3ndQ36i 5 i 9 nn63nmb i 9 nm636nb bnîh&3. i 9 i 33 6 nigi 59 n Al3+ îh6n S^cn-
jnb3&i bni6b iy3636b iiCl5nirnCDh 0U36ih336b ((jbh. 3 . 10 ). iQ}9 n6 nb 01*16360
n^ 3 3 6 3 6 j)i0(5n39nb nm6;)6nb ihbb, hi(j n6333b ii0b n39-iic9nigxn6nb bnb^^Qnb
93*JK*JibD6ib. i 9 ghmb, 9 i&., bg3&i Qn^mbnbi g i 33h3gnb 3 m0 ih x n 5iB 3 p 3n b
3hmi53b3&nb 9^0^15363!. jjnh6ihnb 9 m dx 3 9b36ih336b ^n 933d0nbcn 9 dn9 ^ 6 3 ^ -
i036nb j)m9303 jb3hig g i 69 i g i A l3+ nm63&n 9 ioioi3nb 33636300!.
03gih3&ncnn gnbnmgm^nnb ib^j^ncn 03nd03&i 5i9mnymb 9(j36ih3cni
] 9 nb 93tj>i6f»i0nb9nb mhn (jnSn (uo)nbnml5n336oH) [ 1 1 7 ]. >)SCUnd^m°3DCn 9 ( 5 3 6 ^ 3 3 -
6n n9ihi53&36 6yi03n bb6ig 9 i0 i(jb i g i ^n^hi^cniB giK i30nh36x . j>i0Un09k; i 9
930oib333i9n 3jh3gob 6336n 93nî3b jqjhm 93,5 Ca2+, 3ngh3 K+. 9 h i3 iQ n
B rassicace ae , fab aceae, G eraniaceae, Euphorbiaceae gi, hm&mhtf iji9 g 3 i i56m6ncn,
y 330 i C rassu laceae ihnb jfidjn^hm qjjQ n. ^pynqjm bjh n 0|j36ih336n Ô ljn ^ S n b
9i(piC im6i336(jhi^n36b 36hd3ni6 Ca-nb j,igiy3i5ncn îgy n30-mjb i0 i^ 9 n g i
33j^n6cni6 9nbn g i69noi (6ib. 3 .23). rvjbiOi^nb ^nib & i6 3i30 i36 3& n i6
C actaceae, Polygonaceae, C henopodiaceae, L am iaceae g i Caryophyllaceae-b m j^b ^-
6nb 303ii3bn 6ih9m9ig53630n. ii0^n30nb 93oi3nb36i b30b 36ym6b mjbignb 9g;>3-
36nb bn6oi3bb. i 9 m jib36nb 6ih0m0!g53603&b 9bm0mg 0i9 n6 933dgn.ion
gibib0g636 j>nh6ihb3, hmg3biu 9ioin gi6j,6ignb 03ly>6m0nb0n nd033i m J jiQ ^ n b
bij,9 im himg36m6ib. îo^nÔnb h35X H 5nn^ 5nQ33 3hoi 93j i 6 nb9b 6ih 9 m ig 5 3 6 b
- 206 -
806066=57)6 6030)006066016 03^01 0)6301067)606660 Ub36«?6b b 36 366066630
(133)13(50*, hr*i830O(5 "bpôhoi SaxifragaceaeA» g* Plumbaginaceae-dn logi^PO -

bnb b i^ o ^ in c n b * ]j 3s m g ijiin ^ h ig 3n 8 gn 6 *h 3m 6 b.
633)1 8380^333*80, 9 jj3 6 ^ 3 3 6 ^ 5*3(1(5x36*^ 9* Ca-nb qjobomooôbb 9 mhnb
*hb36r*i6b 5 ih 333X n *P39nhn (6*b. 3 .24 ), 013815* 3b sh 6*h3m*9536b (1*083
brnftbg (535g36(5f»*b. jjipynohnqgx mj*b36b* 9* 53*113680 *1^360636 objovi b*b-
3m636n, h0180360(5 53 m6 mo*5 ot,3h ibS^j^So _j*ctf/jj/*?cj/j*p & i*OtSn<3JdDh*Q nj(50-
30*6. 3b 3*5*000)0 3633636b, hm8 331*10^503(10 606*30)1(^6360 0830*01*9 801^83-
93636 IPQ43X <3(1^11*101*150^ 113*^036^3. biôhm* y3X > sb io j^ b 5*oi3*pob-
60636*. *b3, 8*5., 8|536*h30i* b*b]m636nb* 9* 33mlyi336ob 5*3)1153036* 3oh6*h
60*9*5^3 5 * 0 m633xrc» ^ 8*h|yT' 35h39ob 3*01503806 (5300b oi*3ob363h3636ooi,
*(1*839 ^5)130133 *3(50036300 6omx3936o36ob (8*5., (13060b) 530i3ob^6ob o6*hob
5363^033)119 (goJb0)136x^1 8oJ*6o<b836ooi.
Silene Anthyllis
1 .6 -
c
2
61b. 3.23 ôuoqjno^h 3s6 ryji^ 3n dmbihg

bb3igibb3i qjnVifnjrOnb djmS] 9u]6ihnb
qjmoimpgn iipn^dnb, iiCUn]3 nb p* 3i{,6n]0nb
933u33Cfi6i (Iljin, 1940; Horak & Kinzel, 1971)
Ju n e u s N ard lis Origanum
6ab. 354 iicunjQnb bb3*p*bb3i 339^33pm6nb

3nhm&3&3n i(jneino)no3hn !PC(ifx5J363hn (Juncus
sqtiarrosus), ^00005353)10 (Nardus stricta) gs
>)*Cl5n.!>m0!3hn (Origanum vulgare) 0(j36:>h33&nb
bhgnb oii3nb363h36i 5o (Jefferis & Willis, cit. 4 10 50 100 500
Kinzel (1982) 8ob3p3noi) Ca added to nutrient solution (mg-l'1]
- 207 -
0O6Orf^;3<*> 6030)00(60660)6 300)30l>066
3.6.3 monanibhmqoe rfcfiiWiM&an anrbên aisD6^hDD6 n
m cnan^rKgxn 3^60^^360 nb^cnn ip^np^dni, b^gx5 9(j365h]at> "bhgobi Qi

5i63ooiih3&ob dnhnoogn 0^35300(136300 5ilv)0m q jij^ h n i bij,33&n 6030103(1363-
6nb 6iiC36r»i6i. j>i6bii)30ih36ooi 36 n03630Pi3i 6 oi ^ 30 10 X3®3®\!)3^n^ 6iyvj6m -
6i, hm5mhn(3ii i'bn^n g i qjmbcgmhn. iQnb 9o'b3<boi: QnBghiQjhn 6030*131136361101
piho6o b^&b^hijjn (9 i&., j,3ih(5ni6n j 3n36 ihn); ^3^3, g3j)hbgnh363Qn 6nigi£)0; mh-
j,i63pn Ri0 m6i(5336nb 6xf» 0 ^> ihibh30o gs0 Qi- 06 nh, b3 nh:>p n/noqhmjQSQ ©3^33
g i J3n0ni3i6 6 0 ^ 5 36 ^ )3 3001^936(1^6 biODhmqjnc^hn 6 3 ^ 3 ^ 6 0 q)mh9ii5036n
(fc>3ihi0O, ©i^mhbpn, qjo66nbn g i bb3.). ihJdOixn (j^Sghnbi gi 9ipi00oiob 1503*1
6oiQij,n Ri0m6i(5336o bi^)6ob 9 o63hipo'bnh3&ob 6300 (j 303nb, 3303bob g>i6-
6bb. 3.25 ibm^nb

6iQi5bn Rhpncmjcn
93393cnnb îmd^&Sn
(Rosswall et al.,
1975). 9n63hi036nbi
p j 6ib 3 nh5 ipnb b ip -
}6bnb 9 m p x n
ihiynxo
^sro&36nUj»3nb nb.
Sonesson (1980)
Ubhnpn 3 .11 63n9nb Snjh 9n63hi<v)hfi DCD9o6j 36nb (5(1 9 ® s'*) n93mh^n (Golley et al., 1975;
Kallio & Veum, 1975; Likens et al., 1977)
^ m S n iX " 63n9ro5n Ojmcnmc3i53o6n J35$hs (bj5Sen&>3n>)

(li06i^tiC3hn iS^hny.) ojihcnmojnoicmjsen
(Rhgnenrjcn iSahnji)
N 2.07 0.07-0.1
S 1.88 0.5-0.6
P 0.10 0.0004
ca 2.93 0.22 0.25-0.54
Mg 0.49 0.06 0.05-0.15
K 0.95 0.09 0.08-0.12
Na 3.07 0.16 0.1-0.4
Fe 0.30
- 208 -
8060A 6c;t)(f) 6030)00(^06^0)^ 03«=?0b 01i 30b06U(f)06660 bb3^ b b 3a 366066630
353ni6Q6nb g i b3 nhn Qi,i36n!636nb 2,i3m, 6ih9oi:>g536b i’boijjnoii g i qjmlxgmhncn

Cfihn6 igjjnQbiQymqg^cb (6ib. 3 .25 ). 3 o63hi03 hn Br^cnn^h^b^fiincn îSbiiônh^fencn
(j'ihnbni îoi6nbi g i ^mhqjni6n 6niQi536n. ij, 338360b Bi^hoijftnb gngn himg3-
6oi6nb, donjh gsbspn pH-nbi gi fth^j^nb 6yoo&nb gi^îjftnb &!0 m, QnB^hip^h
6n30in3h36i0ii yn6t536l>)h!UO! dipni6 gi&ipn!. i 3 n^m0 , gngn 66033630016! 360-
^ 36 ! 9n63hi03h 6n3oin3h3&!Oii 338mlj)i5ib (jihncn am^Bnpn 0v|)3hnb! g i B3n3nb
bibnoi) ( "ndbm^hmajnb " ybh. 3 .11).
3n63hi03&ncn pihnb 3i6nl!>!li)363n 3m6n6igh3 8i536ih336n, dnhootigig, 6*63-
i3cn363&ni6 5>ihj)333Cn fT'hj>i 5 ntbi(jnnbi 013 ‘bhgnb qjmhSib. ^30 1360!: 13030 h3-
j,nrn636nb 00^36360 (6i6o0oi6ho3 i‘bm(j3ignjbnh3630n q)Oioi6noi6lj3&n g i 33nqjn-
l!i36n); bi3b3&n ^ g h i b i g i ^!0i6363n; 6ipibm3i6n 9(j36ih336n, 0!Oi Smhnb t,mh-
gni6n 6ih|j3pm3636n g i nbp3&n; 3isnh3 boi8nb mhQ366ni6n 6i0ibm3i6n 8(536!-
(13360, 5i6bij>3onh36noi h o r t ^ x 0 ojnhSnb; 3 3 5 ! 63(^360, bSohig S i^ h i 3 ihig 66 -
3i63 <30101036001; 3b3333hj6oi6n 8i536ih33&o, hoi330 oiiu ibibnioi36cn 0301010360b
b303hm0mhqj30m6nb 3ipi0n n6g3jbn (% 63g0n 6oi^$oib 93q]ihg36i % 63g0
5hrnlj3n6cni6) [ 147 ]. Ericaceae-ebisa gi 8nbn 8m6ioi3bi33 oi3ib36nb 6 ih 0 oi0 ig-
5360360, hri8036nu bib0g36ni6 iboi^noi (jiihn6 Si&n^^bSn, 63330n63^ni6 hm-
2>oih[j j30m(i>ijbm6m3n3hn 030^001001503(10 ^nSo: 6nî^h3Q3j(ii'bnb ij^n3mbi
3 im3 n dpn3h g!6ipn! (nb. (jbh. 3 .7 ); s0 oi3ib36nb 633b 6ih6m0iQ53630b ij3b
9 ij>ihn, 005606001 3 gngihn 0301010360; 03ii!6oiQo'b0n 0o0 ihoixoi 136083606 Qij,-
hm336nbi36 gi 8n63hi03hn 6o30io3h3&36nb 93ai3ob36i dpn3hg36i qj3b3nb bn8-
60016^0 ^31500303(10 b01501360b 83333016001 (3(1030105300 gi ih63(iOingxn 00301-
hn'bi).
0n63hi03hn 6030103(13636001 5i6bi330ih36n0i cpihn6 3i6nî^363n 536 3536!
3 i(3'i 0b53(jn!0 n'bnh3i ) x n'J i*jnl3mlS*:)OfT1 <3^(10360, 1101501(1015!* 330030,331™ 03hj6ni6n
3l336ih336n gi 801(10301001503(1^ 013 (i)ijbm6m3n3hig 3hi3i0<33hm3i6n 0lî|53-
60360, (10183001! Soihob ihni6 (101501(115 330030,5360, 0G3 îm&nb 01536^3360.
330030^360 156013(101636 mhj)i6x b3&bj(hil!)lb3 g i 3n63hi03h 6n3oin3h3636b
noi3nb3636 !^3mbq)3hmQ!6. 6 n3 nbi g i 3h6nb qjoih0 nb 0301010360 Qj&jhmq)nQnx
3 ij,., Platycerium-nb g i Dischidia-b 030cnb333!0n) g i di6hnb3&hn hort^^&o
(0 !^., Asplenium-nb bib3oi63&n gi Bromeliaceae-nb 6ih0m3ig53603&n) !5hoi33636
6i03j36bi g i g3£hnli)b. 030101036^3 5 i 60 i 6 36300 33660130 J3h(j036n iig3n03636
noi63&nb 33c 633ib Ri6oi6ig36n 6yo36ngi6. 8oihcgoi0oij)n3h 6ih6oi6iJ6636b
333 bi6 i 836i cg36junoi6i03hn 33 &336i 6n. 33nqjnl53hn 53n8h3&nbi g i y3i3nooi3i6n
6l536ih336nb 8n3h gifthoi3n0 g^hn^Bn ndyoi^36ni6 6ij^3hn36n, boi >(01360 gi
40n
M 350 m
□ 1150 m
30-
rffl
20-
10-
J Bib. 3 .2 6 fihp. 333eocnnb g i b^Qn g i 0 i(n ip n
bfidi(pQ 336n g i6 ih 3& x Q u jB ih ^ o ii qjmcnp369n i'bm ^n b
\ 5i 6i 6n036i . 0i(n iC bn9i(pc33*^>3. 93b i 6i 3 n b ig , jih ^ b
M 35f>Cm6(i]h 3nhm6363n 9mbihg 9(5]eih]b ijsb 3qjhm
50 100 150 200 U i^ ih i ‘bmSnb cgmcnmcn g i, i3 g 3 6 ig . i t m ^ b
Nitrogen concentration [mmol N-m 2] 9 3 9 u 3 X 'T* i 3 ib 9 n b ^ i^ n b ^ iijh ig O ^ n i (K o m e r, 1989)
- 209 -
80636^36 6030 )0 3 6 0 6 ^0 )^ 330 )30106^
(5601339360, (10180360(5 01^151630 5 ihgi;j96 ni6 9(5361(10601306 ig 3 n 0 > e -

03013063619 <301(1819.
9 vW5D^Xn 9l3D6 bh33&n bjopn 36901301^93(10 h3onj)XD9 noi gicgihxo jnhj)3 C rn'
3160 33(139360916 5180^010936 80186303630 36'bo8366 (3hoi(i)3i'b36n, 333oog.i<fc>3-
60) pi 30*33 32^393606 83833016001 0013063636 18060183133661 9* 0 0 16 36 b .
51(19* 18061, 8016393630 (5601339366 61^3(10360 363336 8063)1100130)13616. ©O^bD-
&pm6i ihnb b3hbn, hm9 pnb 618310360011(5 8(336*113 i2>o)8336boh3&6 i‘bm(j)nani q >
cgmbqjmhncn piho6o 3*600*03606 1(116396^(139 51393616. 150006 8m63p3£>nb
03033m6nai Pinguiculab 6163016360 0(^3636 61^0(101 I’bm^nb 20 -6 0 % 91 03m b -
ajmhob 35 -80 % [ 111 ]. 01(1516300 61213360 6030103(1363606 383*0019 5 18 0 ^ 3 6 3 6 .^
b3pb 36ym6b <3010103606 66(1109 5161603616 9*, *^39*6 5180189061(13, 0136 9 36 0 6 ^
9 1 8018*3*90 1(1180161(53606101306 bi^nhm 01(1516015363(10 6*2)3360 6o30i03h363£>nb
915(10133616 [216 ].
b*j)336n 6030103(13636001 ("1(106 3i 6 ooioo60 o 9161693639 8(5361(13366 * b * -
1*11013601 3(33^03^01606 6ohi00 &n* [59 ], 8obo 83833016001, 612)3360 6 0 3 0 10 3 -
(136360b 6i2)C36oi6ob 803639*3*9, nbn6n 061(16363636 2)0162)3(136(5006 361(16. 0 i c n
033dpmoi 5110)903(1016 8n63hip3hn 6n3cnn3h3636nb 03oi3nb36i (sbbmhbynnb j c g y -
jiijh m dx 815., <33633606 303(101 88913(119 5*630011(13606 9* 51801(10 33h 3P3bn b
(110193601606 5i‘bhgob bi83ip36ooi) 91 5ii383oi63brTi6 8o63hip3ho 6o3cnn3h3&36nb
8ihi5o 093633606 603(1(5380 (dmdnQnbjdnb jqjjjtfjhmdy, 815., <33633606 B n j h
83133606 91 d3P*ii>nb 8382)3(1303606 5>8mymgnoi). 8n63h3p3ho 6030103(13636001
piho6 9 1 89*33 6oigi536b3 Smbihg 8^36*1136 *6*60101366 dmbdshjdnb j q g j j -
tfjhm di (h3ohi6bppij)Hioob 3<33dti»3hrn6i), h*u 8ib 6183193616 idp33b 6 12)336 0
6o3OT03h3636ob 6398301(139 5191616093606 83833016001 01130916 5i8oioy36m b
obo6o ib*po mh5i6m36ob 83bij863pi9. 3(10169016 8(5361(133680, a^bpo&nb
33 j0 6 ob 3(101(53680, 612)3360 603Oi03h363&0 (5*66*21301(136001 qjoibqjmho) 3 3 ^ 3 -
0i(jr>3ho mh5i6m36ob 61(53919 d*poi6 9090 (1101936016001 8036019361 hoShm-
93ji2)n3P 01(151601366. 8(1131969016 8(5361(133680 6001393836(5360 5(1013936^
5i0m‘bi0cnh36nb 01(1516013600; 6o3cno3h36icni 8(1131933(1190 h3ljhi6bpoi2)*i5r»^
610 3193616 idp33b 8(5361)136 0(56013(1016 612)3360 6030103(13636001 pihnfo
3i 6 o0 * 0368 o.
(303 h35om6360o 0h*3ip6pm6o 8(5361(13360 ^1(1601636 3(10^016366. 8 1(119 0 6 -
316 3 <3010103606 60(501(56906 6165(1890301616 05033 3(33^0 ij3b; gmaip^&nb
yori33p6po3hn (5391 86019019 61609016(103 69361, (10011(5 8(5361(13 01130916 0150936b
<3010193606 (5336001 5180163339 8o63hip3h 6030103(1361011 2>ih53i6. 369015363(1^01
913(1015(11836390 “bhgob 5331 <301(181 (536(11^033(119 91 801(103019015031119 5 ^ 6 -
30(101636390 “6(1906 3hoi(53bob ( 1 3 5 x ^ 3 6 1 301(180163606 83833016001) 119 3 0 0 3 6 b
bi333&o 6030103(13636001 bi2)8ioi hioi936oi6ooi 80181(1153616 91 I'bm^nbi 91 qjm b-
qjoihob 303(101 3093^43^19 5i9oiy3636ib. 163010 b1831036360b 5180^3636001, (30 3
9 1 1800018 8063(1393(10 6030103)13636001 piho& 3i0ii00aiob (13500163680, 6 i0 ib m -
316 0(536ih336b 833800101 iboi^nb 309(101 ©3,50 2)^6(5360(11(503606 83^861, 3119(13
916901606 8(5361(13366 (61b. 3 .26 ). 06301 31600*153680, (101501(10(511 (5369(11 pi,
81(^108010163010, y330i'b3 61(181(53630360 ihoi6 ob 8(5361(13360, 1101893661(5 1^300
ib ndnp^j&nb hjgndnb ^m6bjh3^jQn ^nZn. 1600103(119 33<bp 39 x 51(1380180 1( 16 3 -
60161 80100160136 83O*6oi0o‘b8o6 91619 0603660301616 - 080610130b, hoi8 lb h 9n b
03‘bp393ob bih3'b3 83oi6oi0o'b8o6 y330i 3hm(53bo 8089061(13016936 21(18016030^9.
- 210 -
9063(6 6 ^ ( 6 6030)00(60560)6 (603080 30366(637) ^ 1636(630
3.7 ari63hXDh 6n30in3h36^ h39nan

3l536I)h33C b ^ h g n
3 .7 .1 ai3 3 6 ^h 3 DCn b ^ h f l b 9 n 6 3 h^CDh 6 n 3 0 in3 hD6 X n ^ 6 ^ W 6 b fl
0U36ih o x n biqjihob 9o63hi03h 6030103(13650^1 hogn9 n 6ij)i39oh3630oi 6ib-

9 nh6 ignb h3go9oii6. 9o63hi03h 6030103(13650^ 33ai3ob365 5h3530oh36b 9u;j6 ihnb
9 i>bnb 9m0b^35bb, bmpm 65b3oh&5gob 5bo8o05uoob Shm^jbn 3<bh363309mq)b n9
9ibi0nb 03j96ib, hrn3309n(3 Rinjb(ii336i 9063(^03(10 6030103(136360.
3i)rnCrn£>n3hn 3hni03 jljn30 (ii6 ob 5i6li)(ii036o0i6 559m9go65h3, 33bid93&3905 8153-
6ih330n bicg^hnb 903(1 9063(1593(10 6o30io3h3636ob 60(i3hn 5ai3nb36ob 55911101395,
013 u6n6n0(ii 6it5ihob 939 i5330m6 i 95 33859536011165. 8(3365(13300 b5qj5hob 9n3h
cg^hcnmbnb 3hoi330lb3 600b 5 i 69 i 30(ii6 i 9 o icn jn b jd jp dnBjhspjh 6njcnnjhj&^cn^
tumgjBmdsb (Matis) 5^99365 ^>9^33 639b h3ih3(jnnb bibnoi 558mymq}090 9063(1-
593(10 60301031136360b (15(11936(1165 (Mr), (1111939015 559(110113(56365 6593;)360b 9n3h.
8(3365(1339 bbqgihSn (16365 8063(1503(1 Bnjcnnjhjbscrts n6jmh3mhnhjdjpn 6s 6npn
(Mj).
Mabs = Mi + Mr (i 5 3b-lyl). (3 .1)
815365(1330 3 ib>3 n 95(1636000 8063(1503(10 6(i30in3h3636n, 3hrn03 ji>)O30 (ii6 ob

5i6^rn036nb (2.21) 33bi6b3nbbg, 65600936056 690b 5i 69i 30m6 i 3 o d m d ^ jb jp
dnmdsbsbj (AB), 6i9m(3336ob 3393559 (93ljhoôb) 98(1590 6m6ob 6003b 39365^3
(M[ ) 95 ^m BbjdjB^jbnb dnjh dsbnb dm ijnQjdsbj (Mq).
Mj = AMb + M l + Mq (1,5 3s’1 y 1). (3 .2 )
559010139360 6 ih3 m36b 33(150 3 ibi 3 n 8063(1503(10 6o3oio3h3636nb (93 bi 6 i-

3 obig 6i(}sh3 o) h5m036m6nb 9595360b 93333(116001. (159556 8(5365(1390 60(11303-
936^36(1 ihiOii6 i&h5g 656099365 9s 9063(1593(10 6n3om3h3636ob 339lj3O0rri6i
0(539365 600b 5568539016530, 8063(1503(1 603010^36^01^ 6i0i6bob 5590101395 3hm-
93jljonb 5i6^(ii036ob 633(1015 36(15001 55958(153936001 6i^hnb 3391533911160b b^33>
0n 8(116x53336^3 ih 33039365. ob333 (101501(1(5 6 ib 3 oh6 igob &50i 6 bob 9595360-
bib, ijiu bi ôhfiii 3ibi9ob 5^365 bb35g5bb35 ghmb gi bb3igibb3i mh5i6mb
3ob3g300i. (jbh. 3.12 3mi339390i ^50^39863563 95 3ihig963i63 ,593360b 8063(1-
i03h 6o30i03h3650ii 6i0i6bo.
013 hi 6090 9 mgob 690b 5569539016580 061)01(1301(10(13639 9o63hi03h 6030103-
113636133 gi hi 6090 - Ri0nh3ubo9 g i (15801(533609 S ib i^ , 83039365 959506-
93b 6(136306 ^1*3030(3036^ kM bi33i936ooi [257 ]:
kM = (ML + M r) / M abs. (3 .3 )
86nhn j330>yob 8^0163 6id36ih9n ioi3ob3&30o K-ob 80 %, N-ob, P-ob g i Ca-nb

70 -75 % 95 Mg-nb 955690136001 60 % 3556 36(1369365 605955b [2 5 8 ]. 6i0iboi-
3i6bi0)h0i6 6003965(1 ,59380 N-nb, P-nb, K-ob 6h363ob ogij^mhoi giib0fii36ooi
- 211 -
Ubhrxyi 3.12 3n6ohiC3h 5n3oin3h]6ioii (y33Ci 3n6i|339n jîo^cnen* e>h 6-e y 1 ) 611363* ^33660 (Duvigneaud et al., 1969; Rapp, 1969; 1971) (TC
ihnb 3n63hiODh 6r>3cnrp3h3&:>oi> jsSn, hmSjcny 33^ns N, P, K p* Ca b i3hcnm j>3 t>3, 3n6inpi6 Bi^hnb bb3> SCOÔ^^&biu)
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3 0 6 3 ( 6 6 C ? 7)(6 6 0 3 0 ) 0 9 ( 6 9 6 6 0 ) 6 (6 0 3 0 9 0 9 0 3 6 6 (6071 ^ 0 6 9 6 (6 3 0
9 0 %, brnQm Ca-nOi g i Mg-nb - giib0Pi36noi 80 % . ^hmin^x? 63n9nb ^yxjbSn

Ca-nb g i Mg-nb 6)136306 qjij^mhn 83595360 95 - 100% [ 171] (ij3gi6 1/3 3ihjnb
Ri0 mh3i3b3i<b3 9 mgnb), K-nb - 85 % (6ib33ihn 9 mgnb 3ihjnb fii9 <nh3(jb3i b 3),
bmpm Fe-nb - 50 % [7 9 ].
3.7.2 ^bfliWiian an6Dh^C3h 6n30in3hD6^01i 6h363i
3n63hi03hn 6n30in3h3&36nb 6h363i9n 9u36ih330n biqjihn 96093630^316

hm0b ibhxD^*- 9n63hi036n <33033600 Oi93i036noi 6nigi&ngi6 93061*1336^6,
6nigij,nb ^giSnhnO 13330(31 &igiig5n0g36ni6 g i qjmoiri0U336nb 930 g 3& nb33
6nigi&0 36)136936016. <bmj)n3hcnn 91536:^3 (8^5., 93b!) dpn3hig g>i63noiih3630n
0330301} Onb^30nb Byigm&noi, 9n63hi03h 6n3om3h3636b n6m3b 6nigij,n0 516-
Oiôih^bnoi ph9 i 9 h336ngs6 . qjmaipiQ^Bnb 939935 9063(1103)10 6n3cnn3h363&n
2>i0 ibm3i 6 9 (536ih336biU b9 ihg36i (61b. 3 .2 7 ).
3 >i39r>dhifTii>nb 93 ji 6 n'b9 n 6mgi&bi g i 9 ^ 36 ^ 330 biqjihb SmhnO 9 n63 hi03h
6030^3(1361011 U30nb 96n9363pm3i6n >)m93ni636ijni. n0 , hm9 1*1(12,16300 Ri9 m-
6ii5336n (hmQpnO himg36fn6i 9|j36ihob Shm gjj^nx^nO 3hm3 fnhyn30 ni) n033
3im0n0j)39ib 36(i36g36i, qjhnig 060936300131601. bib3jnb 93î6nb 3j,m0mj)O3h
Oicg3d33Q<b3 gi 9 yih3630 O 9 md(r>3h35i 36gi 30 yih 36mg3b i 9 9 m3036iO: 130093-
63901 9 mbi30nb i(j>3&nbi0 9 (T>303^30° 6133360 6030103(1363600 gi6ij)0obn 9303-
.1\\ , ;i
Vv;V \ y :'M '
6sb. 3.27 &x&nnb
(djbi, (p)3b3n, Bnqjxfi) 0n6yi- Leaching
sQ^n bnjrTfijnybxra ^njjjvib 0me- from crown
X3>5 n
^ynbsc^nb nb. Johnson et al.
(1982); ns3m6nnb qjronrn03^3360
gi 3ihse963i 63 ljy336nbjoi3nb
nb. Tsutsumi (1987); £hfn3n£xn
63n9ns6n ^ynbso^nb nb. Edwards
(1982); b:>3s63&nbs<7t3nb nb.
Egunjobi (1974); tjmcJnQhn
bih^ypnb 9p x m0^n':>;’cn3n'J nb-
Rychnovska (1993a);
0n !li6n^>^3&nbi<n3nb nb.
Gimingham (1972); <b(p:>cdcnnb Uptake
(Retained + Relurned)
63fij6sh36nbiO>3nb nb. Larcher
(1977); ih ^ y ijx 0 iE|6ehnbicn3nb
nb. Bliss et al. (1981), Chapin &
Shaver (1985); i*n&3&nbioi3nb
nb. Dykyjova & Kvet (1978)
- 213 -
80606 . ^ 7)6 6030)0360660)6 930) 30b 3W
bmb bib^jo&nb 93bi6 i 9 nbn himp36m&noi. iSn^mQ, 9 n6 ohicohn 6(i3om3h3636nb

6h]63nb îQmcngpib &iiR6(ii m&r^j^hn 0(1^3^36(1.
SmbD^X 3imbnbo39i9n 3j)30mdh:>(Ti6nb bnRjihnb bibmSb 6ih9m ig536b
QDvb^ni!)^n i'br’i^nb mhj^S^cn bi^hcnQdnb 63(ijni-0n63hi0fi‘b3bnb n6o36bn3fn6i (ubh.
3 . 13 ). (lfcn 033bi6i936i 0n63hiO3hn i'bmonb bnîhb^b, hm030(H5 hR]6i i'bm^nb
5^'bnb bibnoi ihb]6xn B^hcn^bnb imhcrrjc^bnbi pb 0n3hmmh5i6n'b03&nb Qn^h
9 mb9 ih 36x n ibm^nb 5i0PiiP36nb 339935.
bnbo39i3n: 91536:1(13 - 0n>)hp-imhj)36(1^0360 - 6nigbj,n dn63h*cghn 6*130103(13-
b^bnb (jhi6 b3mhon 33g3n3ig 0n0gn6ih;)m&b gi 0ioin 6i6npn bi9 ^gi9 mg
6i. (16333b bij)0img 96033630^13^611 hirng36m6nb bij)336n 6 0 3 -
0103(136361^ iihgîb (0130153, 50nb ^ 3 6 ^ ^ 6303^363 d63pn:>). 0n63hiO3h 6 0 3-
oin3h3636b 333d0(iioi 5igbig2)(ip36i 5h36(jnb 6yQ36nb, qj3hgm636nb Rb0m6i-
h3(5bnb:> g i J)i5n0 ih30 n Bygnb 6ij)ig36nb bi33i036noi. 6i03j36nb 03933(116001 9(33-
63(13^33 g* 6nbgij,‘b3 np3j36i ihimh5i63pn 60301(13(13636(1, hn9g36ni5 5i'bnb,
0(53hnb, 6nbgnb, 013 ^hm'bmgnb bibnoi ihni6 i(j,9(nbq}3hm9n. 9n63hi03h 6(1301(1-
3h36ioc> 3jb3mh^n bg3&i 6i0ib6ihnb 0m(5np3&nb ghmb (i5bm33036nb 0n3h, 9 m-
b i 30 nb ip36nbib), 3hm‘bn36nb 930 oib333i 9 n, Ri9 mh3i5b3fibib gi 6 yip 9 n bb6bgn
6(1301(13(136361^ J,igih3i5b3dbi 013 936m3nb ghmb.
(jbhn(Yi 3 .13 (jnigijnb ^ 39^ Sh^Sn itryjnb SnB^hbcn'biiyii g i 33&3i^l}nnb SjhnmgSn g3(jhn(i)r>Qi6
3n6]hiC3hn i'brynb 3mji36|yi3hn (j,h N 9"®) (Rodin & Bazilevich, 1967; Lossaint & R ap p .
19 7 1; EUenberg, 1977; Janiesch, 1978; Rehder & Schafer, 1978; Nadelhoffer et al., 1992)
0 (5^ ih 3X m&i 3n6;jh:>on'b:>^nnb dn&^hic^hn :><bmlint>

6315^1-06^3^ 03(1161 fciamymgs
Qm^Biy^hn)
(j/imlnyftn
6308(1} 6n (jy] 10-60 26
99hipn qjmoimp0(j336o ^ 3 10-20 22.5
îpnb otO 7
b i3i 6i 0.34.5
b^bmqjcm-bi33]h63m 006360 7-10
btndnjhn luhfjyxn
b03Q<ns33i'b(n33Oinb birahmqjnp^hn (jy3 2-2.5
0ih ig 063i 63 6n63m3i 6n jy ] 3-12 5-18
qjihcnmq)ncn(}m3i5n qjmaimQ0u336o jy ] (2) 10-20 (25)
Fraxinus-Aceretum 15-38
Ônb ijy] (2.5) 10-20 (50)
J»SC*6\!)f>i3hn 5^^0636300 63fiJ6iho (0.5) 1-3 (5)
0g3Om 14-26
9g3pm 13-23
6mi5nm obQni6o 9g3cm (0) 1 (4)
03hipn 3ih(530m3>6o 0g3CPi, b>j33o 1-3 2-6.5
h^gjhiQ^hn 0)336ih3X(,'6i 4-30
go36360b 9(i36ih3X rr|6i 1-3
3iCm<33n(iD*,n 0.2-1 1-1.5
(jnjn hj&rmSjbn
6mh3iC3ho g i 9onob ^3360 1-5 1.5-2.5 (8)
iy>56h3(jocn ^ 3 ("Kjummholz'') 1-2
03hy5iho 15
0ifiC9cnob 535s 63fij6iho 0.1-1
9ipiC9cnnb 093pm (0.5) 2-10
ihJ(jOi3(Vi (536ghi 0.03-0.5 2-5
5S(nihn&363pn în6n 00.5
- 214 -
a n n jo ^ n a o ^ T i^ o ^ 3 n 8 n 030^(n^ 0 8 i)0 6 6 6 aaa 0
4. eycnb hoanan
bnurriijbc3 6 ih0 mnj86 i Byip3 n pi y33pi tfmijbipn 6yipoii6 ihnb gi-

i!33nh3&3pn. Byipn nb {jih ^ri!, bigitf 8n8gn6ih3m6b bnmpmg^hn 3hmu3b36n.
3hm^m3pib 0 i bn^mijbpnb ^6ihb Sbmpmg Bypnb fcbhâmSn i8gp!363&b. J>!0 m3 -
hm&nbib 0nbn gitpj^b ih ihnb i3(5nC3^3Cn> 3 ^>&hi0 !0 ghmb nj,n Qi^^S^^h bi-
bn^mijbpm 8gg,m8!h3m6i3n &ig!gnb.
0^36^3, 303ii3b6npip, 6ypnb5i6 339536!. 3hm^)m3pi'b0 i 03niji3b bi33 ippig
85 -90 % Byipb. 35h3gnb unpncni gi pn3ng3&ncn 3 gngih 6i6np3&3n, hm&mhn^jii
jpmhrn3pib^36n g i 0n^njm6ghn3bn, 50 % 6yipni. Bypncn 5i6bij)3cnh36noi 8pn-
gihni bmhi5m3i6n 6:>yncg3&n (63gpn Brn6nb 85 -95 % ), Bitn q3mcnp36n (80 -9 0 %)
g i qj3b33&n (70 -95 % ). 63950 03hji6n 33n(ji3b 50 %-3 g3 Byipb. Bypnoi y33pi'b3
U>ihn6ni 36 nqj3 oi3bp3&n (y33Pi'b3 jg h n b3 nhig 10 -15%), ‘bmj)n3hcnn (jbn0 nb
g ! 0!{,hm33&3pn cn3bpn 33 nî3b 8brnpmg 5-7 % 6yipb [ 123 ].
4.1 ^nnijncm^nQhDcn amannrranehaen

31536 ^ 336 0
b03P3<jinb 8ij36ih33&3n, hm8p36n(j 3!3hncn ihni6 J,ih330mh^y03pn g i 03g0n-

3ig imhcnjp3636 6yipb, 5i6m6ib6mh363pn Bypnb h3gn0n ihnb bibnijm^bpm 3hm-
(53b3ftnb 6mh0ip3hn 0n0gn6ih3rn6nb i 3bnCO^DOn 6 n6 i 3nhm6i. n8nb 0nb3g3noi, 013
hmftmh 333dpnicn 0u36ih336b Bypnb 0 ihi&nb bi 50 mi)p3 I53!p3&igm6nbi gi
imhoijp3bnb h353pnh36i, nbn6n nyngniB I53ip3&ip ^ 36 ^ 6 (jlmdmnm^nghjp) gi
bii3cnhn3 ^36ni6 (!3mn^nQmlnphjp) 0i536ih336 ig [264 ],
3mnyi(vn3nghx*' 0i536ih33&nb Bypnb h3gn0n 3cni6i6hg3&i 0 ioi J>ih30m3n
ihb3d3p j)36noi QmQihij^&nb 3nhm&3&b. 3hfnj)ihnm^36b, bmim3&b, 6yip8(536ih33-
6nb Ibmj)n3hcn bib3rn6ib g i pnj3036b j,iifi6nicn 3iîhi 33h3P36n, hm0p36biu ih
ij3cn U36^)hip3hn 3ii3mpn g i i0n^m0 J)i0 m3 hfn6 nb 330cnb333i3n ni3033&ni6 bi-
bnumiibpm 3hm^3b36nb gij33noi3bnb 5 ^ 3 8 3 . 6ypnb 330l533Cm^nb gij33noi3&ib-
o ii 6 3hcnig, 3 i5nhg36i nb3cnn bibn^mijbpm 3 hmu3b3 &nb n6^ 36bn3R b i, hm &m hnuii
q3m^mbn6oi3'bn g i b36cnJ3i. ^36nb 0m0i^36ibcni6 3hcnig; 3b bibnbmtfbpm 3hm-
(53b36n 0693636 R 33X 11 n6^36bn3m6ncn 0mj03g36ib.
Bypnb 0n6n0ip3hn 3m^36unipn, hm83pnu 5i6i3nhm63&b |jipi33pn bib3m6nb
ij^n3mbnb bibpîhb, b33(jnqjnj)3hni g i 8^36ihnb J>i3h|j3p36nbicn3nb J,igi86y33-
^n 86n3363pm6i ij3b. 6nigi5nb bmjjR36n g i 6iJO)3hn3&n 3bmpmg 0 i 3 n6 nBy3636
giymqjib gi 6mh3ip3h (jbm33p3mj03g36ib, hmg3bi(5 b36b^hi^nb 6ypnb 3rn^3&-
- 215 -
B’dÔb ff)03090
jjbhnQn 4.1 Sijhnb qjshpm&nam ^36ns6m6i (bb6shnb ^33001) gs bb6shnb mbdmb^hn 6633s 20®C
(Walter, 1931)
qjihgm & fjcnn mb9mb3hn aphem dnovi mbSmb^hn
>)3 & ;Ii6 m 6 i (% ) 86331 (d3s) ,$3&rts6md3 (% ) 6633s (33s)
100 0 93.0 9.8
99.5 0.67 92.0 11.2
99.0 1.35 91.0 12.6
98.5 2.03 90.0 14.1
98.0 2.72 80.0 30.1
97.5 3.41 70.0 48.1
97.0 4.10 60.0 68.7
96.0 5.50 50.0 93.3
95.0 6.91 0 CO
94.0 8.32
Polkllohydrlc plants Homoiohydric
61b. 4.1 0^363(13013 Bypnb hjgnSnb 33Pip3(jn3 Siam bd^Q301^3 5sgsb3(<sboi36 psi339nh3&noi
(Walter, 1967)
y n ip n o io c n jb n i, 3 n g h 3 -5 - -30 MPa ( 0 3 3 6 1 6 1 6 3 6 1 Q ssb p m y b n c n 80-95% li^ h n b

q jih g m & n c n ^ 6 n i6 m & s b ; nb. (jb h . 4 . 1). m &nb b m jjf^ b n “b h g i b 0693636 75-80 %
li^ h n b cgihp m dn cn n (i>3 6 n i 6 m 6 n b ghm b, o i^ Q jji ih b ^ b r n ft b bm ^^bn ( 0 15 .,
Xeromyces-ch b ib ^ m ft a b n ) , h m O cg b n u 60% J)3 6 n i6 m 6 n b Sn hm b^ feSn S o ^ d p n ic n
‘b h g i [ 183]. 3 i 0 m g n 0 3 h n b i j ^ h n ^ b n -40 MPa ^ 3 6 m 6 m b n b 3 n h m 6 3 5 6 n n 6 ih -
6363636 iiyi3fn6ib. 633hn cnj36n nBihR^^b qjm^mbn6cn3‘bnb 36111b, 0 i 9 n6 ii 5
y i , h m g ^ b n j c n ip ^ b n b B y p n b 3 fn ijo 6 |jn i0 n -3 MPa - ^ 3 6 3 3 1 1 1 9 j3 3 3 n o r i i h 3 1 5 3 8 1
[ 203 ] . 3mn])n(vn 3n g h x n m h 5i 6n,b 036n 63b3e36i 09h i 0 3i 6n ^ i ^ 363n ^ y i _
- 216 -
81)366601 7)J(A3 ^ 0b 603000
bi3b353n, ^h^Qn356 Jjhn3^m5i93b3n {Selaginella -b bib3mb3&n g i ‘bmfcnphcnn

53n0hi) gi 'bm&n^hoi qjihjpcn;}boni69 n (nb. ou>3n 6 .2 .4 .3 ). 3j>3hnb 9ihu3p3&n gi
oiQbpnb i(p9 m6 i(j36n Bih9 mig536b Smdmnm^ngh^Qn 3 i536ihnb 3mnj)npm!lnghx
bîgnib.
3fn9mm 3nehxn 9 u36 ih 33bn 3B3i 63 5yip 9 (j36ih 336nbij)i 6 5i6bb3!3g3bni6 3i-
i^mpnbnh^&xn 33(13936001. gngn (^B^hip^hn Sii^mpn ihnb SmSmnmlngh^pn 8(33-
bihQQiinb bi^hoim 6 n9 >6n. bypnb g^gnjjn^nb 9Q9oib333i9n1 9nbn 6ii03&m 6i 3hm-
^m 3pi'b9 i 9n yi3336bn h g3& ! 3bj)3rip3n ihb^SrjQn 6ypnb 9ihij,ncn g i i 9 n^m9
Bypnb h]gn9 n 5 ^ 3 8 0 1 3nhm636'b3 3cghm 6iX 0& >6 ^ n b g^m^ng^bjpn. }0 !3 3
ghmb, gngn 3ij,3mpnb ihbQbmbi ij33ncn36b bn09 hipnbig 9 n 3(j36ihnb 9 gj)higrn-
bib. i 9n^m3 3m3mnm3ngh3pn 0u36ih]36nb Bn6i3li3bn S^nghmg 093636 gi])i3-
3nh3&3pn ^36ni6 6nigij,cni6, i6 3]g 3n 3ig g ilj 36n! 636x 3ibnîôii 6 (6ib. 4 . 1).
0bmpng 3ib 933gQ5, hiy 81336ihnb bb33pnb ‘bQgiinhn ginqjihi imhoijpQbnb 83-
3iii336 3p n jj^njj^pnb Shncn g i ginbym ^hi 6b 3nhiunnb h353pnh3b! 6ij,33bnb bi-
33ip3bnai, 33bidp3&3pn <y>bgi Bypnb h3gn8nb n3 gm63'b3 336ihfi3636i, hmg3bi(j
,536ni6rn6nb bBhiqjn L53CnCDbnb 3 n3b3g i 3ig, 3hm^m3pi‘b3 i 9 jg 0 n3ig ij^n3hn
hRjbi. i 3i 6 5i3m n63ni 3hmg3i|)n3pfT>bnb bij,hd6m6n 3m 3il53di, hiSijj, oii3nb
3bhn3, bi93!p3& ! Snbî 3i536ih]b j,i633noiih36n6i bi5hd6mbpig gngn qjihcnmbnb
3 jm63 bb33pn.
4 .2 9l536^hnb 3X h3P flb 6 y C flb h3 9fia n
4.2.1 eyxn DXhoean
jn9n3hig b3 x n Byipn ihb3bpibb 5m6b^n^3(jn3hn bibnoi; Snghiûnxn Byipn

gij)i33nh3i)3pni nm63&oii6, 3 ib 3 n 5ibb6np m h5i6x 6i3hoi3boii6 g i i(p033b
3jh3gnb 3pvihib^h3jl53hi3n; e j 3m'bnl}jhn Byipn !3b3&b 3ii3npb; n6&jhb&xsnjhn
Byipn cni3nb 0 03^3 np^bb 6n3oin3h363bnb ^hi6 b3rohj)b ^jhQpnb 33gp3b9n gi
33h3g3mhnb363n; y tb iX '^ X ^1 6yipn 3 mdhimbb ^3h^pm3i6n 3i336ih33&nb fciaîh
bnb(i30i 3 n.
3nghi(|)i(jn3pn Byipn. cni3nbn gn3mp3hn cn3nb3bngi6 J,i3m3gn6ih3, Bypnb
3 fTiP3i3P36n 5hm3g36ni6 3hcnig i6 g iS jb ^ x 'bQgi^nP^bt^ 3qjhm gngn 'bmSnb
lb33mp3i3p3h qjmh3363n (b^h3j£)hnh3& xn 6yipn; 6ib. 4 .2 ). dpn3h g id ^b ^ x 0
nm63&n Bypnb i 3 3mp3j)3p36b 3 noi jqjhm 3 «Jnghmg 3j)i33nhg36ni6, hijj 3q)hm
gngni 03b^n gi 3 il5ihii nm6nb hign^bn. ^Sîpmg nm6nb (b3gi3nh‘b3 ihb^^pn
bypnb 3(T'P3i3PObn, gngn 9 pb^nb ihb3bm6nb 5i0rn, 0^nghmg g i oinotj3 nb
jSmdhimg ihni6 nn6cni6 g i i i 33nh3b x n- n6 n3 D 3nhm63bni 33 j 36 npn ijnpnbi
3mpnbijihng3bnb 3rop3j)x 3 & ',>b (b3gi3nh(b3. Bypnb 3mp353p3bn 6iBnpg3bni6
3mpih3p 5530336^3 (Snghmjbnpnb, iihbmjbnpnb g i i9n6m553qj3bn) gi J96 ni6
0hi3ip, 3hcn3i630i'b3 bi6pij)3bx pi 3 n&6 ngi6 j>ih3cn ig3npig 5igiigj,np3big
bO)h3Jii3hnh3b xn Bypnb 9h33&b. Snghiî^njpn Byipn 33ig&36b DjhQgnb Bypnb
bi3hcnm 3 ihij,nb Sbmpmg 5 -10%, 0138(3! 3 ib bibnumijbpm 36n3363pm6i ij3b.
3nphbly^n3pn Bypnb 336n9363pm gi^p3bi(3 ^n nB333b 3hmj)rri3pib 9 nb 3dn03
- 217 -
tfodoao
bOh3J»!iDhX 93hp3p336b.
3hmofTi3pb'b9 nb 3nghi(jil 5f' x n 6y0fib ^gng^bn 6i6nQn y>3o03h3039 ihnb 6030*1
9 (j36 bhnb 7jh3gnb 4390360, ago&hnpjbnb bn9 ^nghfn3nb 606393001, 0601336 6 y 30 b
1.5-15 MPa-nb ^0361503006 3nhm6363n. 66333b, hm00nou>u *63363 6y30o 03 ^hn-
I5nb (33h3gnb i393cn, 303‘bQnb imQmngn) bofôD^X DOD^D^D^- 36019363
03^1102)3031™ 66333.
933 mbno3hn 6y30O. 6 y3pn y33Q3<b3 3qjhm 0mdh33n3 33h3gnb n3 636003660,
hm9036 ni3 3bhxD&D6 bb63 h3&nb h3lb3h333h36 nb q^jyn bb. <3010103600 6y0ob
03 h35 nb 636333!™ 3b3onn cgmh0 ncn 3hnb 6:>h0mp536non. 6y0nb 3b cgmh03 i3 ^
3hnb Oi33nb3q303g b306 nb363gm0 o: nj,n 932)330011363003 53 bb6 nQ 633(101360136
(03 j 3hn, mh53630n 693336(1, 63(^390 6331101360 93 nm636n) mb0 mb3ho 6633001.
bb63 hob Fib9 mb3hn 66333 ( 71*) 336'i2)-3mcgnb j)36n 6 noi:
7i * = n . R . T = 2 . n T /273 [MPa] ( 4 . 1)
bb63hnb mb0mb3hn 6 6333 03O30»T'6b 36bmQ3j,3ho 0D9^Dh303hnb3 (T) 9 3 g>3bb-

6n0O 636o03i36nb h3PiQ36m6nb (n) 63036360136 3hoi39. 1 mbdmon . 2&'1 0 9 3-
303h bb63hb 25°C-nb 5nhfn63&6o 63363636363 mb0mb3ho 66333 2.48 M P a ( 3 6 3
9 8 .2 % 333hnb qj3hgri6oa>o 036o36m6nb 3 ^ 3 3 0 3 6 0 ^ . 6m0 3 *)X3h 633(101366, 90 90
6ro6nb 536m, 633dpn3cn nb3 9 3^ ^13363, hm6 mb8mb3hn 66330b 6363363630 0 3h
^n ^h gm b. mb0mb3h39 d0O3hn 0m032)3036n6 5m0fi03hn‘b3yonoi (635., 03 j*hn -
b3b303dQ3g) 93 36 3 ^ (5 3 6 0 6 32>3to3noi Snghmcnbnb 63033^16001, 6y0ob mb9 m-
b3hn 60nb 63633636300 b6h3qj30 93 06n0363Qm3639 01530363. bOô^COhn 332,3m-
0nb 0)m63 3hmom3Q3bo368n, 33h3gnb 633660 6y0nb mb6mb3h 663b3 93 3hmofT>-
Sgbbenb 6yc<ib 83(1356 Smhnb 0 ^nghm 3hoin3hoij)335nhn3. 30nli)m0, 3m o3^linDhn
mb0mb3ho 66333 33nd03&3 530m3ny36noi 3hm(jm303cb0 nb 6ycnb 03h3&nb
63633636039.
- 218 -
3 0 0 6 6 6 0 b 3 X6 3 ^ 01) 63 3 0 3 0
4.2.2 ai536ihnb DXhDeilb 6yCflb 3Hi!)D5lSnX?n
3hm1>)oi30:>‘b3nb bnmjndn^hn *jon3fr'&n*xri3nk gngn 06n03630fn6* shs

6yoob 3*h*5b, ihb03g Byonb oi3h0m9o6*0nt)3h dg5m0*h3mb*b. 33h3g0n 6ygnb
013(1301906*0033(10 0950185(130165 8300936* 039*hg3b bjogoui 6ycob jn9njh SoioD^"
I50*gb g* 5*6663*336* 5*3mn6*omb O3h0o6noi 301036^03(10 363(150*. Sycob
qjihgm&noin 301036(50*00 (T) ihnb 030*01606 nb (1*01936016*, (101030*115 b*^nhm*
n0nbioi3nb, hm0 60300 6yQob 5m^36(3n*on 530^3*60101 015306303*00 6y0fib 3mo3&-
Ijnscnb g o ^ b ] [227]. 6yonb qphprnftnotn 3moD6l5n*Cn 5*60106*036* 36Qh5nnb
3(10133030 (J id"1)- hfn03Qnu 66330b 3110133001*6 0389350 o,T,Cfn6ncn ihnb
95353801136x0: 1 J = 1 MPa.
6ycnb 8511550b 30933 3(1010 3*63363630(15 309(1*00(136*. hm3x o u 55301065^3-
6* qjihpm&ncnn 036o56m6nb %-9 n 56 MPa-0 n.
bb6*h3&0n 6y*on mbOmb^hig *hnb 6030^, 580,4018 0nbn 5*80160153^303-
03&nbioi3nb b*^nhm* 363h5nnb g*b*h33*. 6ycnb oib0oib3ho 301036(50*00 ( T n)
^qjhm 956*005, 3ngh3 01*306303500 Bycnb 3mo36un*cn, 50oooi0 n&n nd36b
3*hyoiqjnoi 36 n936 x f T'£|*k. 301001093(1 g* SnghmqjnQ^h b3g*3nh'b3 663c 6y*cb
ij3b 3ihymgnorin 3mo36yn*Qn (6yonb 8*0(103*03(10 301036150*00; Y T). 013 6y*cb
66330b J330 80135^(53301, 0nbn 015306303500 363(1505 05o x fT>&t>- * 9 ghmb 6yonb
6633nb 3011536(50*00 T p 9*9360010*. 6yonb 0*h*5ob (3.0. b*030*mb 03bhx3^nb*
g* li)h>6b3rihonh36nb 36*hnb) 33b*g*b 3&c*e 6yoob 93015330*' bobo33ob (33(1x11,
33h3gnb 6*600360, 5*h3 bb6*ho) 6yoob 301^36(50500 95(^365 oi*3nb3qj*Qn 690*16
301^36^0^06. 6ypob 0*h*5n 0ooi 303(101 6(50113*, h*(5 303(101 35(^0*300105 55801-
b*i3033o bobo30ob 301,336150*00.
4 .2.3 DXhoenb eycnb erân w h n 6n6^benhn6^
6yoob 5(1*01*656*110 301036(505006 0Jm63 *9500366 3 mhob 5hb3&m6b 3mo3&-

l5O2>0ai* 663*016*. 303i!»hxn g5d*6xoi6ob *6*001503(159, (1016300(5 6*h0mnJ063bi
bb3*g*bb3* X D i>^X n 3mo36(5O50nb 8^0163 *95nQ36b 9 mhnb, *j*|5 03ndo36* 30-
bi^bhmcn gid*63om6*lb3 - 3b** 33601306 gsdi&30oi2>:>.
301^36150^03606 ym33on bb3*m&i j36nb 85010 55015656(13606 6n6 *3nhm6*b.
33h xob 6yonb (1390606 030onb33358n 3b 3hoi(53bo 601(1(50309365 6yonb* 9 * bb3*
6030103(1363606 5*9*0*6001; 6yoob oh*6b3oihoo 0n0*hoi3on* n0 bnbo30nb*336,
hm03Qb*i5 303(101 9*6*00 301036(50500 yj3b. 013 gnqp'boobioqob 5(1*30015(10 6n6*-
*C0g35m6* *h *hb36m6b, 33h3gnb 8056001, 33(139366* g i 8ioi 55113801b 0mhnb
9yihg36i Oi3h0mgn6*0n33hn 601656601(10165. 353(180 6ygob oihoijgnb 05e>*0O 93030-
bnoo 9* 3o33hom630n 5^3601 (3*5-. ‘bp*3nb 6y5on, 56 6059550b ©0*83 6y*oo)
n6m336 6y*ob 33(13909*6 Q* *0noi *^330013636 bnboo8nb 6y0nb 301036(50*06.
dpo3h 3ihymgooio 301036(505906 0JO163 33h393b0o y i 6yb0n 5^30019*6 33h3g0o
80390636*.
33h3gob 6y0ob o^*6b3mhoob 3*6m6'boi0n3h3636n 33b 30933 03-19 6*3336380
nym J56m6noo. 6*b. 4.3 6*63363605 dnhncn^go 3hmi53b36n: 6m6*b6mhm6nb 63b-
- 219 -
p a ^ o b rtoaoao
ôqo 6ycob 3 mi533x n 396 m35h3m6ob5ai3nb (Bypnb bbQhoim Sm^Buaico ’f-jjhên)

6ih9mig2,36b 6y0ob mb3 mb]hn 3m|j)36uo50nb5 (Tjt) 0 ^> Djh^pnb 339 Cn^
g 5d5i> x rn^n^ 3m^35(3nipnb (Yp) 8mhob j>56bb3533656 [4 0 ]:
’ F 3Sh3en = ( - ) T K + ( + ) T P ( 4 .2 )
mbSmb^ho 3^1,336(50500 (T ^) ym330Oi3nb 35hymq}noi05, bmgm 66330b 3 m^Q6 -

150500 ( Y P) 83080365 nymb 959360010, 6300b (jriQn 56 083050159 - îhymgncnnu.
33h3g6o 5hb36m6b 551130133 g365gm6ob 859360, 4 .2 -b 3695 953-
Si^mb 3c i“b0 nbi 95 33^13906 33903606 85^03593)10 3m^36(jnipn (-)Y X. 333,536
036016333530, 85,51103503(10 3m^36^niQn, 0153060 bnd^nhnb 558m, S^ndp^bi 353-
C3^DC3ymcn (653m65j)0nbn5 331139360 1536,511503110 3533m0ob 5511393).
Bytyicn ^ -y jn j/ b x (iWh6 Dt^ 50&iSX) 9g5m 9ah3m 650n S h m ^ m ^ b ^ o a^ jb n a^ cD -
hbg o'bhggbi 95 3311390b i3gC3&‘63 QnQ 66335b 530015(1366. ^hfcrohob 83633(^6001
3jh3gnb 339300 33og3h365g 0630365. 59n^m8 i53h53bu36^x 8g&m85h3m653n
33h3gnb 33g0o6 33366335 53^33660)1366 33h3gob 63360b 8386013 d505b,
06333b 6y0ob 83601306 9363)13656. Bygocn 5533)13606 638016333580 -
0 95 = Tp. 33h3gob 603b 6y0ob 9^165)1553606 365ho (C -jjt^ - 5nph^>3Cr>33hn
365ho [4 0 ]) 5(j>633b gng 36o93630m&3&b 86mQmg 85806, hmg3b5i5 6y0nb Sm ^ B -
150590b 386083630m 15300036360b ghmb 33h3gob a m y x ^ ^ ^ 3 5 ^ 3 ^ ) d0O3h
0 ^ 9 36 5:
< W [83 M Pa- 1] (4 .3 )
3b oi3ob365 9581*1309363905 33h3gnb 33900b 305bÔ3hm65<b3. 050 ob5<bp3h365

3 m(530m6 noi59, x i 6(!>0]hm6nb 3 mgxnb (e) bsbnoi [4 0 ]:
a b
65b. 43 a,b îyiQn'bnhj&DC'n 3^ 39361^ 6ycnb Sf’ViaSyniQn 3n33f\jPi6nj)3h 995m93h3md33n (a);

3i i>3rnCn'bnhoi>xn 33)13936^ 6yoob 3m,i36|yi3cn 533^33 5398h3c qjmoimcSn (b). 6ypnb 663306
imûn^c0 (^p) 9unhp36i 939360010 96113363(^160936 6x ^ 9e3 es mbSmb^hn 3(’nlj36ttn30fi (T^)
(39363 b x 3'# im 3 » ^ e Dshymgnam (Hofler, 1920; Bans, 1968; Kyriakopoulos & Larcher. 1975;
Pospishilova, 1975)
- 220 -
9 0 3 6 6 6 0 b T jJCA O em jjÔ b 6 33090
e = A P / ( A V / V ) [MPa] (4 .4 )
bsgiu, AP 3633636b 153(1^(1300 6633nb (53000365b, hm030O(5 95353801136x05

35h]gnb 0 m(530m6 ob 035(19016001 (539003651x7156 (AV / V). 3Qsb^fr)hm6nb 0 mg30 ob
g5&5go 85rt9363pm63bn (5 MPa -693; 055., 6505bm356 0(5365113015 65^5 03010103600)
63 r'^ 33^D^)b, hm9 33h3gob 3 muxm&nb L53CnC3^nb^b, 0 obo 5390360 i 93 nC;>6 063-
036056 95 i0n^m0 ,53(15(11110 01569501561^16001 0^1(19365. ^30 1 33h393&b 033d0O5cn
633(10 Bypnb 0m0ih55365. (159556 b336nb5 g i cgmoi03&n 303(101 363805,
3ngh3 6i0ibm3i6n 0(j365(i336ob5, 05(^500 ^h^mhnb 3nhm63&9n 0icnn 3Qib^n3hm-
6nb 0 rri930 n 5^6336 0ijbn0iQ3h 06n03630m636b - 10-20 MPa ^3^0163(159
063563 gmoip369n) g5 30-50 MPa (05h5g063563 03010103690) [ 199 ].
Bygnb ^hj^nbib 0(50(^365 3i53fT'0nb 0fTH53prn6i g i 0i^ 30m6 b 33h3gob 633-
6nb 5m6(j36^hi(3ni. 50 ghmb 3jh3gob 539300 b30 3qjhm 653936 66335b 530
cri5h3&b 3hm^m305b^36'b3, bb6 i 9 ih 0oo(p6335 nb <bc’3h60n (^h^mhnb 630(113560
63(1,5000), hm00nb 830g355(j 33l>3gnb 53930b 5(j'5h 933dpni 939^09(11^13365. 013
33h3gn bnoibnb 55(1300100 n0ymqj3&5, 3hm,5m305b,5o b^n0g36i 33h3gnb 53930b
(bibi‘b(j'3hm 305tb0m0n'bo; 65b. 3i). 50 0gj,m0ih3m6i9n T p = 0 g5, 5j3g56 550 m0-
9065(13, 'i'-jjh^pn = b030 3010b 8(5365113360 By^gb 35I155336 33h3gob 55113 ^>3-
g i 3nhngi6 5mnoij036nb 5<bnoi. J,i0m09h50n 3jh3gnb 33903680 Bysgo 003906365
3hm^m305b^36ng56 . 33h3g3&n 0693636 ^36m6ib. b0303 Oinb 0(5365(133690 3gi'b-
amontn n9 n^m0 bg3&5, hm0 33h3g3&nb 3300360 ^53hb 5h 5,55113636. 50n^ma
3399360 0 n3y33&n56 3hmj)m30ib^ 36b, nbn6n 935,6001 930^160;)36056 95 50 ghmb
93od036b 0mbg3b 33h3g3&nb 9553(53^ (ijo^mhn'bobo). 013 3^,5(130503(10 550 m9-
hm6nbib 6nm0306hi636n ginh(j'3335, 33h3gnb mbOmbjhn bnb^30i 3050 000365
(6ib. 4 .36 ).
Jbr»3nc0n 33h3gnb 3390ob 66335 n'bhg3&5 56 0(5ohg365 93^016300 33h3g36nb
03333m6 ooi. 50n^m0 50 ghmb bi^nhro 55(130133 jbm3ngnb 05di630m6nb 550135-
0nb6n6365. 013 03’bm&30o 33(139360 33900b 66330b 03b5650nb5g 0mj03g3636,
,53(1^1536,5300 0 g5 m05h3m&5 000(^6335 3333 Bygob g5650O 930^330m6ob 3 nhm-
63690 (3533m0nb 3qjhm 35*55(15 0m(53pm6ob ghmb). 3b <3(1059 060836300135605
6i“bo 33g036nb 0 jm63 33h3g 36 nb5cn3nb, hm003 &O(5 50nb 550m ^h^mhnb b^5&n0-
3hm65b 6y0nb 0(50113 h5mg36m6nb 036m3noi 5(3*63336. Jjl^gnb 033hn3n 3390360b
930016333590 obo6o 933o6hm336ob 3hm(53b8o 0ymqj 3hm^m3o5b^b 33h 3633056;
50 ghmb 65h0moj06365 ^h^mhob 35hymqjooio 301,536(50500 g5, 93 b5&50 ob5g, jbm-
300360b g5d5630m65, hoi03005(5 93od03&i - l - - 2 MPa-b 0n5(p6omb (055., 3500ob
0301010360, Zygophyllum dumosum; [ 172 ]). 00 830016333500, 013 5hb36m6b
93bi659nbo mb0mb3hn 5(159036,50, 3hoi0563oinb 533(^001 5560553630 351^ 36 8 0 ,
hm930oi5(3 51)301 556bb3533630o 33900b bobj3 g5 305bÔ3hm65, Byoooi (J3g5g j%-
h36330ymq}ob 3ohm6369o 93 od03&5 0mbg3b Bygob 55g565(530365 65”bn 33gg36nb
9jm63 33h3936og56 (055., Bygob g5055hm33&3oo b33X3B^3hn qjmoi036n g5
P3hm36n) 3qjhm bj3033go36o56 3jh 3g 3&9 o (055., 03bhob3&3ho 3ih 36jo0 ob 35h3-
9360).
9(5365hnb 656oo36obioi3ob g505b5bo5oi36xn Bygob 3m^36(jn50nb (53503650m-
6nb 050^30936x365 B x n^> 0 m05h553&5,b 3 550mnb5^365 66335-0 m(3x rn2)n'J 9 ni6"
h50ocn (P / V-go55h505), hm0 gob 0ob3g3ooi5(j, 6 mo0 -05hnmôb 356m6ob 550V
350ob6o636noi ( P . V = 3m6bv556^5b), 3jh3gnb 6633nb 30365, 6y0nb 35h53ob 3o-
hm63&9n g5oy356365 ©muxf’i^nb 30365^3. 66335b 550 mb5^>3b 6y0ob 301^36(50500
'^ 33bD9 n' ® rnUXfT1‘->;>^ - 6y0nb g5hgm6noio 030i5330m6i (RWC, ob. 55h30i33
qjmn0305 4 .13 ). P/V-go 55h505lb3 03od03&5 0mod366mb 6y0ob mb3mb3ho 3m^36-
- 221 -
633080
Bib. 4.4a,b fcidroahmbnbib qjmcnmc3n 6y(yib 5 ^ 16^ 30 361 6ypnb gi33ih i53&3Qn
Jbm3nQngi6 (3nph36jn3i) ibnSncujn^h jbpynciQgO (^0^ 36^091). a qjmaipnb ^hnon 6yonai
iP 33h36nb qji’biSn (3ihub36i b^hiovi) g i 5i 3 r>3hm&nb g i6y3dngi6 3 gc'nb 933g3& (611133361
b3hnnn). 3ngh36jn3i 3qjhm d3£ig 11^3366361, 3 ^ 3 Jop1*136Jn3i; b 3ngh36Jn9ibi gi
JOiTih36jn3 i 9n 6ycnb qjihgmiinoin 333(j3X m&> 6ycnb îh^nb SjhnmgSn. 3i3n6, hmyi gmcnmon
iih{,i3b 6ycnb 50%-b, 3ngh36Jn3i 33fHji3b Sbmcmg 25% 6yiob. bmpm jcmh36jn3i - 33h (>0933
75% (6yonci j ,i j 3h36ign himg36m6nb) (Haberlandt, 1924; Schmidt & Kaiser, 1987)
61b. 4.5 8(536111301 6ycnb h3gn3 nb 5i69bi'b(p3h3cn 5ih i33(jh3&nb b ^ i^ jh n 3mg3on (Richter,
1978; Schulte & Hinckley, 1985; Richter & Kikuta, 1989)
I5fiicn 6ycncn 3nhm63&9n (0hxnb 6h03030 0m6ii)33cnnb 3jb^hi3m c^

linncn RWC = 1 960336300161093) pi Bycnb 3ml436^nx'n ^h^mhnb 630^3^6
6 ]honc9 n (nb Boh^ncn, bigiu 3 h]gn 6hq3n3n 69361). b h x OD^6 D^5D®bn;>'J>;> Qi
O^hftrnhnb 6 x ^ 3 ^ BQh^ncb 3 mhnb ih b36xn 9 h]pnb 0m6ii33cnnb qjmh3 nbi gi
Qibhn0m6nb 0 n b x 3ncn 9ondc36i Qi3igj)n6moi 30ib(j(n3hm6nb 0m gxn; 3 ^3 ^ n3-
j)X0ni6n 3jhx36nb 0^0163 Jbm3n0369n e 0ipi0ni, 3.0. 6ypnb 9360131^ 3hmu3b3n
- 222 -
813DG660 U f f a e m 603080
33h3gnb ioeCD^nb 5ibi630ip bi^nhmi 3ipi0n (j^h^mhxn 6633^. 83mh3 9bhn3,

6ypnb abmpmg 0unh30n gi6iiih536nb ghmb f p n6y3&b 3(336 xD&^b. P/V -gn >
5 hi3 npi6 i5h3ai33 33 ndp36 i 9^9506936 mb0mb3hn 33j,336nb 3hmi53b3&n: 33(13-
gnb B336nb mbSmbahig i^r^hn 6i6n036nb 3m3i^36nbib, ii3hj,mhnb 6 x fT’3^6n
B^h^nQn nbh36i RWC-b Qi6b0n 96 n3363 pm636 nbbt)36.
4.3 ai506^hnb 6ycnb hosnan
b33P30inb 6(536ihob SnBnb'bQgi 6i6o0n 3bi‘b(j>3h3&} S^ghb g i imhoij036ob

3hni33b9n 33h3 ^636 1j x i e iih6i3b Byigb, hm3 pnb Sihij/i 36gi 33fi3bmb 6nigi5n-
go6. d)hi6b3nhi(3ni, By0nb 836013} g i 3 nbn qj3b336ngi6 bi^hi6 b 3nhi-
I5nm rnh5i6m36i3g3 ihnb 3u3Bihnb Bygob h3gn3nb 36y33lijig 3n3gn6ih3 3hm(53-
b3&nb 3hcnm6pnm6i. 3i3h3n Bypnb mhcrrjpnb g^qjnu^n ihnb imhcnjQ36nb
3i 3 mdhi33&30n di0i, bm0m 6nigij)8n ih b3& xn Byipn - i 3 Shm^bnb 3^(13633-
0ymcgnb 335n33630m3i63bn 5 yihm. 6y0nb hggnSnb 8 3 6 ^ 3 6 3 6 } bg3&i Bygnb 639-
3n3n 6i3ignb bi03ip3&ncri, bmgm 3 nbn 6m6ib6mhm6i ihnb 8mdhi3n 6m6ib6mhm-
6i, i 3 bnlj)y3nb 5nhgi3nhn 36 n 336 x fT'6noi.
4.3.1 eyOflb 3D013nb36i
0(335ih336b 6y0nb 33cn3nb36i 333dpnioi 601300 ^giinhnoi, 3 ij,hi 8 dnhnoii-

9ip 3ioin Gypnoi 3lbh3633cymq}i bg3&i 6nigij,ngi6. ^ Sifj^ b 3l336ih33&3n i 3
q336j(jnib ^ 3 0 3 6 3 6 bûnionbnt^bxn mh5i6m36n - qj3b33bn. 30gi6o3b 8(536}-
(1336b 03363360 ih 5iiR6nicn gi iSn^mS 6yi0b ooi3ob3b36 3n6nb‘b3gi 6i6np36nb
33833^6001.
4.3.1.1 eycnb snhe^nhn doms^ cn^cDbnbi ycmhibDbfib

a 3 3 3 Dn6 ncn
oii0ftiqjnlj)36n 6yi0b nBri336 3i 3n0 ih30 ig (*i36ni6n b36b^hil>ingi6, i6 hm|5i

63n3i, 6 i 3 n gi 6yihm36n ^360663636 8ion <b3gi3nhb. bh]0n 6i33^36nb 3nhm-
63680 bmy^bnb b b g x 11. 9303b363hn 0nj363&n g i ^mhcgob bi3b3&n ^(10133636
15-33(1 ^jaghm 83^ Byipb, 3ngh3 obo6n 33n(5i336 5 i 3 m83 hi0 8gj,m8ih3m6i3o
[240 ], 6yi0n 83061*1336} bBhiqjig. oii03bn ngpn6oi36i b x hi3g 36n33 630inb fci6-
3 i 30m6 i 3 n, bmgm giibpmabnoi 6ib33ih biioi9 n 3nn(j»633i bhxn 6^33h36i. 6 ^ 3 -
(10366, 33gi603b bmim3&b, bmin36nb 3nqj36b, bmj,n3hcn Byi03i536ih3bi g i 0n-
foBb Bypnb 336m3i 333d0nioi Bm^nm 3i3hnpi6. i 3 j/bnoi 336m3n0n 6yi0n ym330-
0130b 303(101 3unh3i, 3ngh3 333i0m gi^36ni636nb ghmb g i 6y0nb 338(5330m6nb
- 223 -
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2,ih30mboii6 5iai>6i6li36i, hmj,mhi3 6]bn, 0n0gn6ih3m6b 0hi3i0o gjnob

&i63 i 3pm&i3 n.
^h^0fii6n 0(336ah336nb b 3g i 3nho giqjih3oni imhotjp36nb biBnBiiCpQg^m 53-
(jnjiX^b 3 hncn. i ^ n ji X i 00333 3°}3iw hig ^Scnb b^pb Byonb 93pB33ib gi^36ni-
63611b ghmb, hmj,mh(j 3 nb imho)jp36ib 9u36ihob bb33cob ^giSohogiB. 013150
bb33Qr*b tb3gi3nhogi6 6ypob 836^13^ bi3hoimg 3o3go6ih3m6b, nb bg3&i SngiOirn-
g3&nb (33ng3h0nbnb ByicasQ^hn ig50p36n) bi 33 i 03 &ooi. 330030,53(1
Bromeliaceae-b 6 ih3 m0 ig 2,36036b ^R bn io i Qioioôb 6 3 ^ 9 0 6093630 ^60 338-
6m3n J3h(j036n. 0U3hi3n 6yio 3 b36ih336o <3frioic36ngi6 Bycnbi gi 0n63hiQ3hn
6n3om3h3636nb 93biBm3ig ny363636 6i2>336nb bib 383i53ono iSihi^b.
4.3.1.2 eycnb g o e rm 6rbe:)&ne:)6
6y^cn 6nsgsj,Bn
Byipn 6oigi&8n b3g36i 6soDjo6nb biSîc 36001 e* M T ^ b fth ^ n b 6 yoD^ 9 -
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83 g3i3n33^hn. 6nigi2>8n 83(3163300 Byonb 6i6non hRp6i qjmh3&0n (djbm jnQn
BysQn). 6nigi5nb ,503^3 g i qjmh36ob b m O i^ i gi 3 mj)ng363cn 83(p63Xn Bypnb
nb 6i6non, hi(j 3 mgnb 8361^00 6yio'b3 gi nb 6iBnen, hiy 0 ^ 2 ,3 6 } Snigifjnb
(ph9 i 03363680 (6ib. 4 .6 ). 10 |i3-3g3 bmSnb <3^1(1360 0,5,50(539 6yieb
ii3noih3on di03&ob 33933m6ocn; 3cghm gngn bm0ob <3^ 360 (60 jxd-V) 03c>n)
3 ib b6hi<3ig ^ (13 6 3 6 .
6nigi&ob 336m^non Byoooi 2>i33h36nb 3g2,m0ih3m6ib 36mg36i 6nigidnb b i3 3 -
03 (J36^)33igri6i gi 2,i 0 fnnbi&)6 i &h H20 himg36m6ncn 100 9p 0 m(53pm6 ob 0 jm6 3
6nigij,8n. 3oBhmqjmh36ni6n, impmng36ooii g i mh2p63po 6i3hoi3&ncn Sgngihn
6nigij,n 3ghm 0 ^^ Byipb n6ibi3b, 3ngh3 0bb3np<3mh36ni6n. 6oigi2,ob b i33C 3
ii36(J33igm6i 8i£)pm&b 838932,0 hn^nb 3nb3g3noi: j3n8i, oinb6ihn, ombi, îmbn.
bi65hdon3n B3O0nb 33hnmg36 nbi gi ^ ib iq jb jp ^ oim3pnb g6m6nb 830936 ByiQn
6hm3g36i 6oigi£,nb oi3nb: 3qjhm ig 300ig
635 36i03Jm 33(1001936^
<b3g i 3ohx^ di03&ob 038331*16001 33&m^non Byipn np3j3&i 6nigi{,nb ynpmng3-
6^3; *,i3opi(i3po di03&nb 9mJ03g36nb 83932^9 R3hg3&i Bnigi^nb <3mh363n g i
83odo3&i oymb mb0rnb3hig 836môoo nm63&nb 3n3h (5i6bi530ih36nai 3ihnQni6
6nigi5dn). ob333 hm5(Tih(3 0(336^380 ih b ]6 x Byipb, ihy Boigi^nb 6yipb 833d-
OOi oni3ob3<30ig 0mdhirn6i. Bnigi^nb 303<53b ^n3368n nm636ob 0n]h Bypnb mb-
0mb3hn gij>i336i gi qjmh36nb Bypnb 3nghmbî)n|)3hn B633i (Tp) n0g36ig 6(^n-
h3i, hm0 33ndp36i 0ioin 36X 3^ 3Cy(T'°J;>- 6nigi^boi3nb 06n936x f T'3^6n^ 6yQnb
ds^hntSJCn înQ shjQ n) 3m^36(3nion T t - 363(15^, hm0onoii(3 836m^nQn
Byipn ii3936i ‘bpgiinhx^ dip3&nb 0n3h. 0il5hn(j3pn 3m^36(jnipn 5i6bi530ih36ncn
gngni 63hopgnih36ni6 6oigij)8o. 0il5hn(j3pn 3m^36ynipnb 330i96363pn ^>3 n-
0ih3cn 3ri(!)36(jnicn iQHiB3h36i 838932,0 <3 fnh03pnoi [72 ]:
- 224 -
9096660 U BTdc?Ob 603090
' X x" ' k
6sb. 4.6 Byonb 6p5maih]m6i 6ycnai jp j^ h jd x

(3ih(jb36i 6i6ncn) gi 5i 6m03hsc (0sh3336i
6s6nen) 6nigi53n (Lerch, 1991)
= - 4a/d = - 290/d [J i ^ 1] (4.5)
bigiy, 0 ihnb Bycnb VpiSnh^cn gidi&3cmbi; d - qjmh^nb gni^^hn (|J.3 ).

Bycnb 33&m^3i dcn]h 0iO3cm6b, hmg3biu 6nigi2>n 5i0m9h3i>i. gngn tbm3 nb
cgmh^dn BycnbjpB 05036^ g i 0bm0mg 0.2 118-^3 S i^ h i bnSnb qjmh3&9n hfi3&i
Jp3n0ih30n Byion. ^ 3 0 1 3nhm&3d9n ;)3n3ni6 6nigi53n Bycnb 3mo35 LSn^Cn bBhi-
qjig 3sh3bg36i, bm0m cnnb6 ih3 n 3b 3hmu3bn 3qjhm cni6i0n0933h30>g g^> 630^
9 n3gn6ih3m6 b (6ib. 4 .7 ).
6ypnb 836013^ qj3b33&nb 3n3h

8^36^3 6nigi5)ngi6 njs8g3 n6m3b Byicb, bi 6 i 3 qj3b336nb Bycnb Um^Bunicn
ih QinB33b qobgcni 9 hnb 6ni95j,3n ihb3&30 Bycnb 3m^36i5ni>c'b3 gi&0i. Bycnb
936^13^ 3 ncn 3q)h(Ti n6o36bn3hni, hitf 3qghm gngni q}3b30ii bnbo30nb 9306m3n
<b39iSnhn g i g)3b33dnb 3n3h 6 0 ^ 5 0 9 ^ 6 Bycnb 936m3nb 36ihn. 3b 3ncnih3&i
Xpn63h36i 3339350 qjmh93 cnai:
W abs " A ( Y 6 n ie i& nb - ’P q ^ n b ) ^ ( 4 -6 )
qj3b3<jii bnb^33nb 3n3h ghmnb 3hoi33c3n 93Bm3n0n Byonb himg36m6i (Wabs)

ihnb gi033b3ni63i)3cn bn3hynb (qj3b33&nb ijo nD^n ^giSnhn) 3n9mu3cnb ^ g iS n -
hnbi (A) gi q)3b33&bi g i 6nigij>b 9 mhnb 3m^36uni0nb bb3im&nb 3hm3mhunxn;
n^n 6nigi£,9 n Sypnb fr^ h jb n b 6n6a(pdQ3&mdnbs gi 6nigig>ngi6 q jjb jjb d n
Bycnb dj(p6jjnb 6n6^(pdQj^,mdnb (djpB j^nb 6n6^(pdQj(,mb^) 3 ])33hm3mhijnxrc>-
33<bD6X5nrib 33hnmg9n qj3b3ou> bnb^30nb 9 md3X 02)X n (3hmjbn0 i 03hn) 6i6non
n3ncru>h36b jjmhSnb qj36ib, hnb &i0mu qj3b33&nb gnq}3<bn3hn 0ibibniOi36c3i>n
- 225 -
b60)67)(f,n
(ttb. 4.7 3hoin3hcnei9mi|np36xDbi 5nigij)0n

6ycnb 336u33cm6ibi gi 6niQ;>2>nb 6y0nb
Sm^jEiunicb 3mhnb (Kramer, 1949; Laatsch.
1954; Slavikova, 1965)
0(33936}. ibiQn <33633606 039903(1 bhgnb b ih j'b j ;>Q6nCn ^36 0336301} 6n6lj30nb
6n6bi(p0g3j)m6bb. i 9 n^m9 6i^nhoii <336301} bnb^Snb 63013063606 ;)6}hn 5i6bn-
O30 nJ63b ghmnb 9o6}9nj)}9n.
03363360, dohnoiigig, 03001^3636 6yonb }ihypiqjncn 3m^36unacb, hm930 n(3
MPa-nb hi9g36o03 93^01396 3^019936^ hi(5 6^3636(101 6 ij>9 iho6 oi 60^9^506 363-
(i)3bn ^n336ri9}6 93601^00 Byonb 3gng3bn 6iBnonb 93biBm3ig. 36 jp6ri6'bm-
0n3h3&i jjih&ig fii6b 6ib. 4 .7 -gi6 : 3n^36(jfi}0nb 0boi0oig -0.2 MPa-noi gi^CD-
bnbib 03363366 333d0nioi j 3n96 iho 6nigi£,ngi6 63601^00 6yonb 9 ihig,o6 2 /3 -nb
9361113}; cnnb6}hn, hm03Qn(3 63ho0cgmho}6n 6}6o0ij)36nb 63333016001 3<gh(Ti don3h
}j)i33&b By}0b, 5ib(330b oi}3obo 93601^000 Byonb 0 ihi&ob 6}b33}hb 3333 - 0 .6
MPa-nb Soi^BôiOob 6 ih 0 oij06 obib. <boij,n3hoi 6u36}h3b 933d0n} &ihj)33x qgih-
503680 }j^03hig giBnoib oi}3nbn Byonb Sm^B^nion, hi(3 0 }b b}33>036}b ^ 9 3 3 6
939}h36noi 83^0 By}0n 93n6oi3mb 6n}g}509i6. In^hmajn^jbb (03g0O3}g bm^nm
j>}h39oib 0oiy3}h3on 01536^3360) 933d0oioi 9}Bnoi6 oi}3nbn Byonb Sm^buniQn
-1 MPa-093, 330n93hn h32>ooi636ob i 3C0)3h x 0(536ih336b — 1-9}6 -2 MPa-893, bm-
O01 03'boiqjn^36b - -4 MPa-093. 09 hiCn 3s6nli)} (|,36nb, 3.n. bn09 h}0 nb 0 my3;>lixn
6(536^3360 Ijbjhm qjn^jbn) i300iih3636 -6 MPa-nb 03bi6i0ob 3oil536un}ob g i ^rv
933 Xjhoi 6}i036biu. b3-8(536^33630 3b 0oi6i(530n 03hy3m6b -2-g}6 -4 MPa-993.
Byonb 336013} q)3b336nb ibpoib 0 9 3 6 ^ 3 6nigij>nb bn3h(5ngi6 nB333b 6nb
J,}g0oiig5no36ib boigiôb 3q)hoi (i,36n}6n 033636ng}6. 6yonb fcigiig^nc^b} 0n6-
gnB}h3oi6b J)i3n0 }h30 ig, 630} gi 9boi0oig ibooi 0i6dno36'b3 (hi0g36n03 88-9^6
1 b8-0g3). qjoih36nb bnb(j38ngi6 Byonb jjihj^nbib Gygnb g^^hjbnb 6n6s±(pd-
gj&mbx 5i6bii3aih3&noi oinb6 ih9 n, 0i33oihig ribhg^&i. gngn <301113606 6^0163
6oi9ij>6o, 3333 gi6ion gidibxoibnb 3ohm&369o, Byonb »>i3o0 ih xn b ob ^^n b
"diO)36nB 39i0 6yg36i g i 5îh36nb Shoi^bn nh(ji333i.
Bnigij,9 n 6yi0n 93ndo3&i o9ym<336mg36 oihoijgnb bibnoii^. (^63 6nigi5nb
- 226 -
90366 (n0l> PaÔU 633090
bpgi 03363611 bBhiqjig 15039365 g i By^pn jqjhm o*6o(Vi (j+105 0336360956 mhcnjpnb
bibocn 50oignb q)3b336nb bn3h(53093 g i yn6g36boh936i ( cnjhdmi,m6p j 6b^jnj).
83 hi0 h35nm6369o, 50 3hm(53bnbi g i 6 i 9 nb 03933016001 6nigij,ob bags <3363630
069360 Byipo 2,hm3g36i, hiu bi^dihnbni i j dnVbihgn 8(336ih336nbicn3ob. bn08 h>
9380 69365 55h 30133 Byonb 33hooi5o;)hn 5ig5ig5O036i qj3b336nb bob^30ob 805-
6001. 0i03 836013000 Byi^n 8036^9365 ih i 0bm0fng 9(j36ihnb 8 0 6 0 ^ 3 9 5 6iB-
0936b, ihi839 (b 39 i 3r»hx qj3b33&bi(5. hm93bi|3 6059550b 13395 0336360b Bycnb
301,536(30^0 3qjhm 9565905, 3ogh3 q)3b33&nbi, 595090 5j3b Byonb 550mypiq)5b
<331)3360956 60595580 (65b. 4 .8 ). bagiin h xn q)3b336ob 0)0163 b5b3oi636b, hm8-
9360(5 (ph85 qj3b336ob 0)0163 9|536ih336cni6 3hou>g n"bhg36ni6, ibôm
Bycnb g(?j-(?sdjhn sjsfijn b 83833016001, 333d(V>50i 33301 55950036016 53503360.
55h38mb oi53ob363h363&ogi6 5580189065(13 (6059550b b^lrjtejhi, 0nbn bnph-
83), 8(536^38 830(^0365 556030015(111^ (b 3g 53ohx,n. C+i0i9 g50<Ti03 nCn 56 biho»3-
Onb36ho q33b3aii bobo30i (6ib. 4 .9 ). 0(536ihob 0336^5001650^ oi53ob363h3636bi
95 qj3b3ci5 bobo30nb ^n3b 8mhob ihb3&oi6b 3 ^nghoi 3hoi03hoij)533ohn. n6^ 6bn-
jhsg 9*<33b3n$ 6jb jc n (3 * 6 - 3598360; 633hn, 0 i 5 h50 ihi y330i. 65956013560 0(53-
6ih3) 0(5365(13360 qj3b336nb bj^n^h lb3g 53nhb lbhgn56 63b3b36nb 5i63noiih36ob
b5hj'b3; jjbtfjB bn jhsg psajjbjnsBj& jcn 0(536ih336n 0311015(13636 035(101019 556,50V
0>3nC <33^3015 bnbo38ib (855., 00^9393630^ 356005,5360b 633hoi bib3m6i; 355-
^3b360), 5695 - 6nigi53n ph0i0 8085350 <33t>336b (055., 5h36onb 6yo36nb
550my3636i‘b3 b33(5Oi0n{boh363Qo 39i66oib5 95 ioe0^ 8(5365(13360).
6yonb d3660b 3hrTi(53b3n qj3b336n 03980359 0mdhim636. 6059550b 0156-
Night Day
Bib. 4.8 Artemisia tridentata-Sn Bypnb {(iQ}6}U3C'3&:> <30l>33bngi5 anESnbtrxp 6:>6np9n g i dnbn
imhaiJcjbi ppjfniaob J>b60:>3em&i3n (Richards & Caldwell, 1987; Caldwell, 1988). 3b&>3bn UJqSqJki
bbjigibbsb bibniynijbom qjmhSnb 3|x)6ih]3bnb!><n3nb nb. Dawson (1993)
- 227 -
fnoaoao
50
100
Pituranthos - 150
- 200
250
Lolium
300
350
Rock at 6 m | r500
6ib. 4.9 n6^36bn]hn (Lolium multiflorum) gs jjb^^fjbn^hn (Piituranthos tortuosus, Zygophyllurr,

album, Ferocactus wislizemi) qgbjoui bnb£)9nb d^ft^cnot^bn (Cannon (jn^. Kutschera (1960), W alter
(1960 g i Kausch (1959, 1968) 3nb3e3noi)
- 228 -
8006660 b e a ^ o b ôaooo
Qiai}6m6noin 5i9m9hm6nbib <336300 bnbÔnb 6i6n0O igigrigifc i3e o 6 i, bi9i-

fc^hmg W>3i *95003630 - hs9g36n03 9 3 ^ 0 0 1 o‘bhg3&i g i n6£)6bn3hig n ^ m ^
6 i. 3b 0 13 ^ 3 6 * j,i6 b i 330^36001 gi9ibibfiioi3630n i 93hi0n 3i6n(îlj)36nb 8^36}-
h33&nbiOi3nb. < 3^ 30 1* bnb(i)30nb 6ibD®y33ibDCn 'bhgi ihnb 9(i36ihnb 6ygoo> 0m-
9ihij,36nb 396n 036xm 3i63bn 3nhm6i. 9]>3hn3n, g i li,35ni53630n 6nigij>36n b3Qb
300ni6 (bmcm ^ 3 0 6oigi536o } 6 x 3 6 3 6 ) <336301} bnb(j30nb 'bhgibi g i &}63fi-
oiih3&ib. ib30> 3i6n(ji^369n qj3b33&nb 'bhgob 03<33hb36i ihnb 9(536ihnb Bygnoi
ihibij)3ihnbn 3<bh36330ym«3ob dnhnoiign Srfb^fbn.
}6bmh6(jnnb 3nhm&36nb g}big2>36ig 0bmpmg 6nigi5nb 3ih i0 3(jh3&nb 150196}
ihibij)0ihnbni. bi^nhmi *2^30133 d y jS sh jd n Bygnb dg^m dihjm bnb (5mg6i. i0 0n-
0ihoi36noi 06n03630m 3i6ni qjmoigob 6ygob 3m^36(5ni0nb 33b6i 30i 6m6ib6mhm-
6nb 3nhm&360n (0 i&., p i 3 nb g i0033obib, i 6 b x m 363h ig 030i5nh3&3Qn (3hi6b-
Snhi^nnb ghmb), h iy giibom36noi 0 33b i6 i0 36 i hn'bmbq^hmb 5m^36(5ni0b.
4.3.2 9l3362)h3 6nêi6bi » >!>anb°33hnb aPlhflb eycnb

3 Hi!)3 6 l5niCnb 6 hiQn 3 6 C>nb 3 nhn 6 3 6 3 fl (b flb W a i: 6 n ie ^ > n -a i 536 ih 3 -
ô>anb<33 h n )
n0936ig, hi9g35igii5 9l536ihQ }hb3&m&b 3i3h3n 6y0nb mhoijgnb 6533nb g3qjn-

Un^nb (3.0. 3qjhm gibipn Sm û n ign b) 3nhm6369n, 9ib9n 6ih0m nj3636i 6y0nb
djgdnjn 6i»|ign. 3ml535(5ni0nb y330ib 3 ^qjhm 0dicjhn bb3im 6i 6ih0mnj0636i
cgmcn036nb <b3gi3nhbi g i 09hi0 3i3hb 9mhnb. i 9 ghmb &igib30nb > £ 3 X ^ 3 6n6n
6n6ii(p3g3j)m6i ij3<7i 3mdmnm5nghjc 0(536ih33&b (6ib. 4.10). j,igib30nb 3ip i0n
6n6ii[p0g35m6i 6ih0mnj06Q6i n0n(j|m9, hm0 6ygnb imhoij03&ib (2.45 3J."1 H2O
20 ®C) gi 0nb 9 3 (^ 6 3 3 ^ Jbm3n03&9 n b^nhg3&i 363h5nnb gngn gi6ibih3n.
Byonb fc i^ l^ n b g ib ib ib n io ^ g ig ihb3&m6b nb3oin33 3i 6 m6‘bm9 n3h36 i, hm-
&mhni5}} m0nb 3i6m6n X3ib^m6nb bijpgob 9 30 0 *333^ 0 . i 9 ol!)m9 3^hni6n 0^63-
6mgi i6i0m&n3hig 6 ih9 mg)3X 6n6i 6nigi£>n - 9 u36 ih3 - 3i 3hn-nb bnb^39 i Rih-
01300 Bh3gnb bibnoi. 6nigi&n - 6(536^3 - HiQhn bnbli,39i ihob yn6»yi6330n gi
0 ib9 n ihb36xf> 3m^36(5ni036nb 5 ( 1 ^ 3 6 ^ 6ih9mig536b 0(536^390 6ycnb (i>hi6b-
5mh^nb 0i0m dhi336x diQib. Bygnb ^m^Bynion ( T z) 0i536ihnb {>ihj>33X *©■
j,n09n 0noi 3qjhm îhymqjnoini, hi(5 3qjhm gi6i0Oi Bnigi^nb 6ypnb 3m(335(5ni0n,
gngni fch^n^^nnb d ig i ( T g), gngni bb3igibb3i &igib30nbi 013 5igi(ji6nb 6n-
6ii(n0g3j)m6i (rj) 6nigi&bi gi 0mi530x 63hljn0b (z) 9 mhnb g i dQ03hoi Bygnb
6iiign 9 i536ih39 n ((5^ 3330° 6i3ig3&nb 3 i 0 n S j j ) [ 197 ].
* * = ^6<i}ei5n + + ^ bnigi^n Jl • rl ^
}9 qjmh030ngi6 5i0m0gn6ih3, 3m|i)36yoi036ob gngn bb3im6i 03bid03630ni

9bm0mg Bygnb d0n3ho 6ijpgnb 3nhm63&9n, 3_n. jjhi6b!3nhii5nnb gngn n6^36bn3m-
6nb ghmb (6ib. 4.11 g i 6ib. 4 .12 ).
- 229 -
F a ^ n u rtoiioan
— U -7 < n
Dry air \|fa » - 100MPa
Va= ap----- h / - 1-°
(48% RH) -100

^ t ^
9
^stem
M b -
Humid air -10 MPa
(93% RH)
Dry soil \)»0 = -2.5 MPafrf r -at r

1soil | 'root
Vo=0 ]j< = h
Humid soil l|/„ - 0 MPa Vroor-0-6 MPa êoil■ I \
- 0.6
6ib. 4.10 6ycnb !3nvj35unicnb j,h;>pn36^11 91 6ypnb ,jh i6b 3mh,jnh;)&nb (5nigijn-0(}36ih3-3^9fnb<33hm )
h3‘bnboo61j x ,T|6:> (Cowan, 1965; Boyer, 1974; Kreeb, 1974a; Schulze, 1986; Nobel, 1991a)
Bypob 5^35 3(536511300

0(536ih30n 6yi0O 5i055Q6n0g36b 301,136150503606 51159036^06 806393001 33113-
90956 ( s b o m ^hiG bH m h^n) g i ^6003606 558^5110 6n6d30n6 833330161101
( d m h jjQ n ijh^ 6 bl3m h^n).
69006 5*0*6^153036* j6m3no363n 80890651130)66 3311300956 33h3050g3 Snghm-
b^bôôhn 51^036,106 55663ho3 gi 33h3go6 2,3903680 - Si^trpjob 3ho6yn 3 nb
( s Z m lp s b d j h n b$gs6su3Qjbs) 83656580659. 8365658060 069035^01113606 558 0 ^ 3-
636001 33 bidc36x n^ Bypob 0ih3h3^nbi g i 5595595003606 6ofijiho6 959536^
(6ib. 4 .13 ). jphjob 3ihQ6 ;)o0 ogi6 6 y50O 5595906 36901931100680. qj3b3nb fohjn
ibh30D&b 6145065155306 0336^(5056. n6 6583503656 ^ 0 3 3 6 8(536^36 5iioi56i6hmb
6y0ob 00160)93606 b568mj)03 (53503659016360. o6gmg3h0nb3o iDgoob 55663(1030
i3m305b^3ho ^hi6 b3rihôb 5g5O036fl 80165(53030166 3nghngm6 x fi56ihoi36on:>6
(iib 3ihob lbm036o), i6 J>390ob 556333630 65600360156. 50 5'boOT 69365 6yQob
6ij>igob "ii6i0n‘bnh36i" 36901931180606 6y50558^5h >95003660.
0336306 (536(|)hi03h (3o0o6gh0n 6y50O 803906365 dm hjjcn i)hs6b 5mhtfnb bnb-
^3806 03333016001. (536,111503110 1500069(106 3ih36jo030o ^ 3 9 3 6 0 65110015953636
<336306 663306 65(1801801606 59500366. 06 6ih0moJ0636i 0i8n6, h01(55 6yi0o (om-
6360156 3hoi5g) 03061*13365 ^3(1^03606 6nb^30i9o. £)h ^03606 606,1305 0635 0m6-
ym6o0o, hm8 833d0mb 836013000 6y0ob 66(150359 806019365 8i336ihob y330i 65-
6 o00 o. ^)h^036nb 550îh 606(130500 6 y50n dnhooiig5g 55950(^56365 hm5mhi5 9 >-
bo3hn 653590 (.3 s b t )jc s h jQ n Gspgri). 03010103680 jbo030nb 800360 0,101,136056 jq j-
hm 631100 5566(101363659, bm0m 8icnn (lhij3O03OO 601001360956 6yi0o 8306013365
95652^3311(136300 35(136^0606 0n3h, 6509565(5 5595065(530366 83^10103000b 33113-
93680.
6y0o6 6535906 6nRj5h3

6y0o6 h5oi036oi65, h018300(3 6igi505O0g36i 558^5(1 606^38580 ghoiob 3(1013-
300n, 05801109363005 8n6 06301 633(50q303h oi3ob3&36'b31 hoi50iho(355 550 l>)5 hn
- 230 -
30366 ff.ni> p a^n u rooaoan
Water potential [MPa]
6ib . 4.11 cnid6ijm3n cgncnQnb 6ycnb (lF]eaj-) g i pjhmb 6ycnb (’P stcm)
33h^,ny>03hn jhignQb^n. hmiy> ^hi6b3nhiijnnb n6^36bn3mdi aîem, e^hmb 5ib63hn3 diphg^bi
dpn^hn 3nghmgn6i9nj3hn jh ig n ^ n ; hnî gnoncabn oni3bg3&i SihyW n g i imhcnjcjfei
3nphmpn6i8ni>xn 6hign]6jnb 6^336363(30 njis^&b bha^^c™ Smtnxyiib 6nbgij,nb (Tîj) gi
ajmcnpob ('P]eaf) 3mhnb (Begg & Turner, 1970)
Rain Sunshine Night

12h 5h
MPa -2.5
( - 1 .2 ) -1.2
(-0.9) -0.9
-0.38
(-0.B)
(-0-7)
Juniperus scopulorum Encelia farinosa Hiiaria rigida Agave deserti
5ib. 4.12 bb3ig ib b 3i bibfujmubcm qjmhSnb 9u35ihnb 6yonb 3mlj 36uniCn (MPa) (Wiebe et al., 1970;
Nobel & Iordan, 1983)
- 231 -
6ab. 4 .13 6y(vib 5i 9i i 95n036i. Soleirolia soleirolii-nb qjmoipnb j 33(?i 33np3h9nb3 n Bypnb 5 19 119 5-
(10361 (0ihub36i b3hicnn); 6ycnb ,ihi6b3mhljnh36i cnhip3bu>6ljni ip6nojpmh>4j j339i 33n93h9obnb
33h3936nb i 6i5nj)On63h i3903^9n. qjC3mh3bu36uni 3633636b 3jh 3gnb i 3PC3&9n 6ycnb iSm^cib^n^^h
^lebpm iujnib (3ih3336i b3hiain) (Strugger, 1938)
bnb^Snb qjihcnm&n g i Sypnb j^gngjjnQQ&nbipSn 6n6ii(Ti9g35Pi6i, S^^Bihnb qjn-

<bnrnQfTij)ri!)hn (615., 61533606 &ibb6n0m6i) gi j,bh]8m 3nhm6 ] 6 n.
Bycnb 6ijpgn (JH20 ) 9 ncn jcghm gngni, hi|j 3qjhm gngoi fci^h ô b 'b^giSnhn
(A ) g i Biîgnb bnRjih^, hi^ ftiQmnb^bi cgmhSxnoi:
jH20 = v . A [ i 6 69 ' 1 ] (4 .8 )
qjmoiQnb 56 pjhrnb j>i9 îho 'bjgiinhn ihnb jbnco9 nb y330i 5i9îhn 303636-
ijnb 5i6n3n ^cn n b ophcnn&3&nb 5i9n. 6n63ri3i6o g i q)mcnpm3i6n b] 9(3361(133-
6nb 'bhm3&3n gi b j x ^ ,536 9 0 , hm003&bi(3 ij3cn j,i666330n gi 9^nh3qjmhni6n
100 9-133 0jjnh3 (9 njjhmq3mh36 n) 03hji6n, 6yi0b i^h^bb b jx n 9 i 6 ^ni, bm0m
9 ]hj 6 nb 33i53p9n 6yi0n ipih 9o3go636i. ^hm3nj)30 n b ] 9^336^33606 gngqjmh^-
6ni6 (dsjfhmqjmhjbn) 93 hji 69 ni3 nj,rt33 b n ^ ^ n i i 93 j9 6 n0 n. brnQn^h bih^yQoSn,
nqj6nb, i5h3 ijjiynnb, 01300b, 6i60nbi g i 93bnb hg)m0nb363hn qj(Tih36nb (un3Cm-
ggmhjdn) 9jm63 93hji59n ijmîi nb3cnn 00n3hn (15(^0360, hm903&bii3 933d0nioi
5y0nb bhxymcgnpig j,iîh36i (Bib. 4 .14 ). i 9 9i536ih30ii 93hji59n 6yi0n fcig;>-
igj)n0g36i 33(10033(101^3 51601&3&30O 6003(10 h6mQ3&nb 039331116001. 9i536ihnb
6y0nb h3£)o9nb bhjQymqjncip gibibibnicn3&0ig, ap>bgm6noin ‘b^Qi^nhnb
9noinoi36i 3qjhm 9 hi30 ob9 oi;j930 oi, 3ngh3 516030 ^330106 qjihoimbnbi. qphgm-
6nom lb3gi3nhn [99 ; 201 ] ihnb jbo030ob 510^(10 t^giinhnb ( A j ^ ) 93<3ihg36i
cgmoi0nb ^giSnhnb q)ihoim6ois6 (Aj), i6 ajmoigob 9 ibiboii 6 (163 cgmoionb 639-
0n 6m6i (FWj). qjihgmbnoin ftiSîhn <b3gi3nhn ( A j ^ J A } ) 5^ ^ 3 0 1(136 0 0 1 gn-
gni n9 9i536ih3369n, hm9036nu 633b Byipb 1(11(101^03636 [63 ]: 3Qi66mb 9^3-
61(13011 qjihgmbocno ^gi^ o h o b ^ ig m g 3 ^ ( 1^ 3 6 1 2-3 99 s . 5-I bm0Pi ‘b m j^ h
3jbl|(h39iC3h 3ncnih36i9n 5-7 99 s . &'1 ifp633b; b ^ ^ d n b 8(336111336(16, 9 bni 6 n
- 232 -
8ua66ff>ou Ebc?ob 6 oao3 o
i>95 n0bi0 ymq3x n'J 9my3ihxn 6:>nh6iQib3&nbr Ericaceae -nb 6 ^ 9 0 ^ 9 5 3 6 3 9 0 333b

63^3606 gi b830oii93i(bc’33cnnb ^0 81536^33606 qjihprri6ncnn ‘b^gbSnhn jghnb
0 .5-2 39s . 5'1; <bm0n3hn bih^ygnb b] 8(336^33606 ^S^^bnftnbi - Qiib0rri3&nan
0.5 33s . 5 1 • Rhgngob i 3 l>)i 5 n 9u36ih336obi - 0.2 33 ® . 5 '1; 6yig9(j36ih33&bi
g i bîx^BviObb ifooi 5i6bi33cnh 36001 gibigo 8iR336363go - 0 .02 -0.1 33 s . 5 '1.
cgihgmbncnn ‘b^giSnhnb bngng3 Qi0nj)OQ3&3Qni i5h30i33 8(536ihob (bm336b3. 8(53-
6ihnb bhgnb 3hm(j3b9o ^36oi6 0160b 3ohm63&ob (33i03&:>9m6ibaii6 3hcru>g, 0(3393-
6 i b3 8^33636906 6ogog3(5-
,$0190 fciSv^hn ^giinh^fenb 3ohm6369 n, b33(jnq)n3ho 3oghi3gn33ho 5^8^136^-
hrii6m&i 9 i 3 m3ng36x n i ^3(1^93606 5i6o3 33301^3 gi 8icno 03*0(13606 li,n3^>3. 3>-
536-33^3906 3i6m6ob 806393001, 8090b 5i0,y>h36ihni6oi&i 8nbn hign^bnb 3hoi
33mcnb3g bihnbb3 n iy3i6nb 3hn3mh(50x n i [3 5 ]. ftiS^hn £3(1^9360, hm83goiiu
ij3cn 306(101 cgmh36n g i qjng^Xjnnb 8>C1iCn 6n6ii(j>8g35fTi6i (855., 6063*0363606
(jhij3og36n), i8(joh3636 633^00303(1 309(^39033(1 5 i 3îh 36ihoi 6 n 6 ib. b6mh3p
iSoôiS 606301363690 6ypnb 5^ (136 0 6 36iho 01(133(1 6^393601, 3ngh3 9 ihig 86 -
3^63 qjfTi0191*13^6 b3 3(j36ih3363n, bmgm i 9 33i6ib363gaii 6ygob 5 i (jih3&ob 36^-
hn, cru>3ob 3bhn3, mh33h 8ijnh3i, 3ngh3 ^03639863^63 63 8u36ih336obi (ybh.
4 .2 ). q3ihoimq3mh36ni6n ^3(1^93606 8^0163 03363360 gi 90i63&0 5i6bi330ih36ooi
3ih5ig ^ ( 13 6 3 6 6yi0b. 633(500303(10 5 i3 l»>ih36ihni6m6iij 93 od03 &i 9^33^390(10101
03010103606 0ibiboioi36936b (qjmcnpnb b3j(jnq}nj)jhn îd^>hj6shns6mds [281 ]).
t>3 8(536^300 “611013630 050 y339i"b3 9 ipigoi, bmgm j)i69li)m36nb ig5og3bob
33695018 - 8903(1 3 9 3 6 x ^ 6 6 (6ib. 4 .15 ). i 9 n^m8 6ypnb 5 ^ ( 1 3 6 1 b3 91536^390
36n93639Pi3i66n9ig gi8oi3og36xni (101501(1(5 33(1393606 i6 iljm9o3h 000331136(50(13-
6 ^ 3 , ^ 3 3 3 3ih3nb 5^6^01^306 biboioi^.
ljhi6b3nhi(5nnb 6ii»ignb 9ijbo8ig3hig 93bidg363go 06^36603^16} gi9oi3og363-
0ni £,i9 vj>ihn bobli39nb i6i^m9n3h ^ 3 6 x 3 6 ^ 3 g i 3u36ihob bb3igibb3i ,j)o39o
5i6bb3i336xn i (libh. 4.3 gi 6ib. 4 . 16 ). Byonb 6i3igob boRjih3 bb3igibb3ii
3(536ihnb bb3igibb3i 6 i 6 ng9 o (^mô, ("3(101, 033630) gi jjiSîho bnb^38nb 805-
6000(5 (gngoi boRjih3 q3ihaim(nh30i6 'bhm3 n gi 303(101 gi6igoi - jbog39ob bb3i-
5sb. 4.14 9^bnb uoyYnojmhx 33hji63n

jjh jg x n 6ycnb îfjbSmhôhQ&i (Braun,
1970; Cermak et al., 1992; Anfodillo et al.,
1993)
- 233 -
g > b b 3 i 6i 6np9 n). b i 6 i 9 Byenb 9Q6n3> ih S^ q j^ h b g^ b i, 6ygnb 6>j>>gnb b n fijih 3,
^hi6b3nhi(3nnb n6^35bn3m 6nb lb h g ib o ii6 jh o iig , n“b h g3& i. gngn ,^ 6 n b b^ 9^3-
6 ih 3 3 6 0 n Bygnb 9 m dhim & i gnpnb b iio i3 6 9 n n 6y3& i 33663hm gi6 g i 5b6^m ^3nb
> g5 n03& ngi6 g i 6 i 3 i g n 9 mdhiPi6b bhmb dnhnb]>o6. >9 ghmb n6y3& i ^ hi6b3n hi(3n-
nb 6>i>gn b n6l336bn3hn g n 6 3 d i g i 9b n b i9 m b 3g n b 939g3J> ip 6 3 3 b 9 ijb n 9 iC 3 h
9 bfi33636Q3&b. bipidmb ^hiBbinhi^nnb 6i3ign 9ynhg3&i, 9i 5hi9 306 n3363 Qm
9 m dhim 6 i 9in 6(5 93n6 n963<bi 5 3 n i6 (J>i930g 3 - ib 3 9 ^ 3 6 ^ 3 i5h m 33& b B ycn b
h3i
3nhgi3nh3hm3mhyn3Q g i 0 m3ng3& x 3i>i9 n qphoimbnb 3110133(^3 ihb3&3C i!>hi6b-
3 nh>ijnnb n6^35bn3m6sboii6 (Tr), nb333 hmfcmhu Bypnb j^>h32)i qgmoimp3 n
(Jh2 °) gi 3333hm3mhun3c<9n - qjihgmbnoi 2>i9(jih b3gi3nhoii6 ( A ^ nc / A 1 ):
v = T r/(A Jbnc/A 1) [9 -69 -1 ] (4 .9 )
(3hi6b3nhi|yinb 6i>pgn n9g36ig b6hiopg 3<jmC932)i ^hi5b5nhi(jnnb n6^35bn3r>

&ib, hm9 9ibb3 bi69 ri303 93hy3m6iu yi inbib365. i 3 nt!)m9 bnb tbhm0 n ByQnb
6i3ignb j)i‘bm93noi 93ndp3&i 3n3 b33pmoi 601300 3ih3nb (jhitjblJnhiiyii^ (6ib. 4 .17 ).
h in 9 3 bibnb 0 3 ji6 n 3 3 h n g i ‘b n i6 3& n b ib (9 i& ., dgn3hn jih n b 3nhm &363n ^ , 3 3 -
&nb h b 3 3 i g i & i(T 36 3i), i6 , 013 Bypnb (jhifSbSmh^nb b n b ^ 30 i 9 39 im 6 b
(jbhnpfl 4.2 0i536ih3cni 6b3ig ib 63i ^n39n gi mhj>i6m9n jbnp30nb 3nghi3Qni3hn &i9î(v)6shni6m6;>
(b926d"' 03i_1) (Berger, 1931;Huber, 1956; Zimmermann & Brown, 1974;Raven, 1977; Cichan. 1986:
Ogino et al., 1986; Losch, 1990)
6n53m 3i6n 9jh jb n i5n 0(536^3360 5-10
E ricaceae-rh 03hj6ni6n 6ih0m 0ig&36e36n 2-10

9 ih ^ 98 63^ 3 qjihoimq3mcnpm3i6n b^jbn 0n>)hmq)mh3&ni6 n 93hJ6ncn ( 2 ) 3-15
q3moimc9 (5336n <gihoimq)mcnQ(Ti3i6n 6336(1 0nyimqjmh36n i6 n 03hj6 noi 18-50
qjih^ccn^bfviiBn 0^36^336(1 0 iyim q 3mh3&ni6n g i uni)Qmo9mh36ni6n03hJ6 noi 100-350
[yii636nb 03h ji5n 300-500
O]mcrtfnp0i3336n qjihoiPio)rioiQm3i6n b33&nb <336336(1 200-1500
qj3b3nb ;y>8vy>hn ijm636n 1-2

6iQ ib n i3i6 0U36ih ]o ii ^jh^Qnb ^ 6 3 6 (1 30-60 ( 250 )
Ubhncn 4.3 bb3igibb3i ^n3nb 0^36^ 30(1 6y(yib 6iiign b 0ijbn0iC3hn bnfijih3 (0 69"1 ) (Berger.
19 31; Rouschal, 1938; Huber, 1956; Zimmermann & Brown, 1974; Hinckley et al., 1978; Rychnovska et
al., 1980; Yoshikawa et al., 1986)
0ihig063i63 6(536111336(1 1 -2
bj)03hng3np3h(i 63-0(536^ 336(1 1 .5-3
,536(1160 ^hm3(iixn iiy336nb <gihoimq3<Tioi(W3i 6n 6336(1 (9) 18-34
qjmoimc3l5336n qjihcnnq3moipm3i 6n 9n>>hma)mh360^611 33hj6m6(i 6(536^ 336(1 1-4 (6)
qjihcnmq3mcncn3i6n (5n>ipmqjpih36ni6n 93hj6m 6n 0(536^336(1 (6 ) 2045 ( 60 )
6iC ib n 3 i6n 81536:41336(1 10-60
pni63&n 150
- 234 -
90066601 paÔU 603090
Leaf specific hydraulic Tracheid lumina (%)

conductivity (%)
6sb. 4.15 Abies balsamea-Sa Bypnl) 3ih(jb;)6:> b^hsoin: jbnQjSnb bûnqjnj^hn J>i3|iphni6m-
(jip ben-1 Byonb 6>2pen lOkPa d^&h"1 5 n633&nb 99hion 6m6i); 3ih j336s b^hstnn: *15033 9^3-
6>hnb ^ ijjso jh n (nh^jftnb eniS^hn ((i3) (Ewers & Zimmermami, 1984). iBaeroytfhn SmBujjBaJm
bbji 3|336ih336nboi3nb nb. Zimmermann (1983) g i Garter (1991)
3nhm&3&3 n (0 sft., 6 ypnb fc i^ l^ n b O^b^bâbi 63^C3nbib, i 6 Grug^nb 5iyn6-

3nbib), Bypnb 3 *i4 np-inn 6s]>sqo 03nd03bi 31536 6;>6yp3b. i6 ghmb 31538* ^3330-
gi6 ^ 3 6 3 0 0 6633b (jpjn& ijns) £ i jbnp30nb bnb^SiSn cî^CDhbg b3g3&i 5>3hn
(jddmcnnb dm3Qj6s) (6ib. 4 .18) [47 ; 152 ]. bi6 i 0 jjbS^hn b n b ^ Si ov>3ngi6 ih
03n3b3&i 6ypncn, 3i3hnb 636^3536(1 iO}3hb3636 Byonb &igiip5n036ib. b3 81536}-
h336n, hfn030ou>(5 jbnpQdnb 0i(j>i0n b33(5nqjn3hn 5 i 3(i>bh36ihn}6 m6 b 5}ifi6n}cn,
3qjhm ihni6 0ngh3yi06n 306fnQnrtbig0n, 3ngh3 3n6hm j>i0t)ihn (3+1336116 8)0163
81536^33611. fihgnprnooi i03hnyib gngn g i h&m0nb3dhn q)mh36nb 8)0163 63360
53ni6 6 i 3oiih3 n i0^nh3636 Cbiq)b3pnb 3m6i(53036cni6 03Qih36noi) oii3nb
3nghi3pni3h 5i0îh36ihni6mbib bi03iQfng 55 %-noi; j>i66330n cgmh36nb 8^0163
0jj36ih33bn - 17 %-ncn; 6n63m3636n »,n Sbnprng 3861133630010 - 0 -8 %-ncn [267 ].
386oipnnb 8013036366 ig&n0n ^301 6iQiboi3i6 8(536^3363(115.
b3 0(536ih3oii 83hji63n Bypnb 8rodhim&i 33nd036i 5i8m63330n nymb oib8oi-
b3hn dip3&noiii5( 013^1 3oi(j36(5ni036nb bb3im6i 3j536ih3bi g i 6^3801815330 i^-
3mbqj3hmb Soihnb d0n3h gibipni. ib3om 8g&0i8ih30i6i 33nd03 &i 33 nj36 ib ‘bioj-
b3Q063i63 81536^336811 igh3 5ibiq3b30tb3 83cgoioi03nb 6n6. fcihgi i 3 nbi, 630-
3 nbi gi 6nbpnb ^336116 63 81536^33680, bigitf 3i3hn 6i 33h3&3Cni Bypnb mhov
jpnai, j)m6lii^|)3h 3ih36jn3i3n 936i b x n bibi3360nb 3m6n0n(b36nb} g i bb6ign
6ib3nh6yi06ig36nb £ ^ 0 3 6 8 0 gpSmymqjnb 038330161101 81536^36 333dpOi ii0md-
himb 6yi0n mb0mb3hn ^hign^B^nb 808^013036001.
- 235 -
633090
Laurus - 4
-3
- 2
E,
>»
Ia> - 1
0
i Olea
I>.
X
b b . 4.16 3(}]6*h3ou bb3iQibb3i

6 9 12 15 18 21 (yijan Syprb
Time of day bnfijih] (Rouschal, 1938)
I
£§-
!|i
1J §CO£
£
a
P
k 6>b. 4 .17 6ypnb 5igi5i(j3C3&i 16 0
bn3b(T^>nb 6id30n g i Scn^Qn
82. 3(}36ih3xn bsqjshnb ^hi6b3nhi(5ni
(Tr) (Schulze et al., 1985).
îiQfO^bsoisnb 9m6*u393&n nb. D ye
Time of day & Olbrich (1993)
£
1 =
S'
(bb. 4.18 6ycnb *
Jx»c33nb gi& ipn
6yc°b 3m^]5(jnicnb &i9<n -6 -8
(Speny & Sullivan, 1992) Xylem water potential [MPa|
- 236 -
3(3366601 p a s m 633090
4 .3.3 eycnb W h63i ai336^hne^6
9^36^3 6yi0b fciOmymqjb mhoijgob bibool (ljh^ 6b3nhn}n^) g i <bmj,33h anb^-

3>gn biboaii^ (dD&>l5n*)- 630îyi^>, bn0ynhnb 5i0m, ByQnb h3go0oboi3ob
96 n936x fTi&^ ^hi i^ b , iQn^nQ 93093580 ”6ycnb fciQmymcgob" 1)330 6 i 5 X nbb03-
6n oj6;)6i 0bm0mg ,jhi6b 3ohiuoi.
4 .3 .3.1 33i 3n h il 5f1^ i!>D6fl^6n u 3ghm gogoi mhcnjQnb 66jjn b fthspnjGtfn >0 ^QiSohbi g i i(j)0 mbqj3hmb
3 mhob. i(?6086300 j,hig036(i)0 6ih0moj0636i 0i0n6, hmg3biu imhoij036nb
b3gi3nhoii6 ih b 3& x 3i3hob qj36i8o 6y0nb mharjgob 330U330m6i (0mg O’3, i6
5h 9"3) 3qjhm ©ipÔi, 30gh3 i0 (b3gi5ohogi6 0m0mh3&ooi. 050 0ipi0Oi, hmgj-
bi(3 iQimhanjg36300 <b3gi3nhn jjihîg ihnb 6y0oai 3^(1363301^03000 g i ^3(1^3
3ghm cnbogni. 0t>ob d0o3ho higoiôi b30b 36ym6b i9imhoij03630o ^giSnhnb
5iai6ni6ib g i i 0 ooi - imhaij036ob £ ^ 0 0 3(136 ^ (6ib. 4 . 19).
6 ib . 4.19 3i3h9n 6ypnb mhoijQnb y T)6(j36-

bb3igibb3i pi
mhonjpnb 663306 ^hipn^S^n (AC = C ^ , -
Cact) e* 6>330h3630 3i]hb
3mhnb 051133 j)3933h i^ 3h i<b 3: D P - 6idnb
6 ih 9fnJ96 nb 63h^npn; C ^. ^hidbSnhnhjbipn
Temperature [°C]
imhoij036ib, hmg3bi(5 i0imhaij03&30O b3gi3ohobig0o 6yoob 0o6mg36i ^ 6 3 -

bi'bp3h30Oi g i 6y0ob mhoijgnb goqj3'boib ihiqj3ho 36o6ii(p0g3536i, 36mg3&i
5mli36un3hn 33i3mhi(jni ( E A b36vjhm3oi36ob 38 hi0 h3500163680 3mlj36uo3hn
imhcnjp36i i[0633b 10-15 00 . gc’3'1 (= 1 i6 H20 . 9^ t^giSoho . gC’3'1)< b03p-
- 237 -
sycm rioaoao
0 ii 93 i'b(j'330>nb h3&nm69 n 99 hig S^hnmgSn - 5-6 90 . ec1]-1, 3 i3 i1y ih ]c bm6 i 3 n

- 3-4 90 . g p j 1. bm0 noh b ih ^ c B n 3m(i)36(jn3hn 33b3mhiyns biojb30nb 3 bni6
pp^b ip633b 4 00 . e C J 1, 3060ib:>l5nob 33horng9 n biSîQnQ - 2 00 . gpo"1 -
bmpm Cb>0cru)h9 n - Qbrngmg 0 . 1-0.2 99 . q^q'1.
6nigi5nb ^360*60 b 3g i 3ohnb qjyj^f^hn 303153b 030oib333b9 n
jqjhm 0unh3i, 3ngh3 ^rn^jS^njhn jj^ m h ^ n x 3n6ingi6 Bypnb 9 shij,n cnncnjSnb
ym330oi3nb fc iS b ib ^ h x o i.
4.3.3.2 C)hi6b3nh^ni, hFl&mhb °3frbni)3hfl 3hni30bfl
0U36ih3369n ^h^bSnhiuni bmhun3pg3&i (5360560 b3gi3nh3dngi6 6ypnb imh-

cnjQ36nb Jji6m6bm9n3h3636nb 0nb3P3ncn. 6yipn trihoijcg36i 0jj36ihnb y33Q* n3
‘b39i3nhngs6, hm03cn(j 393*9™ 3*3hoi*6. 3b 33odg36:> oymb hmfcmhu
5ih3, ^ 3 3 3 3 og* b3g*3nhn. ai*pmq3n,53b9n 6y*pn mhcnjp036i oiipjbnb 5 ^ 3 b>3-
9 i 3nhngi6 , £jh^0Pi3*6 9 u36 :>h33&3 n - 33ng3h0 nbnb 2)3^116001 g iq p h x 6 ^ 0 ^>39-
93609*6 (hOtf/iiOCjhn tihiGb^nh^m) g i 5*>)mh33&3pn b3g:>!3nh36ng:>6 ( 3jh n gjh -
d jc n t!)hii6b3nh±uns). 3i536*hob mhf,b6m36nb 9 nj>6 noi Byign mhaijpg365 33h3gnb
b39i3nhngi6 33h3g9mhnb3o. Byipn 01633*90 9g6m9*h3m<bng;>6 jpgignb fci'bn-
b563h9n g i 939g 36. &i&33&ob iSihiôb 93933^16001, 3*3h3o (b^nbdnjhn ^h^6 -
bZnhsijns). 0u36*hnb b3g*3ohngi6 Bygnb mhoijpn gjigignb !b3hnb 9fnbibp3h3
0336*90 (bibib03h3 0336*), bmpm njng*6 - :>^3mbqj3hrn3n (6*b. 4 .20 ).
Water vapor conductance RH C H! ° T
[mmolm"2 s '] [cm s ’] [mol-m3]
4
o&° 001 v -
(2-16) (0.005-0.04)' ,, v- L
2000
- - -f - ' ; ‘ W %, ’ , * v '> -
5 : \St< ^ ^ ■W% V „ 99% 1.27 25°C
(1600-4000) (4-10) < > -
200 0.5 ''. -

95% 1.21
(160-800) (0.2-2) -- >
1000 2.5 “ , 47% 0.60

(300-3000) (1-8)
9|; 154 3.8 .... 50% 0.48 2 0°C
61b. 4.20 Stbnbse&i 9n0shoixn ojmoipnb êbSnhi^yiiSn Rshoix0 ShcujDbjbnb bj30i: nbh^&n -
Jbm3nco63n Bycnb J>i6:>6nc3&i, 03iimhi(jni 3jh3g3 mhnb bhjSn 91 i00>nixnli 6^30°°". Bypnb
mharJcnb 5^30^ 6*633680, ambi'brehj a t^ n 6ypnb mhoiJenb 5*hbo6n; 8^ 33603: (5^ 3h ^ 3h i (T).
i>3hnb ophgmdnoin ^360*6m6i (RH) 91 6ypnb mhorjcnb 5m6(j36^hiuni (CH20) 33h3gamhnb
bn3hi}33n, bi&nb j 3X b ih39n, 9nbi'b(p3h3 9h39n p i JiQhnb 3h30n-, 9ih|jb6n3: 6yp n b
mhorjpnb gnqj^'bnjhn 6n6ii(jêTj)m6nb ^Sn^hn biSîem 9^^3363603^° (20°C 3*3hnb g i 2 5 °C
qjmoinnb ^ S h i i J ^ ' b a ) : g,.w H2°i3epf>iXDhn ohi6b3nhit5ni; g j^ O 3I h3e3ne b: g<,H20 6ie,nban3hn;
gaH20 3mbi*bc'3h3 9h^; giH2° qjmoxvib biohcnm 6n6ibCf6902>m5i (Nultsch, 1991; Nobel, 1991a)
- 238 -
3(3366601 ffaÔb 633090
5*h30oi (3*^0^360 *3 09936*0 *693636 5*3036*6, h*6936*9*13 83-

3dpn*cn 3|336*hnb 'b39*3nhb* 9* 3*3hob 0mb*(bc3h3 0336*6 3 oihob Bypnb
ynspnb bhsQnj6 $nb 6090906 93(530*. *b3, 3*5*9001*9, h jh n b bnddhspnbs gs tfjd-
Jjh ^ jh n b 0*(j3&*boi*6 3hoi*9 0 ^ 9 3 6 * jijh*6b30h*i300b 06^36603016*15 (6*b. 4.21
9* 4 .22 ). 0 bnb 9*(j>*0n h*9n*(jnnb 83935*9 3300)936* <301010360 5*9*b3hg3b 9*,
3fi3b39*3*9 3*3hnb 0*(p*on ^360*601606* (0nbn ^36001 5*j3h3&ob 3300^333*8015
50), (i)hi6b3nh*i3n*0 0 oin0 *ôib. *0 *b 9090 06033630016* *j3b Bypnb* 9* 603010-
3h36*oi* kih*6b3mhl|)Ob*oi3ob 330093b h35om63&3o. jshn 639b 36ym6b 03010193606
0o0g36*h3 ^36001 5*33h3&3pn 3*3hob 03360b 601(50036*6 9* *01(101^93606 ^13-
9*3nh'b3 5*90mo6*u3C36b 3*3hnb 3qjhm 09 h*po 0*b36o. 9090 cg*hoioi6ob 0^0163
(3moi036ob*oi3ob (0*5., 6*6*6nb 0301019360) 3j*hoi *006980 b*b*'bp'3hoi qj36nb
6o6**(t>0935oi6* (ob. 6*b. 2 .15) b*033h 03^0*, 309h3 0|3oh3 boi0ob 9*6*633h363-
00 *6 630bnb qjoih0ob g3oioi036ob*oi3ob. 3b 5*6^3*336360 j*hnb 6*^36*601*6
E
0CM
1
o
E
c
o
CL
to
6ib. 4.21 CD^69nb gjmoie^&nb (jhi6-

30 40 50 60 70 80 90 10 0
b^nhi^nnb Qi9myig36x 32>i
pi 3i]hnb lj36ni5m6i'b3 (Burian, 1973) Relative humidity %
61b. 4.82 0^363630 6y(yib SjcJîxvn&nb eC'XP^S^hn

pn6i9ny. oiiQ]bnb g i 3v)hnb
qjihpfn&ntn (j36ni6D6ibau>5 j>*39nh3n (Kershaw, 1985)
- 239 -
e a ^ n u rtaanao
(jbhncn 4.4 &h*6b3ohi(5oob 9s#jn8sc3ho n6^]6bn3fn6i 9rnh0jmcpi&03hip & WCm6n3l'iP 6*6^3^33-

63Q 8(536:^13011 5033680
9[535»hnb j,n3n ô6b3rhi<jnrtb Ho^fbnj^ (ji Qnipn HtO

9® 63"') 55666000 615336^ 339crib333i3n
^360160 ^hm3n2i36n
63080b ,jyob 636(5361113360 leooag)
6ob(Vii5n ^yob 63-8(536:1113360 400 (2000-3000)
0016360 2000-8^3
b38nshngxyi (ihminyjdn
3108360 1200-1800 (2800)
8SisC° 6S03&n 800-1400 (2000)
SiBjhmb 510360 600-1800
b3&ijhm3oi)3po "bmBnb 3Qi66m3&ob 63^360 gi
6sb33hig63fiJ36n 2800-7000 (10000)
b93COtt93><bp33oinb bj)C3hno}oe3ho 8(536^3360 (600) 1500-3000 (4000)
lbm8o3hn bih^ypnb (jy33&nb gmonmp8(5336o 63360
60610100b 9my3ihxn 6i63in636o (1500) 2500-3700
Khgopob 185160 61631*16360 (780) 1200-2200
9ihig953i5] 6n63m3s6n 63^536^3360 1400-1700
335* 63ft^360
• (iO&ehfib 150450
9i(niC6anob 1800-3000
rohQ365ni6n 6i0ibfTi3:>6o 8(536^3360
9iCpiC6iCib330m6i 9000-11000 (16000)
^jpornqjryjjin 5200-7500
6(50(^(30^360 15003000
91^1080106 9^36^3360 (1500) 30006000
3^(530(1136360, ob03&n pi ôpob36h6o
536ghob 200-350
9e30mb 3000-4500
03h59nibn 3160515360b 5000-10000
83hi0o 3^05^3606 (1800) 4500 (9300)
6i6i3ohm go363&nb 20004000
39*66mb 8(536^3360 10005000 (8000)
liQmojn^&n 1200-2500 (4500)
8(5361(13360 633303653110 9001036001 800-1800

i>ib3b3&o 600-1800
8(53(1130 8(5361(13360 5000-12000
QhcnbQ 3 (5nhQ3&h: hmgjbiij Jihnb bnRjbho x?633^ 2 3 69"1 g> 9 ^^b, bib>'bc»3hm
crj36nb 6n6sip0g35m6> 6 >iC0&n> 0.5 69 bd-1; >0 ghmb 050 mg6>3 gibiQni 6 ^ 533-
6nb gncgj'bn^h 6r*6iXP06O6fT^)i<':,D- Q^nghm 6>Q>b6>hb> g> bnb 3^336611 jihnb
bnRj>h3 dgnoh 0unhg3&:> gb ^n^mS ^hitjbinhiunnb n6^36bn3m6bu i)03 £>30 m6 b.
8 *jbn& 03hn vjhs6 b3nhs(jn>. d>jbn3 >03 hn ^hi5 b3nh>unb 636361103
- 240 -
313066601 f f y s m 603030
5^6^0163630*1 33i 3mhi|jnxf' p ^nh o^n b 3nhoi6368n ihnb 9u36ih30ii 3n3h

Byonb 833^(^39^30 ioihcnjo3&i. (jbh. 4 .4 -8 n 9 oi(53930oi b9303Oinb 8(536ih336obiov-
3nb j)hi6 b3nhiunnb n6lJ36bn30i6nb 9 i;jbo9 i 03 ho 6^336360360 pni 6i533&nb 838-
cnb333i3n. 'bp3nb 6yi09u36ih3368n, 00^3636^ Qi bi3b368n 6y0ocn bh30n 5^33(13-
6nb 930cnb333i3n ioihoij036i, dnhooiigig, gi8oiing3630ni 8ioi 801110301001503(1
cru>3nb363h3636'b3, 6^5., b3gi3nhnbi g i 9 oi(530oi6 nb oii6iq3ihgoi6ib3, b3gi3nhnb
bo>h3jii3hi'b3, giôi^nb bnb0n(i3b3 g i 6i0n3nb 5i63ncnih36ib3.
(jhi6 b3nhiijnnb 6 ijbn 3 i 03hn 0 6 ^ 3 6 ^ 3 0 ^ gi8oijjng3630ni bhgnbi g i bibnu-
oi(3b0oi qgoih0ib3 ( 0 i 6 -, 6i0iboi3i6n 01(10366^6360 gi 9 ih 15300136360, 3 ihig 3B-
3i63 gi <301010106(53360 b3 6(536^(13360, b3iXD®(b3^ft Q* ^>3.) g i 350100^3(1
(1)031)3 (3 ^6 - 3lJDhi3n 8(536^3360, îOi6nb 6(536^3360, Rhgn0nb i 3 ^>6 n 6(536^-
h336n, 9bni6 n g i 83 hi0 n 3i6nl5i(>i36ob 8(536^3360 g i bb3.). Rhgn0 oi-ip3 oibi303oi
ibnob 0gn6ih336nb 3nhib 3 mbihg 0ipi06i0ib330oi6ib1 îoi6ob g i 6(53hi3 8(536^-
h33&b ibibnioi36oi (i)hi6b3nhi|5nnb y330ib3 6ipi0n o6(j)36bn3oi6i. 633b Byi0b
ioihcnj03636 i5h3cn33 6boi6o 3i6n(j i (j36nb 6i0iboi3i6o 8(536^3360(5. 8ioioii6
939ih36ncn, Rhgngnb i8 v|)i6 6inh6i0ib36bi g i 0301010108(5336 b3 3(536ih336b ifocn
mhj3h 6^30360 (jhi6b3nhii5nnb n6(i)36bn30i6i, boi0oi 6n630i3636b - j>ng33 303(101
6^0360. 33i3oihi(jnnb oii6i6ih 3ohm&369n b3j)3036^36ob 303lJ3boi6i 601001 ig6(i-
0b3 ip8oifi6g36oigi, oi36(5i 8ioi &363&hn3 3i6nîl»)368n (8bni6n, 88hi0n igjn-
0360) obo6o 6 33(1^ 83^ 6yi0b ioihcrrj03636, 3ngh3 8^nghoi 6i0ib6ih8n i6 ^36-
ni6 ^ 6 0 ^ ^ 36 8 0 5i3h(5303&30o 6(536ih336o. B3n8obi gi 6ob0ob 1iy33&ob b3
0(536ih336b ibibniOi36oi J)hi6b3nhii5nnb 83bi3R633ig 3qjhoi gi6i0n n6^36b030i-
6i, 3ngh3 boi3n3hn bih^yonb b3 8(536ih336b, hijj 5i8oi63330ni ioihoij03&nb
b33i5nqjo3ho 3nhoi63&noi (i8 030cnb333i8n, gi6ign 33^01(^(50300 9^30(1013001).
jjhiBbSnhiUfii, hmôihy gngjbnjhn ^hm^^bn. 0(536ih3aii jboi3n03&ngi6 Byonb
mhcnj0o 5i0oignb gno^bnnbi gi gn63&ob 8 3 8 3 3 0 ^ 0 1 (hoig3bi(j 3jh3g8oihnb368o
3i3hnb B633i 5i6bb3i33&30ni). 013 imhoiJoQbib 6i63nbn0i3oi h0i50ih(5 goa^bo-
ob 3hoi(53bb, ^hiBbinhiijnnb n6£)6bn30i6i (oih, 1101501(1(5 Byonb 5 i li)ih 36i 0301a*-
oi09n - Jh2 ° ) 2>i3mnbil!)35i Byonb othotJonb B53336nb bb3iOi6nbi (ACH2° ; 5i6ov-
biiiXn. h0>50ih(5 H20 -b hioig36oi6i Soi^xfiôb 3hoi330b3, 3. n. 5(1 6"3 i6
9oi0n 9"3) g i Bycnb mhoijcnb gnqfjbnjhn frsd&hjSshnsBmbnb ( g ^ O .
300, hoi5oih(5 b8 B6"1 i6 |i6oi0n 8"2 B3 "1 ) 338 330 ^ 0 1. ib3, hoi3 :
jH 20 = gH20 A C H2 ° (jxSnon H20 9 * 69° ) (4 . 10 )
l|)hi6b3nhi(5nnb n6jj36bn3oi&i 5i9oinbi^36i, hoi50ih(5 Byonb 5i(530i ajihaioi6nbi

(qjoiai036nb 338oib333i8n, 3 iom 3 oiQni6 n q}ihoio»6n, i63 b 30 i g i j33Qi b3Qi3n-
(1360b q)ihoioi6orii 3i8n g i ihi 3hoi3unh3&30 n qjihonoi6n) g i ghoinb 3hori33o3n.
gnqj3bn3hn 5 i6 l!>ih36ihni6oi6i (gH2 °) 6ih6oiig536b gnqE)bn3hn 6 n6ii(p 39350>-
6nb ("li)hi6 b3 nhii5n3on Bn6ii{n0g35oi6iH [2 2 1]) b i 3nhob3nhoi bogng3b. 03010103680
Byonb oihoiJonb gnq)3bn3hn BnBiipag^mbi 93g636i bibib^vjhoi q^Gob,
iDiSn.M0obi g i 6i5336nb Bn6iiQ>0g35oi&36nbi5i6 (nb. 6ib. 4 .2 0 ). 6i5336nbi
(r. \ ns kakniiTmhn (r ^ RnRssnv)nnxmh^n Ronnnnhhno ^hoin>6036b hf^^rnh^J
^ ^ ~ I r&i63 + ^ ^ (4 .11)
& &3pioiH20 = e5b6DH2° + gj)>iH20 (^ 9 ^ 0 ° 9 -8 B9 -1. i 5 b9 B9 -1 ) (4 .12)
- 241 -
Pulmonaria Coronilla Arctostaphylos
( Upper side
Oxalis
Wind Wind
6sb. 4J 3a-d 6*5nb bb3*g*bb3* ^n3n gi Bygnb mhcrrjQnb 5*1^361. a ohoijx^ e* 3r>5hn-
0*9 6*6o*63bxn 6*533609*6 *pihoijo36*. Snh3x 936016333*90 0^8636* lP0J33Cn 6*b33h*gbo)3h-
mb3&ho 336093110 Shj, bmpm 83^3 336016333*30 - 5*38010*636300 336093110 9h3; b 6*5336110 5*6-
C*536nb bb3*g*bb3* (joSn - 6sh(j3C(n363&ob (3hai9*630inb dnymQjbnoi 5*60*536x0). mhQ366n-
*63&ob (3m3m5363h*g 5*60*536300; 3b ,jo3o y3X i<b0 <3*hoimg 6*3h(jX363Cni) P* 563°Dhn 6*5-
0*5360b (33iS3hrn5363Q*g) tyi3o; c 6*533&ob 5*6030 ^hocn. 0*h^b6ng*6 6*833603: 3oh3xn -
bunmq3nl!>36obi 9* 3n5hmq)ol536nbiOi3nb g*3*b*b0*a>363Q0; 33mh3 - fihgopob *3>j*6n
6(}36*h336nb*oi3nb p*9*b*bo*oi3630o; 93b*83 - 33Qflmq)ri)j36nb*oi3nb g*9*b*bn*oi353on; d 6*533-
609*6 5*3mb3onb 836935 6ycy>6 9mo3iX!l^n^ poqotioob 5^1 (Haberlandt, 1924; Esau. 19 53;
Oehlkers, 1956; Pisek et al., 1970; Braune et al., 1987; Mauseth, 1988)
qjmOTQnb Qn^h 6y0ob mhoijQnb &*0li)*hn*6md* ( g ^ ^ ^ 0 ) 0*,j30Pi6b 6*53361-^

J>ibb6n0m6nb 9m3*(j36*bcn*6 ^haiig. d^ nb d±jbnd±Qjhn frid^hnsBmd* giOmôp;}-
6300* b*0 J>330 0U36*h3<n* (b^b^mb^n, 303360 g* (169030936013 30) &*j,nb * 3*h>-
^nb ai*3nb3&;)h3&3&<b3 Cbm0*, 0m6ym6o0m6*, 5*6*60036*, bnS^nghn^; 6*b.
- 242 -
9(30666013 ffdÔb 603090
4 .23 ) gi i 9 n^m0 6ih0rii0536b 06093630^1316 9m6n330b 0 ijbn0 i 03 ho ^hb6 b3o-

hi^nnb 361(106 gibig536ig. 5166121301(136001 9ipi0o gncg^'bn^hn 5i9(])ihoi6ni6i
ibiboicn36ai 2>DCvb3h30 0i536ih336bi g i Sjpnmign^Qbb (300-500 |i9m0n 9 "2 60"1);
01(11530^1363600 3b 9m6x530n 03hy3m6b 250-400 ji9m0n 9 "2 60"1 qphj,0369n (C4
91(1(5390136366 303(1m 0 >(pigo 1^301, 3 0 ^ 3 C 3); 338936 0 mgoi6 qjmoi0fn3>6n b3
0(5361(13360 g i 63fij3&o ( 160-250 |i9n0O 3 2 63 "1); ySDC^D DQjhm gi6i0O 0m6i(53-
03611 ij3cn 6063(1136366, 15(130133 b2)03hmq3O03h- 9b 632)3036^3)103111010^6 9 (336b-
h33&b (gbbb0m36ncn 100 |i9m0o 0"2 60 "1 ) [122]. 606301316 0i536bh3369n 6150b
bSbhb^nb 6n6b 6b6n00n 30011)^361 j>nih3nb363hn bbi3m6r», (11(3 1/3 -ncn b0(30)1366
6y0nb mhoij0nb go ^b o o b n6^36bn3m6bb. y3i3O03&b, hm^mhy B^bn, 6153360
0)jnh3 hbm036m6ncn ijsoi, b6 b30b|j bh j>bbFi6nbcn, >6 0336115031119 5191531)13630(1
ihm6 (015., (jhm3n2>30n mhjng336n [9 3 ], 31(190636(1011 mjibnb 61(10^01953603611
[85 ]). i 0 n(jm0 931300360 6yi0b 51(311036001 0150(13 him936m6ooi 9 1 6301 y>h-
51336, 309113 <301010360.
i3jjnj>XD^n »bhs6 b3nhiunnb ghmb 6y0ob 0 (^033X 3^ ° 5igib06n0936nb6 33093(1-
0obnb 51(13 i390ob 33^06016 0(133600, i3(!)Oi>30i0o g i 33oi 3ôi303hn 'bhgob 033-
63600. i 3oni303ho ^hb6 b3nhii50i 03009361 5i63nbo0moi hn^mhu 00033^01
3oghnq3fri630 5bh30m9o. Byoob mhcnjgob j)IW)r'J)303hn 5b0^bhm6m6b (gi;wH20)
d0O3h 9161901. 6inh6i0iboib 61b qjmcn0360o 3b 0m6n330o 1^16336 10-20 (i 9m0O
3 2 63-1; 63 0(336ih3oii cgmoi0360o - 6-10 (i 0(Ti0O 3 2 60 "1; 01(119063163 qjmcn-
03600 91 60633660 - 3-5 |i 0m0O 3 2 B0 "1; brn0m 3gi66mb b320936(j3h 0133-
61(133600 - 0.5 |i 0m0o 3 2 60-1 [ 122 ], 'b3g i 3oh30 n imhorj036nb5i6 91^(51319
9090 06003630^161 1^36 ‘bhgobi 91 51(130™ 3ohm&36b. ib3, 0151900119, 8(5361-
11336b, hri0036oi3 6019150b! 91 3i3hob bo00hi0ob 3nhm6369o o‘bhg36oi6, 303)1m
bJ30O i)3O)ni30nb 0h3 ij3cn, 3ogh3 ^36016 3nhm6369o 9 m(bihg 0(5361(1336^ 3^0-
93h9obob 51(13011 j)390ob 5b9m0hm6ob 030cnb333i0o 3oghmqjm&30 o 0336360 3qjhm
0^09(1^9 32)3hni6 3hoi0i63cnb, hob 5i0m|3 2>Dibn.MCDho goq33'bo3ho 6o6np0g35m6i
mh0i5936i. 0336360b 5i0îhob6m6i 916190 ^3033)1i^ h o b 3ohrn&369n
0(50(19361, bm0fn 0i("i0O ^ © ^ h i^ h o b ghmb - o‘bhg36i. 03133 6103^360 1^0163-
636 qjmoi036obi gb 6063360b "bhgobi g i i3O)f'i30Dh 033636b.
61b. 4.S4 lbm3n3hn bsh^ypnb bb3spibb3i b3

8(335shnb (bbjigibb3i jîSnb jjhjnb 6^ 63)
33hn93h9xn J|hi5b3nhi(}ni. ^hi6b3nhi(jnnb
bnfijih3 ps9mi,np3&xni (JDS^hi^lhibs pi,
° M A M J J A S O N
6b6b ii3cnh36ncn, Sijhnb bn09hi(X)'b3 (Geurten,
1950) Months
- 243
gi p 3hni3&nb 3ohng]h0 x n »jha6tonhi(jnnb n6vj)36bn3m6i Qi8minp3-
&30 ni i 3hng3h3 nb *536 30 36 ^ 3, mb3ob3&3ho ^jh^gD^nb bnbdnh^'b] g i 6 * 6 ^ -
h36ihni6m6ib3 gi jjmhSnb <3363690 6i 3hip36 nb ihb36m6i’b3. b6mh3Q i8nljm0 iQ-
30b bob, 93bnb, 63i3hfib0obi g i qjn^jnb ^<^360 3qjhm 83^ 6yi0b imhotjp3636,
3ngh3 6id3nb, Sncgonbi g i ihyob bnb ^m^36n, hm830cniu 50330 g i bfogn ynh3 n
ij3cn (6ib. 4 .24 ). 'bm0 o3ho bih^yonb b3 8^36^33600 ^ 0^3090 33hng3h930 n
(ihi6 b3nhi(5nnb n6li)36bn3m6i 03hy3m6b 0 .015 -0.15 |X 0ni0O 8"2 60 "1, i6 3 1 %
3rti(J36un3hn 33i 3mhi(jnngi6 . (j)36ni6n ljhm3ni36 nb g>033j 3hjni6 n b3 0i536ih336nbi
g i 09 hi0 n h3&nm636nb bj30imh3 ni6 n b3 8 5 3 6 ^ 3 3 6 ^ 93gih36ncnn 6 ^9 ni 3CD3 D-
6n 33h ih fiiîh3630i.
4 . 3 . 3.3 i!ih i6 b 3n h ^ 5n n b <3 iTbnm em &n:)hn 2>m6i!>hmen
^hi6b3nhi(3nnb 06^36^3(116* 8bmQFig 0 i 9 n6 ihnb imhcnj036ob 3nhm636'b3

9 oiQni6 ig gi8myig3630o, hmg3biy biftnb i 3ihi^n ih 15300b 5ibb6n0m6nb b i-
hnbbb, 3. n. hmg3biu 6i533&n 03g0n3ig 5ibb6n0n i6 gii3(inpni. 9bm0mg ib3cn
5nhm6363ni Bygnb 6X53830° himg36m6i imhcnj036nb 36ihnb ^hmimh^n^Qn.
6i2>nb i! 3ihilyib <336^50m6nh360b (13530(^136^ 03033^6001 0(j36ih3b 033d0fii (h i
6ib. 4.25 0ihyb36i gnij,hi9i: ^Ki6b3nhiunnb ejnnjhn pn6a9nj)i. A 6ypnoi jtih&ip 9m9:>hij)3&3C'n
8(536^3 (S jjtic x iS n <jhi5b3nhi(}ni); B 9(j]6i.h]. hm9pnb 6ycnCTI 9ro9shi{,36s n'be>3g3&;>
^hnrnpSn; C 915361113, hm9pnb 615,3011 îbbnpm&nb bshnbbn b3(y,i363hip h3j>3enhg3&i
33iimhi(}nnb (Ep) 9nb393001. 811133361 gmjhiQi: ^hi6b3nhii5finbi pi 33ijmhnjnnb on6iq}ihpmdi.
&i9m9pn6:>ti3 9iriub36i 6ibibnb 3m6iu38360916. 013 ^hi6b3nta(sni sh rft>Q>3g3&i, 9ib i;)3b 6hqjn3n
fciambi^xO^ 3-n- <lhi6b3nhny\i 3nhei3nh5hm3mh^yi3cni 33>>3mhnjnnbi. (5hi6b3nhi(jnnb 33<b(73p3b
(16333b *9 gi9myip3&X3&nb pih("333ib. ,jhi6b:9nhn5nnb bnfijih3 9unhp36i phmnb I2 9m036l$ngi6
t3 9m936,ji9p3. ym33tyi 6ê^t>333^ Tr/Ep 6hqjn3n cm6:>q)ihgm6npi6 90301001366 ,jh:>6b3nhn5nnb 3hm-
U3bnb 6ij)nb9n3h
- 244 -
8 (3D66 fn0b e a ^ o u Aodoao
ovj0i 3695, 9 pbid036 Qm6 nb qjih50369n) (5hi6 b3nhi(3nnb

cru>3nbn 6y0nb h^gnQnb 9mcnbm36n0363611b 93bi6i9nbig. 6i5336nb 5ibb6n0*n6nb
bihnbbnb himg36m6hn3n 5^^0163^ 93nd03&i 3mhm03^hnb bi03i 036no». cgn'bnmQPV
5n^hn 3^35301^36300 03 j5 n'b336nb Rih33i njngi6ii5 50 Ri6b, hm9 ^l^bSnhi^n-
nb n6li36bn3m6i ipih 933bi6i336:> 33i 3mh:>i5n x gi^n hoîb (6ib. 4 .25 ). 6^533-
6nb phn36ncn gib3h3i 93ndp3&i 5i9*nn63nmb 5i6iOi3630m6nb 9^33*10136^9, ^ 3-
hnb 5^9n9hn6i9 (556^530^361101 jihoii6 3haiig), ByQnon 0m0ihi536nb g 3030(3(i^-
3 i. 3Jb(jh39i 03 h9 i ^ 3033hil53h369i gi ^mjbn53h0i 5^ 36 0 5.
6i633&nb bjhjQn, dm jdjgjdnb bnfijshj p i jqjjjtfjhm bs 9^36ihnbi g i 3i 6 fi(ji-
^nbo^nb b33i3nqjn53hni. Rhgngnb i9(5i6n 9(536ih33&n 6i^336b ^9i0 63^ 336
6y0nb 39609363001 6:>5036m6nb ghmbiij 50, bmpm bn6ioi0nb0my3ih30n 6i0iboi3i-
6n 0^36:^3360 - &33hig 3qjhm 53ni6 g i cni6gioii6oi6n0i. i8 9bhn3, bb3igibb-
3i bib3m636n g i 53601,503360, i5h30i33 3hcnn g i *151133 9(336ihnb 0301010360 ib iy
nbi (6i5336nb ihi 6 mh9i 03hn j|333i 5i63iicnih36nbi gi bn63hnb 03^3690) gi
9u36ih3'b3 9icnn 5i60i536nb 0nb3g3noi, bb3igibb3i6inhig nji533di6.
6i5336nb gii3ii3nb bi^tyicgn^n TarrU~ihm6i 5i3mnbi^36i j)hi6 b5fihiy*inb
0nig30iun3h i930n^3gi9n, 3. n. 933<b(p3gi3n ji93hn lj,hi6b3nhii5nnbi g i 53^05X3"
hn l|jhi6b3nhit5nnb 93qjihg36i9n. Rhgn0ni6n g i ^36*^611 3i6nl5il536nb 9y36ih3369n
iDiini 303hn ^hi6b 3nhiuni 93ig 536b 3i 93 hn ^hi6 b3nhi(3nnb 3hai 03 bi93 gb.
9i3hn3j3hjni6 9ihig963i63 6(163(^136369*1 g i x^fSm b &3fij369n 6i5336nb gii3(j-
3nb 5i 3 m 6y0nb 5 il 530i J33*ioig36i 9 ijbn9 i 03 h*i (5hi6 b3nhi[jnnb 3 - 10 %-9 g 3, b3-
5X06(5369*1 - 1-2%-0g3. 3b n9 ib 6 *i96 i 3b, hm9 imhovj036nb5i6 bi3530i3bmg gi-
UX 9 l306ih3369n 53(5fi5X3hn (|)hi6 b3fihi(3ni 9 u*ihg36 i ^36*1^60 <b 3g i 3 nh36 nb 3m-
(536un3hn 33i3mhiunnb 0 . 1-0 .05 %-9 g 3 [ 181 ]. 3b 9m6iU3036n 0nc'3&xni 3i-
6nl5il^9n ^ 3 6 3 0 * 1 imhoij036nb 3nhm636nb 303i0m 5 i‘bm03*ib 93 g 36 ig- 6553360b
gi 53ii3r|b 3<33ji53hm6i 0 m|330x 35m0m5*i3h 3ohm6363 o 35301 5 i 9 mnbil536i 53-
(5n5303hn gnoj3ebn3hn' 5 i 9 (5ihni6 m6 nbi g i ji03hn 5i0l5ihni6m6nb 33 q3ihg 36noi:
^ &wnmH2° - 9 o9oib333i 3 n 0*13(1013601 939g 35 9 m6 ii53036b: 0 .04 -0.1
(6i0ibm3i6 ai336ih30ii0i3*ib), 0 .03-0.06 (030101010913336*1 b336nbiOi3db), 0 .01 -0.02
(9 ihig 963i 63 b336nbi g i 63Rj3&nbiOi3nb) g i 0.005 ^353036(5360^0130^.
4.3.4 0l536^h(lb ByCflb 6^C^6bfl
4 .3 .4 . 1 eyoflb 6^ 6bfl: Qn6i 9flWhfl 6n6ib6fnhrn6i

6y0ob 930i3*ib36i, 55,5511365 gi 5513305 ihnb 9i336ihnb 6y0nb hQgn0nb dnhn-
oiign 3hm(33b36n, hm0036ni3 3hoi0i63oi(b 3i gi9m5ng3630n. 013 hi9g36ig 5ih5ig
bg36i ghmnb 0 mi539x 9m036v!)9n 6y0nb 93cn3nb36nbi gi 5i(339nb h35xnh36i,
93bid03&30 ni gig5n6g3b Bygnb 936m3n0n g i imhoij03 &xn hi*ng36n636nb 9 n-
bx3noi. 6y0nb 6i0i6bn 5i9mnbil536i hni5mhi3 i6bmh6unib 9n63b ^hi6b3nhii3ni.
6xnb 6i0i6bn 3ihymq3noini, hmg3bi^ 6y0nb 936m3i 6iX 3^ n i li)hi6b3nhii5ni‘b3.
3ihyoicgnoin 6i0i6bnb ghmb 6i533&n nb3h3&ni6 n9 <bm0i0g3, hm0 (5hi6b3nhi^n-
nb n6l536bn3m6i 933bi6i036mg3b 936m3n0n 6y0nb himg36m6ib. 3b n6333b
- 245 -
6ab. 4 £ 6 6i66nb qjmonmc8n Sypnb 6ifi5bnbi g j Bypnb 3ry;)6(jni(Yib bs66miC3 <3C3dilDi 0nD£|n
b6hiqjn ^hi6b3nhi(jnnb ghmb qjnouncSn Bycyib 3o9u3XfT'2>i nAC3^J 8* 9nbn SyQnb Sn^^Gityiion
bp3&:> b x Dojhm jbhynqjncnn (Lang et al., 1969)
Bycnb 6ip:>6bnb 5*Bm6*b6mh36*b g* nj,n bg^bi 9*9360010.

9y36*hnb By^nb 6*9*6bn 03Q0n3*g Q^hy^mbb 3*hymq}noi g* 9*93600) 96063-
9mhnb, 9*5h*9 bs^nhm* 5*3*hfinmoi 3ho>0*63oinb*5*6 b*60miCD b S ^ O
6*gm6*6n gb bi6j,hdQn3n 9* 9*0*^*636390 (3300936360. b*6dm i£j U3±pjdspm-
dnb ghmb Bn6* 39*6^3 0633b Bycnb h3go0nb h35X*bnnb bb3*g*bb3* 03j*6n‘b-
0o, 5*6bii3<j)h36no) 06o03630m3*6n* 6*533&ob 03j*6n<b0ob 0mj03936* (6>b.
4.26). 6*9*6bnb 3qjhn don3hn 5*9*^360 030608636* gpvib 5*69*39016*90, 5*6-
b*j>3cnh36no) 9(pnb* g* (p*9nb 0m6*U393ro6nb ghmb (6*b. 4.27). cnncnj0nb ym33Q-
030b, gpob 5*60*39m6*0o Byoob 6*9*6bo 3*h3bg36* (3gjhm g* 3qjhm bg3&* j*h-
ymojocnn) g* 0obn 5*6m6*b6mh36* (>936* 0bm9mg b*p*0rt)b *6 p*03. hmg3b*(5
6*b . 4.E7 jhaiiijnhruCn 63iC3nb

&i 3C3®:> 9 fowpflb, (33b 3obb g i
6<i3ei5nb 6y(vib 5mvj36unic'b3. 1^33-
pi'b] gngn gc>n3hn djhyjmiyjbn
3306036361 cjmcnmoSn, hmdjpnu
n9ymq£)&i gf>fib j,i69i 3Qm6i9n
1jh}6b3nhny>x WjhobSn. pi9rvb
5b69i3£VTiii3n (6y>enb djjn
9m6ij)33or)6n) 6ypnb &;<>s6bn bh>
C ie d i i9eo 5ig 6ypnb
3ot^36(inicnb 930(jnh;(i>i eCw 'nGPD
bif,hd6m6n bg3&i (Kozlowski et al.,
1991)
- 246 -
01306660b B ^ O L 603090
6ib . 4.B8 Quercus albab Bycnb hjgnSnb

gpvvjhn pn6i9ny> ( j )i'bm 9 n0n 1.3 p i 7.7 3
bnSipc^'b^ 6mpi5nb bQgiJnhngiB) JbnQjSnb
6ypnb 3m^36(jnsf<ni6 91 6ij,n b gnq^'bnjh
6n5ii(^993j|m6ibcni6 ip39nh9n (Hinckley et
al., 1978; l!»hm3ni)Xn bajdnbior^nb nb. Yoda
& Sato, 1975) Time of day
cgpicnQ3&9n Bypnb 6*0*6bn cn*6p*oi*6m6ncn 3*hymqjncnn 6936*, 0*on “3OI1330 9*6-

9ih36*b' 6y0ooi 0m0*h*536*9o 0(336*hob bb3* 6*600360 (0*5-, Johjnb 5*h36jo0*)
363336. *0 9hmb bnb 030608636* (03b*d03630O* 3 obn 5*^ 0 3*15)
6y0nb 5*(j*h36ob p(p03h0 153*036*91^6* (6*b. 4 .28 ). 0(536*11301* 6y0ob h^goOob
h3530*ijo*9o *h(jcn3 306093630^1 hm0b * ^ 30 3 6 3 6 *5(130133 6360*60 6*ymqj3&nu
(0*5„ ijOijh3b36ob 6*ymq336o) - obo6n bn6 jhm6 x *9 0^300360*6 (3.0. 5^153636
6y*0b) gi 03*1101^139360*6 (3.0. 00*3636 6y*0b) b*3hcnm 6y0nb h]go3 ob 83b*-
6*0nb*0. 00h*0 (1350^163680 (p*0ocn 6(536*113 33b *b3hb3&b 6y0nb 6*0*6bob
5*6m6*b6mh36*b 9* *9 n^m0 050 9(?ooio9(P3 3*h3bg36*, b*6*3 *h 9*9536* 0 mho-
£,n 63000*60 ^homgn (6y0nb 6*0*6bob b3<bm63hn (53*036*9^16*; ob. 6*b. 4 .37 ).
4.3.4.2 9l336^hnb Jl5D3i 6y0flb SPrbndfDhfl 6iC^6bnb

3 0 6 ihRD6 3 6 nb^013 nb: a^M&DCflhD&DCn a0:D6rrba 0 6 n
3jdngjhn hjftnm 6 jdnb 0i336*h33&o 6y0ob 6*0*6bob 5*3*h3b36ob ghmb *mhcn-

J03&*b *6(50113636 6*5336ob b3h30ob 8330611^133606* 9* *b3oi (9î3O)<i0) 996 rn*
0*h3m6*3n 0*cno b*65hd0O3*9 5*fi3h36ob b*93*036ooi. 3b Shm^bn 5*0mb*b30o*
6*b. 4.29 -‘b3: 9090b 630935 (jh*6b3oh*i50* 8150(1936* 6bm0m0 9pnb y33C*‘b3 y b x
^hnmgSn - 83*9c>nb*b; 6y0ob Q3qjo(5niyib 'bhpnb ©03639*3*9, 93*9Cpob*b 6*533-
60 116360*6 9*i3O)n0 095m3sh3R6*3o (*3 ghnb (jh*6b3oh*iiO* 0bm0m9 i3ijoj)3-
03hn 5^001 bnhi5O3093&*) 9* obb636o*6 6bri0mg 03mh3 909001. 3b 03j*6o‘b9n
b*03*03&*b *d033b 0^36*h3b 5**6m6*b6mhmb 6y0ob &*0*6bo. 6^361(106 6y0ob
6*0*6bb3 0h*3*09pn*6o bo03 h*0ob ^homgob <b35*3036ob 0*5*0001360 8my3*6n0-
n* 6*b. 4.30 9* 4 .39 .
shngjQn hj&nm6 j&nb 61536*113366, hm5Pihi5 6360, 5**660*01 60*9*580 ph9*9
- 247 -
Edcm rtDdoao
6 ib . 4.89 6nigi{,nb bntjrTyjnsnb 6^30365 j|h i6b3nhi(jnnb

gCpn^h pn6i9ny>'b3 (Stocker, 1956a) Noon Sunset
S*b. 43 0 Pinus radiata-b mh6gni6n

i(p6m6i(33536nb ,ihi6b3nKi|yinb gjnj-jniS^hn
gn6i9nii; a 6ycnoi 9m9ihs636xn; b Bypnb
6 9 12 15 18 21 9n6mg3i)nb 935y33^ngi5 8q-9 gcob; c 12 ge>3
Time of day [h] ] 6yomg (Kaufmann, 1977)
6sb. 4 3 1 biQbhÔ) biqjhiSijcnSn Quercus

ilex-dn bi&jQ&nb 3iJbn9iC3hn îS^hnsembnb
(gs) 9s gr°b 5i 59>3cm6>3n ta&job {>sbb-
6n(vn6nb Sjhnmgnb (At) EScrrjhn en6b0nî.
99hi(yi Sjhnmgnb J>i63h3pmdi3n (n3onbn-
baJ(j3363hn) 6^3360 3qjhm bi69m>,c3 9hmn<n
9> 6iiC 36ig ophoimg npjbi (Lossaint &
Rapp, 1978)
- 248 -
3(306i>(f)0b P aÔ U 6 3 3 0 3 0
Ri3i3i0n g3b30n> bnb^Qi, i6 Byonb biSihi^m jbm3n036n gi, ij39 i6 5i3m3g-

n6>h3, ih ijgoi ^hi&jSnhiunnb (93bi6i3nbig, C 0 2-6i<bcni (j3pnb) bBihiqjig gi
0dci3hig 930unh36 nb iDL5nC3^Cmî- 3^ J)i6 i 3nhm636b 93 hipn 3i 6 nljij) 36 nbig0 n
3 ioi 33533& x m&:>l3- hmg3biij 6 nigij>9 n Bypnb Sihi^n b]c 3°}hm g i 3qjhm Synhg-
3&i, 3b 9u36ih336rnj i0unli3&36 <jhi6 b3 nhiijnib &!&33&nb j,ibb6npm6nb bihnbb-
nbi gi bi65hd0n3m6nb bi03ip36noi (6ib. 4 .31 ).
6i533&nb Rii3iJ3i 5i3mn6333i 3013011 hn&n 3hmu3b36nb 3hcnm6pn3n 3mj03-
936001. bn99hi0dbib 33ng3h3nbnb ^h^mhn 3^3hg36i g i 3 nbn 3^3633090103!
ipih bgo6i qjmoiQnb Snjp Bypnb 90636001. 6y0nb mhcrrjQnb S ifip n 93qjni3nljnb
ghmb 6i&336nb Rii3(i3nbig0n 8^(132,0036! nt>hg36i (6ib. 4 .32 ). ^h^mhnb
fob. 4 38 Bidjnb i(^0m6i^35au>

Bn63363n pnqn'bn^hn J>i9(5ihni5n6nb iC3&i
jbnpjSnb 6ycnb 3ri|j36(}n:>(xib uspnpjbjbaiiG gi
i^SmbqjjhmSn 6yonb mhaijpnb p^grnyi^nb
bhp>boii6 (a - f) J)i33nh9n (Hallgren, yivj. Lassoie
etal. (1985) 9nb]e3noi) Xylem water potential |MPa]
6ib. 433 bi6&hd0n3n f)3bp3nbib

Bypnb Jm^fjufticnb 5i3C*36i s6b-
Ijntnb dgjjnb (ABA) yn6(536>jhiiyi-
nb {^'bhgi'bj (Wartinger et al., 1990;
Hartung & Davis, 1991). 3mp3(vi.
hm93cn(5 ibibi3b 615336^ yn6^hm-
Cnb 93ji6rfb9b Jbn(X)9nb 633680
i6b(yi'bnb dg^nb (A BA ) im6u36^h>
lini'b] Qi ajmcnpnb 6ycnb bîvKjbV) -1 -2
nb. Tardieu et al. (1993) Predawn water potential [M Pa]
- 249 -
paôi rtoaoao
a) b)
0 1—T—N 1------ !------ ?--

0 20 40 60 60 100 120 140 0 10*4 103 10-2
ABA iin xylem sap ( jjmol • nr3] ABA [ mol • m-3 ]
6*b. 4.34 6i 5336nb 5*0i5*hmdnb 9^ 3300136^ fyftQjdrb 633600 idb^n'bnb 0*p3nb (ABA) >*161336-
^h>(5nnb g^bhgbbo^ Q î39nh36ncn (Zhang & Davis, 1990; Henson & Turner, 1991)
3 38 [5011365 gi jjh3gnb 63360b ioiB^B^hsôob 80105^365 06333b cgm^mbnBcn^'bnb

n5^35bn3m6nb o63o6nh365b, hnb 83935595(5 C 0 2-ob h^^QnhQbnb 03j56o'b036nb
bhiinhQbSnhn 001^39365 5nd3036b 655336b 9inj)3lyTib. Bycnb 65056boi56 3 n3 ih-
0136530, 9i536ihnb 801300 bb^^pnb immhQn65(5053n 556^330^36300 060836300165
5j3b 0 ^ 336 0 9 56 55(33830 3mh3m65Q3h 95 303^01030^(101001503(1 bn56503£>b
[68], 6 n5055nb 550oi8 hoi6 nb Shoi^bob g56y 36nb5b, 0 ^ 3 3 6 0 365033 559 mymg336
q}n^m3mh9 m5b (56byn‘bn6 nb 03535), hoi030 o(5 jbno30 nb 03833016001 b6 h5q)5g 8(13-
6019365 030101036b 95 0nnp6335 65533&nb 03b5650nbn ( 1 3 5 ^ 5 (65b. 4.33 95 4 .3 4 ).
4 . 3 .4.3 e y c n b 6 ^ 6 b f i b a e & n a :)h o n 6 n b n e e n w iim h D b n
6y0nb 65Q56bnb 5500101305 93b50)0363005 9 bm0 mg 053 n6 , hoi93 b5(5 uocghmbhn-

359 550mnb5b365 836013090 95 501(101^036300 6yenb (15019360165. 63636hn3 3 n-
110163680 5b3ain 55lbrn0336nb 65^5(1365, 556b5t>30i(136001 qj3b336ob 0n3h 836013000
Byonb h5mg36oi6ob 5 ^ 0 ^ 3 5 , CJ630O 95 5(i5lb 3bli)05. 50n^m0 Bycnb dspsBbnb 959-
5365 b9365 shs3nhQ}3nhn djcnmgj&ncn - 6yoob 830(5330016001, 56 6yonb 3mo»36i5n-
50 nb 959536001. 6ycnb 35hymqjnoin 65056bn yoi330oi3ob 550 mnb5li)365 ^(ôihnb
836^(136580 95 0(3365hnb jbri3n0ob 6yoob 3oili)36(5n50nb 35hg658n.
6yonb 9T0^<-yim 6nb (535036590165, (101501(1(5 Byonb 65056bnb n6gni>5lynhn. cgm-
0103680 56 yomh^368n Byoob 338153300160b 33(10019x0 55^01835 &5056bnb 095m-
05 h3oi6 ob 9595360b b583503&5b od0335. 03010101080 Byonb 5^3503(10 838153300165
( W * ) 3695 oj63b 95y356o0o 1^695(1^30 ^09093893, hm030oi5 5b5b53b Byonoi
3m05n5536ob 65(133330 0950105(130165^ 3(101-3(1010 5b3oin bngog35 gjoioioi09n
Byonb 838(53300165 6533h360b 30110163680 ( W ^ ) . 6yonb 838(53300165 55lboi03nb
- 250 -
90 D6 ifn 0U ffiaaoan
0 fTHj30x 9m036^9n 33 ndc3£>i ft^mnbi^mb hmj,mh(3 Byonb qjihgmbnom 83815-

330m6i (WWC) [ 169 ], 550mbibxn 5hm|536lj363n 6ycnb n3 339 u330fT'&nes6 ,
hrn33 CnU ihnb 5533(1361^ 3nhm&3&3n:
RWC = ( W ^ / W ^J.lO O (%) (4 .13 )
Byonb 6si)036m&i, 563 Bygnb goqjnyn^n 559mnb5li3<!>5 Bygnot ^jQhpbnb qqojiv

Un^ncn (WSD) [23 7 ], 3b SihiQ^hn 530633636^ 013 6yenb hi himg36m&nb
93q)nun^ni jbm3 np3 n bh^cê 6^33^3630 99 &m3 ih 3m&ibaii6 33gih 3&noi:
WSD = ( W ^ - W ^ ) / WM . 100 (% ) (4 .14)
U3On0 D&3&n 6ycnb 3mvt)36i5nic3 n. ByQnb 838i533em&nb Sohy^nbi 9 mj93 Q36b

33h3Qnb B336nb ^61536^115150565 g i ^h& m h^. Bygnb îhymqjnonn 65C56bnb
ghmb 5mji)36un3hn mbamb^hn 66335 (T ^ ) 9 i^ 30 m6 b. y33C5 939016333580
Osmotic potential [ M Pa ]
-3 -4 -5
Hydrophytes, helophytes
1 Geophytes
1
II •' ^ Meadow plants
9f
i Crop plants
Herbaceous mesophytes of wood lands
Herbs of dry woodlands
1 Deciduous woody plants of the temperate zone
■ Evergreen conifers (summer)
Sclerophylls
Steppe plants
■ ■ ■ ■ Desert plants
Succulents
Salt marsh halophytes
61b. 4.35 9(536^300 3impfn;>n3h5e J>i6bb3i 336x iyi33<b3n Bypnb mb9mb3hn bngnp33-
bn (Walter, 1960; Sveshnikova, 1979; Nobel, 1988)
q p ij^ n . hm 3 vF 7t 0 1 5 3 0 3 6 5 5 h 5 3b m o m g 6 y c n b 3 3 8 15 3 3 -
5 5 b ic n 3 5 Q n b 6 n 6 3 6 3 c n 5 n b
gm 6nb 1530nC 3^>nb 5b , 5 h 5 0 3 g m b 9m h 3530i li m h x n 3 h m ( j3 b 3 6 n b (3 :> jh 3 & n b y n 6 (53 & -
j)h 5(5n n b 9m 95(j 3& 5; 3h m g n 6n g i n m 6 3 6 n 3i i 3m c 363n ) g h m b iu . h m ^ m h ^ & 5Q56bnb
n 6g n i 5^ m hn, g i h g 3&5 m 3,jn 8:>c3h b n g n g 3 b ( h m t fi B y p n b 33OT3n b 3&:> 9 5 iphfc-
- 251 -
PbCJOb 603090
3> gi6>c>6b32)Xni) Q* 3ijbn&j9b bn30h:>0nb iOX^D dQnjhn z>$3Qy

6nb 3nhm6o69n (6sb. 4 .35 ).
qgmoiQnb Byonb 3m^36 un>on (T,) ôghm 'b^b^n bngngji g:> 33301 >b>b>3b
Byonb 6igi>6bnb ( 5 3 0 0 0 3 6 3 6 ^ 3ngh3 mb3mb3hn 3fTili3 5 ^ n ip n (6>b. 4 .3 6 ) . Byonb
6 >o>5 bnb 9id^63orn6> 36^033 nB333b ^h^rnhnb 030ynh36ib. >0 ghmb jbm3n-
03&nb Byonb 3m^36un>cn 3gjhm RJihb 31530^. 309113 mb3mb3hn 3m^35 (in>on.
H am m ada s c o p o r i a
— Water potential
— Otmolic potential
61b. 4 3 6 jgôbr'b 9(536^300

qjmoip3&3n 93hipn Sjhnmgnb 6mpmU
6ypnb Sm^jfiiyiicnbi 9i mbdmb^hn
3nvj36(5nionb g<n>e>idjhn l53CnC3Jl3-
6n b3cm36]hn 6mh6y3nbi 9s 6363-
6hn3 3nhm&3&3n (Kappen et al.,
1976). 0m6a(53036n ^hm3nixn
3ihig963i 63 gi qjr»Knmp0i5336n
b33&nbiOi3nb nb. Sobrado (1986);
3ih i9d63i 63 Sjbnbioqnb d3<bmq3«c-
* I 12 16 3h p i Jb3hmqjnQ3h 3nhm&3&3n nb.
T im e o f d a y Rambal (1992)
4 .3 . 5 e y c n b h 0 3 n a n 0 l5D6 :)hDOD b b 3 ^ G ib b 3 i m obdn
4.3.5.1 snehnb^bficohn e:> 3nehmc:>6ne:)hn 6ycnb bxstbn
<335 jiin3hn ju33nb> g> 3^6 n ^jn b >9 mhR33nb 0 nb3Q3noi, 0(536:>h33&3 n j,>6nhR33>
Byonb 6iQi6bnb mhn 3m6^hib^3on 3^hru>6v5n: ZnghmbtfsdnQjhn p i 3nphmp±dnpjhn
[234].
Znghmb&idncjhn bib3m&36n Byonb 6}0:>6bb >Bni5ib6fnh3635 By*00i,i0ih:>536nb
03hy3m6nb png qjihôs^n, (1:195^6 9 ioin 6^53360 30*033 h3S5nh3&36 Byonb
030y33Qm6nb (5300036:^3 g> 6iiR6ru>oi 6n:>g:>50n ph0ig Rb0 i 3ion qj3b30u> bnb-
^30i. 6:>0i6bnb b^>6nc3hm6>b b3Qb 3Bym6b ^ 3 0 1 3 3 Byonb 6^33330 0 0ihbj,n
b>0>h>5fn mh5>6n36b>, qj3b336b>, 03hjs6b:> gs 'bhrob fohjOn, hm030ou>(j Bruig^n-
3(536ih3-blj0rtbq}3hm b n b ^ S^ n i^nbhnicn Bygnoi 0m3>h>536nb 0n6i5*6n 5>6m6sb-
Bmh36nb qn&lunb (>5 bO<n6 Dh 6h3g0n 3m6g35bivjmh36nb 0bj>i3big. nb. 6ib. 4 . 10).
3py)36yniC3hn mb0mb3hn B633nb> g> qjpioignb Byonb 3r»y!(36unionb b3'bm63hn g:>
gp3-(pi03hn l53CnC3^);> hR36> bi30img 3nBhm 03ih 5036311. >3 ,jn3b 3330i36n>6 6 yiQ0 -
I536ih336n, b333036l536n, Rhgnpnbi9 ^ 6 n 6bnh6i0ib36n, '601503(10*1 3 ih(53Qm3>6 n p i
- 252 -
h35nm636nb bp 8(536^3360 (ubh. 4 .5).
InphmQsdnQjh b*b3^163680 Bypnb @30^1130,33* 8300(036* 5*3mo63omb 6ypnb
gngn h*mg36m6noi 3*h53* g* 3jh3gnb 6336nb Jjm6y36l!)h*unnb d(vi3hn 8rn8*,j36*.
*b3onn 0(536*113360 b*30*mg ^)mp3hi5 (j 30n *hn*6 Bycnb 3m,336(5n*cnb dpn3hn
03hy3m6nb* g* ghm36noi g*£i)6m6nb 3n3*hcn*(5 yi (jjhn^nphjcn b*b3m63&n). *b30i
bib3m636b *;)30i b6h*qjn *(j<g536nb 36*hn, h*g5*6, B3bn, 5**fi6n*cn b3Q-
b iyhxn 33qj*hg36*: q)3b3n / 9n6nb‘b3gi 6*6oQn g i d ^ h n Bycnb 5*3j>*hn bnb-
(!i33i. 3nghrrici6nc3h jinSb 5*6333013636* &*c*bm3*6o 3(536*h336nb gngn 6*6npn,
b^S^&nb 0*h15300136360, *5(130133 b3 0^36^3360, 5*6^3301(136001 3nm63hn b*b-
3m636n. dcn3h 3nghmc*6nc3h36n *hn*6 3 mnyicm3nghxn 8jhj6n*6n 61336^3360,
b*3b3&o g* 00:136360.
libhnpn 4.5 bb3i@ ibb 3i ^ynpfn^njhn s&jqjnb 8(j35bhnb ib n 0nen 5n3h mh5i 6m<ni 6n6 n3:>eohn 6y(vib
3(’M in io n (MPa) (Richter, 1976; Doley, 1981; Nobel, 1988)
0(j36sh3Oi> J53<2n
-1.2
bicibm 3i6n dj'bmqjrvjjfcn - 1.5 - -2.5
3ihu 3(yn3i 5n ^ 'b m q jn jjb n -2 - -3 (- 4 .5)
ljhm3n ix n 63(180*6(1 (jynb bj<}(536ih 33&n - 1.5 - -4
bm6n]hn bih^yQnb 03h;j6ni6n 9uo5ih]36n
cgmcnmp0^3')6n bajbn g i 636^360 - 1.5 - -2.5
3^19063163 b33&n - 1.5 - -2.2 (-6 )
S jhnm gxig 39hi(yi h;){>nm6;j&nb 01536111336(1
bip^hmqjnp^hn 8(5361113360 -3.5 - -5
ihnQ30n 3i 6n £ il!>3&nb 63^^36(1 -3.5 - -8.5
b03(V7Ui03i'bc'33Oinb 6ib33hig63fij36n A - -8
^3hmgnj)36n
3gi66(nb 63^360 -5 - -fl (- 16)
-0.8 - -2
3i 6{,hnb 0(5361(13360 -5 - -6
3i(ynq)n^36n -3 - -6 (-9 )
4 .3 .5 .2 eyoflb 6 ^ 6 b n b JjPiebibniMlSfDhfl ibfl3 D£>n
6y0nb (133080b ^ 30 10 0 150 3(10 93g*h36ocnn *6*0 o“bnb b*30oib3& n n8oi*3noi33

oyoi b*33C 3 3nhoi6368n 8 i3 C 3 3 ;>hcn* y 3 h *g p 36 n b b *5*6n. *8 1)3301*3 3 n ^01503(1010
b3h*oinb 038 330 16 00 1 5*g8m i3303cn nJ63&* *g h 3 R*li)*h3630n j>3C33*-dn36nb 83g3-
5360.
03-20 b*3i36nb 83* 6c36nboi3nb g*5hm3g* b*j>0 *mg gngn (3mg6* y330i 3Cn‘
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b363h3&36nb 03b*b36. 83118036* ocnj3*b, hm8 ^mybsc *hb36*cn* 8(1*3*0033(101363-
6* 303(101 gngn*, 3ngh3 biboijm^bcm 5*h38mbn: 3hcnn h35nm6nb b*b3m6*oi*
b33jlj)h9n 93n6n8636* 3* 6 n(i*,inb*g8 n 9353360b 3*8(1*30 3*hns6^n g* 3018606*130*.
- 253 -
pa«=job fnoaoao
50iboii 6 ghotiQ, &ihi333cn bib3m63&nb, 535(13&nbi 0*3 mjib^nb 5363^1331159

<gn;)bnh3&3pn cnb3nb363h3636*1 (16365 3(530X n- m- QiVîOhnb (0 . Stocker) i'bhncn,
3b 5 3 in d x O ^ j)hn^n 1)3059 559535301660101 6(336ihnb y330i nb ai53nb363h3£>5,
hm030nii n0130365 0m(j3030 356n^5^3n 0nbn ihb3&m&nb 3301001503(159 06n0363-
001356 Snhmbig. jli33nb oi53 nb363 h3 6 3 6 nb 3(15350033(1013636530 0 5 n6 u 03bid-
03&30 ni 550mnymb 33*1103^^0100^31159 5i6bb3i33&3pn 30^^0^31303(111 ,303360
[ 121 , 23 8 ] gi bibnumybom b^hiô^bn, hm003 &n|3 65^501936056 5565,3016363-
0n 5ih30m 3nhm636nbig3n h3ijunnb 3hoini6m&ncn g i 3560,35^0156 3539nhnon.
^3380101 556bn030ni 05550noi36 nb 0o>3cn hn5n.
b3 0 (j35sh335n
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3nhnoi 95 03363360956 <301010365093 gngn 9 i 6 dnonoi, mh556o‘b5500b nb3om cgoih-
0 nb 05550noin5, hm030bi(3 3333 i(?5h 333d0O5 Bypnb gngn himg36ni6noi 35(1535
- nbn6n oii3ngi633 n0ymq}3&n:>6 5i36m6ib6oih3&30n Bypnb 6i0i6bnb 30(1016360*1.
0 bn >6 50 n6g3 n 633b b3 0i53&ih30n 03ig(r>nboi3nb Bypnb &5056bn 35hyoiQgncnn5.
3b n6333b 6i533&nb ((1010036013 b33&nb 333ii3bm650n nb3giu 6y0nb 956500
5533h3&nb 3nhm6369n 038501636) 0n3h Bypnb 5oihavjo36ob 930 ^11365^ 8015-
3ni63&noi, hoi03bii3 6y0nb 65056bn 5i6m6ib6mhg36i, 655336n 0(^6365 95 ,3(156-
b3 nh*ijnnb n6^36bn3*n65 05^ 3o*n6 b (65b. 4 .37 ). bnb 35hjnb y3305 6i6n00n ,3(156-
b3nh5^ni 3(1019(1013059 ih 0130(19365; b3nh59 33663(1016 qgmoi036n h355nh3636 y33-
05^3 53ni6. b3 015365(13015556 5i6bi33aih363pn 3nghoib^56n03hoi6ncn 550 mnh-
R33ni6 ‘bm0n3hn bih^ypnb 6n63m356n 95 Rhgn0nb 50^ i 6 n b5b30i636n. 50533
ghmb, ihb3&m636 b33&n (0i5., 003560, 15(13 iûjni), hoi0036805 6y0nb 6i0i6bn
'601533(1 i(T^33b dpn3h 3ihyoicgnoi 6 n96x b -
b3C3hmqjnp3hn 63 8036^3360 g i 63^36*1

n0 h35nm6360n, hm030 oi5ai3nb5i5 gi0ibibnioi3630ni <biqjb30nb dpn3hn 6350-
35 95 333n6mJljni03hn 6i03fo6n i6 t>50cnhnb b3 nhn 63n036n (b030ot503i<b(Ti33cnnb
h35nm6n, 3i3nb 3hoi3n6(3ni 503(103530, 2>50nq3mh6ni, Rn03, 53b^hi0nnb bi0bh3cn-
gibi303oin gi bb3.), 56h30i33 ^hm in^nb b30 nihngx hinm63&0n 553(11330363-
65b. 437 ^hi6b 3nhi^nnb

gprrjhn 9*1658035 ij^nb^mb
9bns6 b9n5g3n. h rn g ^b iu
Bycncn 3m35h55365 5hs5g3}-
35£)hn bg3&i, 3nh33Qsg
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9jjnhg3&5 i 366h x
y3X i<b3 6565013&X 3663680
Time of day (Pisek & Tranquillini, 19 5 1)
- 254 -
8096 d6 0 b Euôt. fooaoao
90* 9|3o6bh3x ifT'^r^ 6n^n (Q^mhicn: 0*fio*, 5*11053, R*3*h*90, qjo66mbo) 9*6*90
,4360b b] 9u36ih336nbb pi 63^3606 9010060(136001, h»n039Oi*(j 5**660*01 0*h*9-
663463, 303tiobsg b3030n (biC3hm0mhqj3Qo), 3*^*h* *6 b*03*9m ‘bmOnb 09010103-
6n. *0 0u36ih336b, b3<bm63h*9 0)903(1 03*035*90 yyi0i^nb ^0(10163600, 5**860*01
9*6*90, 0*5h*0 6*65(109030 *bn0o0*uoob 36*110. b03ooi*03*,b(n33oinb ,jo3ob bj-
03009010190*60 63&J360 bj^n^hn ‘bhgoboqob 09363636 ^3(100196 030mp5n0ob 630-
93609*6 *9h3 b*qjbx*093- hm93b*(j 60*9*506 ^360 b*^0*hobo*. 0 ]* 9* 530*6
lb*<5bx0o, 6990b 6*393601606 5*001, 0*cno 03^*6mQ3ho *^403016* 0D<bcr393cn*.
b3030gmcnQo*6 0(536^33600 36393600 hmftmhy 3o9hmbljp6oQ3ho, *6333 3og-
(1019*6093110 <336Jjom6*CDho (jnSnb 6*h0m0*9536Q3&b. 0*5*9001*9, L a u ru s-rh ,
A rbutus-rh 9* 0*11*9063*63 Q uercus-ob b*b3m636o 5*0010(16330*6 ^h*6b3oh*uoob
Shmu^bob b36bO|)3hm6ooi - obo6o b6*qpg h3*50h3636 6990b 030ijoh36*,b3; 03m-
h3b 0bho3, O lea- b, PhillyreaK j, M yrtus-b 9* C era to n ia -b b*b3<n636o 6990b 030|J-
339m6nb l}3*936*9m6ob 0o0*hoi 303(101 ©3^0 5*0093016001 b*bo*oig36o*6 9* 5*(jn-
936001 530*6 <bc>39*336 |)h*6b3oh*(50*b. (101501(115 6360, 0*oio 6(190906 0501019360
6*53366 303(101 9096*66 (>>0133636 5*66609 095018*113016*00, 3ogh3 0bob*g0o
6ib. 4-J8 bijQj&nb j>i3^)ihn-

i6mdnb, qjmcnmcSn Byonb
qjihgri&nain 333u3X>,'&nb 8*
cjmcnonb Byonb 3m^36|yiionb
gpn^hn pn6i 9n ii bnynons'b]
ft’i'bihp 0(j35ih3363n ‘bygb^-
ps9e3 (8inbn) pi bncBhionb
Jjhnmgnb dodpDb (b]ij3963hn).
ZTP 6snb3 n ^^h^mhnb 6 x (’i3i6n
Bjhyjnfn (6y3]^non bi'bn) g i
bjilD^D^n (]6y3>yi bi'bn)
(LoGullo & Salleo, 1988) Time of day [h]
- 255 -
ffa c m rtoaoao
pjb3m6 nh36] 0 n S^hnqjQhn^Qn qjmai036n.

9U36sh33&3n Bycnb 330|j33pm6nbb g i Bypnb 3mli)36i3ni0nb 03hy3m6nb bihnbbn
g iS m in g ^ x n i ihi 6bm0mg 6i{>33&nb hîJjjnnb 36^1^3, ihi63g 9 6n 336 x fT13556n-
0ig - 033b3011 bnbvj)30nb 0 1^ 3^ 36 ;^ 36 ^ 3 gi 55630015(1361^3. b303hmcgn03hn 63-
fij3&nb 303lt)3bm6i3n, hm0C36nu 0iOKn3nb b3gbiyhx 3 1 6 0 ^ 3 6 8 ^ ^63001^36300,
'bicgbxnb 535030b 6mpm9g3 rib8mb3hn B633i 3U30i 2-3 MPa-093,
bmQm ih ib 3cbiyh3 0 9 9 hi0 3 i 6 nlj i li>3 6 9 n 6(336^300 6y0nb g3qjnijn^n &33hig ^qjhm
gngni (6ib. 4 .38 ). 3b 5^6^3301(136001 gi 9 ibibnioi36x n 5 33ho3ngh30n bib3m63*
6nbioi3nb, hm0036ni5 ij ^ r ^ i g 36n6ii(j'8g3536ni6 5i9m9hm6ib (nb. 6ib. 6.68).
bi6i0 qj3b336n Byigb n6m336 bi^dihnbri himg36m6ncn, ^hi6b3nhiijnnb 65319b
gngn 660336300165 36n^3&i. y330i b303hmqjn03h b3-6u36ih3bi g i 63ftjb 5556605
55 q)i6 li)Xn Q5 3n6hm ^h^036nb 9Jm63 550îho bnb^30i, hnb 5i8ri(3 336m3nb
diQnb d0O3h 9m9i^36nb 936016333561113 30, j>i3nît5nnb bi3n9hm36i 9(30(135.
Ibf»i0n3hn bih^ycnbi g i (5030 hQ^nmCqbnb 333s 63^^360 g i 659085 6(3365^13360

li)36ghnb 333i 636^360 g i 6ipi06oinbi g i ihj^n^gn (135011163606 6i0n3i 6(33-
6ih336n 553^3036300 ihni6 pni, jihnbj,i>6 g i 3i3330 3i6n^^363n. i6 ,yi3nb 6(33-
6ih336nbion3nb gi6ibibnioi3630ni: 6^nghm 95^1^35; bn3h(j39n 3m635j^}hn 556-
055365; Siîhi g i b3 nhig gib3330n, 33h(30nb3&hn i6 636bnb36hn cgmoi036n. y33-
0 i t 3 gi6ibibnioi3630n 6n3i6-oi3nb36i 30 ihnb 6 0 6 ^ 3 6^nghmg 55hcnb9x'n
bhgnb cgmh6i. ihb36m6b 0(3365(13015 ^03360, hm030Oii 5i3h(53036i 33 cb(j'3g x ni
ih i n6g36ig 6ioin 3nghmq3n'bnm0mj)n3hn oii3nb363h3636ncn, hi3g365g5(5 bncn63hn
h3gn6noi, biOoiihSn aim30 nb b5&3h3300b bo05(pcooi g i 60595580 i'bm^nb bij)9 b-
hnbn Sihi^nb ihb36m6noi. 55(138™ qij^nh36nb ib30in 6n6ib6ih 3 m(530x n 3m0-
6n6iijni, jihnb 5i3036nb J330 0ymqjo 9(336ih336nb 6nn30o6i(jaii6 3hoiig, 5^-
gi66y33v!) 5530365b ib@36b 6y0nb h3gn0t>3.
5 ihoib0x n 335-5 636^360, hmj,mhn(3 ihnb Loiseleuria procumbem, g i 83593-
h30n 0(336^3360, 855., Salix serpyllifolia, 6nigi&nb lb3gi3nhcni6 bnib0m3nb 838-
33m6noi, j06 ni6 5i6m6ib6mh3630 030^013008^^ i0 bib3m636ob 6553015 iSihi^n
qjhnig 85hd6m6oih3i <y>9 m83 hi0 n 3i3hnbig6n, i6n^m6 <3^10193606 55)138018(5330
(03^1019360156 303i0m 33b36nb 8jm63) 3i3hnb <336i3o (i36ni6m6nb 6m6i^36nb
558m, 6i533&n nbb&36ni6 bi36im b6noi, hnb 5i6m(5 5(1(5013 6i(pign ,jhi6b3nhi5n-
nb 3nhm6369n 9 ioin 6y0nb 6i0i6bn d0n3hig ih ndi&36i.
9 bj)i3bn 3033^0, hm83gn(5 33 nd03&i i(no63hmb hm^mh^ 8ipi0o i3hmgn6i0n-
33hn gnqj3'bn3hn 6n6iip6g35m6i, 33n6n363&5 659085 6(336ih3369n. i6iboii6
3hoiig, nbn6n 333<Ji3big 6ihij,g36ni6 6y0noi, hibiy 3<bh36330yrnqjb ph6ig Ri9 ^-
3i0n dnhnoiign <33b3n g i (P3hm36nb 06^36^3(10 giqj3b3ni636i 6i0n6nb J339
g3^hnli,nbi gi 6^3hnb gi5hm336nb bn3h(333n, bigi)3 ^360 bi36ihnbni. 6ib33higb-
q )3h xn 6 i 0 o3 i 8(336ih336n 6y0nb h3go6b i363m63b3636 ^hi6b 3nhi(3nnb ^>39^-
3nhnbi gi 6yonb biQihi^m 3 ngi bn3h(3nb b30b iyh xn 33<3ihg36ob bih3’b3-
63'bmq}nvj3ho 6ioh6i0ib36o g i
330ng3hn h35om63&nb 6i0ibm3i6 6i336ih3363n 63b3g36i 6y0nb h3gn6nb
ainoiJ6nb y3305 J>n3n g i 3ihni(5ni: 3oghfnbî6 n03 hngi6 g i 6 y3& xn g> dgn3h 3np-
hm0i6n03hnai gi6o ii3h 36 xn- 6y0nb h3gn6nb ^n336nb ibgom 6hi3i0q33hm3636i,
33nh3303b ym30nbi, j>i63 nhm636 x n i 6tni6n 3i6nî(j36nb ihb36m6noi, bigii3
03330136^ Boigi^nb J,i6 m3 hm6 nb ^36g36(30i (6ib. 4 .3 9 ).
6y0nb bi36ihnbig 6m6mg36ob 3nhm6363 n d^fbmqyvubnl) 0)hi6 b3ohn3ni
(imhoij036nb 36ihnb 33bi6i6nbig) 5oi30 n gpnb 5 i 63 i 30m6 i 9 n 333cb(j'3gi3ni.
- 256 -
9i)36 ^ o u Aoaoao
Time,of day Time of day
5ib- 439 <i)lu>6b3ntuunnl» gs mb3mlr)hn 3m,j36(jni<v»b g(W)hn go6i9ny> Jbjhm oijhdx'' 3i6n(js^nb
bb3ipibb3i 9^36^ 380, 39hi(vi Sjhnmgob Qibi6ynb3n (0ityb 36i b]hicnn) p i 9-g(nru6n 63^03"^
(3ih j336i bjhiori) 930934, (Muller-Stoll, 1935)
^hi663nhi(5nnb Shmu^bnb i063ihn 6ih0ihoi3i 033d0nioi 3bmpmg n0 8(536^3366,

hm0o36bi(j j)iiR6niOi 6 nigi&9 n (ph9 ig Ri8i3ipn 0336301^ bnb^30i g i 83663301
36shn 66hiO)ig 0 ni6 mgm6 Bypnb bi^nhm himg36m&i qjmo^&b, i6 iQ3nQiQ n^>-
636 bn99hip3b. i 3 n6 oii69 g33n 96n9363pm3b6n 30)3^0 Og&mOih^m&b bnoiftmb
h353Qiijni3n, hm83pni5 8nnp633i imhoijp3i>nb 3hmi53b9n 6b33pn6 5:^ 336 0 0 1. i 9 n-
iim0 ibgoin 2>ni5l>o>nibIX5nDhn ^nin jSn h i^ b ig 5363936^ pni gs 15630 3i6nli(i (»)3&3n
(9cbnb 0i(j>i0n hignii5nn6ig8n 3j63m6 nh3& xn qg3hgm636n). i8i6ou>6 3hou>g, ihb3-
6m6b bb3i < 3 3 6 ^ 3 !™ (j>m6bl5n(i)3(5n3hn) (i>n3n6 8(536^3360(5, hm8p3&ni5 03ig-
pobib i6 gjnnb 03mh3 6ib33ih9n, 6y0n6 6ipi6bnb jihymqjnoin 06n8363pm6nb
ghmb, 3 3^ 3 36 6^53366. bip>i8mb, hmg3bi(5 6y0nb 836m3ibi g i fti(j30i0n 0nnp-
633i gig3&noin 6iQi66n, 6ij,336n nb33 nbb636ni6.
0a(psCîOab3X mî ',> Bypnb 936m3nbi g i 5 ^ ^ 3 6 0 6 oii3n632>3h3i>36n6 oi3ip-
bi'bhnbnoi, j,i0mhR33pn ig&npn 3j>i3ni. mbm^b <bp3nb 6ib33ih£j6d303&lb3 gi
►i36d303^<b3 g i ^ r ^ i g igj,np368n 6i3huxD^Xn hi0g36n03 03^hnb bnOippnb
5hd3oqj3b3hni6n 6i0ib36nb (0^5., H era c leu m lanatum , F ilip e n d u la ca m tsch a tica )
0>hi6b3nhi|5nnb n6^,36bn3m6i bigbjonb 0bni6 ge>33&8n bi03ipfTig 0(536^3^3 &i-
gii62>ihn336no) ip633b 2 gn^h 6yi0b. fcOfc^^hn qjmoipjbnb 0Jm63 ©(536^13
P eta sites am p lu s g i bo3h(539n 0dpi3hig &i3pnpn c,3y6fn>jhni sa c h a lin e n sis
imhoij03636 5-9 pnli)hi093 6yipb [14]. 0n]b3gi3ig n3nbi, hm0 i 0 0(536^300
qjmcnp36n Bypnot &i6bi»Q0)h3&noi 0gngihni (63gpn 6m6nb 90-94% ) gi 6nigig-
- 257 -
f f a s m Aoaoao
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Sndhmajn^^bb, hm003doi3 0^9360^6 ^ 33^380 g i ,i)36ni6, Rhgopoi6 3i 6 o,ji-
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6013 3 bnb Smdhimbob 3ih i 0303 hig 5igio6i|3303636) 5i3m 33^86000 dpn3h I33i 03 -
6igo 5i6iOi36xfT>2>nb 3nhm63&9n ibiboioi3dcn 6^533606 ib333 (33iQ36ign 3mj93-
936i. ghmnb 3 (T>i>03 3 m6 ij)330 i9 n ^hi6 b3nhi(3nnb n6(j36bn3m6i 33 hy3m6 b l/ 3-gi 6
l/5-3g3 [55 ]. 3b 61536^3360 bi6fthd0n3 5i6ioi3630m6ib 33h 0 ^636, hig5i6
6i5336ob bh3Q0 gib3h3nb 3 n3b3gi 3ig, ^3^0^3031111 ,i,hi6 b3nhii30i 3 i(j>i0 n hR3&i
g i q33b336nb b^bô 5i63noiih36ob 5i3m, Bypnoi bi^Sihnbig 3<bh36330ymq3i ih
bg3&i. iSn^mS dpn3ho imhoij036nb 3nhm6369n nbo6n j^nûnnbigSo ihoi6 3 og-
h3j>nc6o (5i6bii3oih 36001 I33i^036b-0b gîhnb bib3m£>36o).
igh3j>i'bigb30nb j)3roq}n(j36b 6y0ob P^gnSnb oii3nb363h3&ob 0ob3g3ooi 33>
30ioi bi33i0m 3g5(Ti0ih3fn6i Rhgo0nbi g i 3 boi 6 o Sibo^^bn b 3(336ih30ii 3 mhob.
obo6o 303(101 3nghmb(î6 o03hn ô3nbi36 ihni6 5igibhnpn. b ^ ^ b o b 5301030,5360,
hm0036ou 6oi0i5nb 6y0ooi bit,3 ihnbig 3‘bh3633cymqjnb 3nhm6363o 3noiihg36ni6,
ihni6 3nghmbvji 6 n03 h36 n - ibibniai36oi ^hi6 b3nhii5nob gibigo n6(j36bo3ri6i gi
qjmoi03 &3 o Bypnb 333 (3330m6 nb 336033630m 33 hy3m6 i. Bypob 3 i(j>i0 O 93030130-
ijob 5i63noiih3&i3g3 obo6o q3moi036b 3 3^ 3 36 .
8^13300136360 g i nbo3&n
6 i 0 ibm3i 63 i3 (336ih 3363 i 3301031503!™ 5i63noiih36nb 3hmi33b3 o gio 3yh3b
bh30Oig 5i6bb3i33630O 3^60^,3360: gi6y3630n 6y0ni6ngi6 6^33360015 gi
3gi66m36noi g i 3 oii3h3&30 o. i 3 ib 333bi&i33&i 6y0ob tgyj^mhnbigSn 3 ioio h3iji30-
360b q336ji503hn 3hi3i0qj3hm3636i: ^63016360, (1018036015 bih3i336 gogo hioi-
6sb. 4.40 bsbi(yi6nb ijhi6-

Wnhiyni ps^ftmhne pi 9pn6ihnb
Juty August Snhgjc (johibi'b] (Morozow & Belaya. 1988)
- 258 -
936016001 6yi0b ^ 30 136 0 4 835., bi 6 i 3nhmoii 946490 imhortjQQftnb
6jm63 8(336^3360 (S^Qbnb^b^h <301010360460 04^300136360); bib3m&36n, hoi003-
6ni5 6453360b 6013033490460 54bb64-04b3h3ob 03033016001 41135x0^13636 Bypob
h3gn9b (64b. 4 .41 ); 451130)33 bibQmbQ&n, hm0036ou 4hn46 309(10104600311360 94
33hn3n9h3036o (54^4330^13600 C4-0i5364h3o&n). 633I10 04^530013460, doh^mhob
943411530b b434b3bmg 40|3oh36b ^h46b30h4(500b 'bogiiohb qjoioi036ob 9433(531^4
94 9463330b b4934Q36ooi (64b. 4 .42 ). Stipa tenacissima-dn 3b 03^460^300, 546900
<301010100146 9 394b36001, 3<bh363xyfTM3b v4h45b3nh4(jnnb o6|j)36bn3oi6ob 40%-O46
030(5oh364b [205 ]. b43463&ob 84^53001363630, 4b43ob 04lj364bai46 3hoi4g, 64533-
60 341154336 0mj6n0m64b 94 634930b 33hooi0ob p46y36nb40)3nb 8(5364(13360
6y0ob h3jo3b 33C'4h 43016^10103636: 6049450 543oi33li40o4, ljh46b3nli4Uonb 3hoi-
l53bo 30 5hd309364, b4640 <301010360 b46oi0oioig 4h 5480^36046.
Bib. 4.41 03hi(yi b^]3nb du^64hnb Stipa

loannis-nb (jhi6b3nhi)yii 63iC3n*J i 8*
9mhBy3nb 5nhro&3&3 n (Rvchnovska, 1965) Time of day [h]
E
0CM
1
O
E
E
*
&bb. 4.42 Orysa sativaAi qjmoiQQ&nb
@ib3330b sgoijn (jhs6b3nhbons'b3 jjshm
gi jihniS (2.4 94 2.8 3 6& 1 ) i9r>6g3n.
gnâhi3nb j33gs dbihjb 648336360:1
gmcncnb gsfc>33311b bb34g4bb34 Smfenijni
(O’Toole et al., 1979; Hsiao et al., 1984) V 0
- 259 -
pa^n b (noaoao
3gi66rob 0jjQ6ah336n
9l536ih30ii bibn^nijbcm cgmhdgbnb 61153500^0136365 &56bij)30ih36ncn 90905
395&6 m3 n: 0 uoh3 <bm9 nb 63^360 jghijpnb^bhn 95/56 Bycnb bi9 ihi&m ojmaio36001;
63^360 , hm03pcn5i5 555660501 b^jjXDîSDhn 95 5bn0 D05(jnnb 36ihnb 9 jm63 j^hjob
jbm3no36n; 6065^3 &ihonb03on 65pn05 0tf36ih336n g i 6ib33hig6;3fi;j3&n; q^b^ho-
560 0i(T»i0650ib330f,*6i; 63^^13560 0^1539/113636(1; b3i3936(i36n; B3(i0nb qjprnhob
3q3303h30d ^3hmcgn^36n 95 bb3- (nb. b^h. 6 .6 4 ).
'bmg^hcno jq jjd jh n bib3m 6i imhotjc36ib b^h otn g ih 59(jnh3&b g i i 9 3hm-
(J3bb &i6i&hdm6b, b i6i 9 ih 5i 0m9h36i. 9h i3iQ6cni6 0(336ih33&9 n 550moymqji
mhn, ghm0n 3hcn0i63cnnb 0n0g33ihn jtf33nb cni3nb363h365: 3nghrooi6nc3hn q ji b i
î'biqjbgo'bg, b i6i 3 6ni9ij,n 33b $0933 &)6ni6ni g i lbiO)b30nb q jib i, hmg3bii5
^hi6 b3nhi|jnnb 05jbo05O3hi9 03'b(j'393nb 03033m6ncn bg3&i bnumubcnbio^nb b >
9 n9 n 550m9hm6nb ou>3ngi6 5(309365. biOonhnbi g i 5ib5q)b3cnb B30dnb 33hnmpnb
ghmb g i 0nb 0309355(3 0(336ih336n 3^359 0 0 ^ 5 3 6 3 6 6y5pb ph0i i6 bo3h(333n
qjihcnmg 55311(33936390 0 ^ 3 0 1 5 bnb^gOnb bi03i936noi. b i6 i0 605955,00 bij>-
0ihnbn ^3605, 3i3hob bn00hiC3 3hm6c30ib ih Bih0m59536b. B3pn6ignb i 0 ghmb
^hi6b3nhi(5nnb bi5hd6m6o 39365 (901339 030cnb333i0n, 65533601* 6565(^0030 95-
b ^ s n b &i0m) ih 03n6n9636i (6ib. 43). 6i&336nb 90033^3(10 5itîhni6m 6nb 9 i-
(P59 9(1163^3 036ihfi3636i 531309363905 6ib 9 nh6 ignb ibn0npi(3nnbicn3nb, bm0m
Byonb 93030(30,50 ( i 0 33hnrngnbicn3nb ih i 303* 3b 20-25%-nbi [241)) g i Bycnb
3mli)36unionb 0(3nh3 giJ33ncn36i i0 3hm(33bb3 gng 5i3C 36 ib ih ibg3636.
'biq)b390n 63i03nbib, 3b5qjhcnbm36ob 9monbm3636ngi6 550m0 gn6 ih 3, 6030103-
>1365015 3hfti@3J[jnnb n6£)6bn3m6i 0(3nhg36i. 6553360 303(53big 952)3^,0905 95
v!ihi6 b 3 nh5i3nnb 030i0unh3&3on bi03iO36i6n (055., &ibjxD 2)3Cn P5 aiiro h ^b ^ c0
j3hjnb jbm3no36n, gjmcncnb 952)31335 g i bb3-) bhgQig ihni6 i0mj039363cn (ybh.
4 .6 ). i 0 n^m0 , 0n;jb3gi3ig gngn gidi63om6obi (00 hion 3i3hn g i bni5b3), b3hbg3-
6 i bni5mi3bonbiOi3nb b i 9 n3 o 550 m9 hm6 nb 00530956 i(5nc3&i.
336035^^0^360
339n3ihi‘bnl!)36n, hm0c36ni3 6ycnb 0mcnbm36nc36ib 0ib3n6d3cn 0(336ihob bih-
3b 3 nj)0iyrT)q3nc3636, 5363936056 33hnmg3Q5g 09 hi0 h350(^163690 g i jb3hmcn3h-
r'in' 8
n
lyj'bmtjnb p ib i6ynb9n. Citrullus colo-
cynthis ihnb 6i'bqjmoi(<ni6n 53m qjn,yi
phdig ft563noish36xn
bnb^jBnoi; Echinops spinosus ihnb
bjpjhmqjnQhn 3335 iijpjn; Anabasis
aretioides ihnb phcfep piqnb3n56;)i>;)-
Limoniaslrum pn 6i(y i3i 9(j] 6ih ]; “n9m6nib^h 39
0 ...........I1" ....... 1 q ay* ihnb h m 'b jjx n bibn(jmyb(vn
6 9 12 15 18 qjmh9nb b33XD®i!3hn SUDîh]. hm9 x ^
nu bib pp ]6i y3X it o 90hiC
Time of day [h] 3nhm&3&3n (Stocker, 1974b)
- 260 -
9 i336^ o u fnoaoan
libhnfxi 4.5 bb35p5bb35 împm^n^hn jaîgnb 9^36ihnb ibnQnciyn^h mh5j6mcni 0n6n9john Byonb
31^ 536(30100 (MPa) (Richter, 1976; Doley, 1981; Nobel, 1988)
/ bibynio 3>93hn efT'tJhn. ,&hi>6b3nh>t}ni > 83h50 bn'bmean, 6309^1560156

839:^36001 ^hi6b3nhjynnb
63n9n56n 99hioa 95j33n<no65 (%)
l i i l i b3bm6n bjbroSn
fihpnpm^cn 50}hnyib 39566m
Nitraria retusa 210 165 78
Zilla spinosa 240 150 62
Zygophyllum coccineum 165 80 48
Pennisetum dichotomum 165 65 39
Haloxylon persicum 280 100 36
Hammada scoparia (4) (1.5) 38
Anabasis articulata (3.1) (1 .0) 32
Retama retam 270 80 29
Artemisia herba-alba (6) (1 .6) 27
Noea imicronata (5.5) (1 .0) 18
oô>d-Aim 39566m
njjSon (soo)
Atriplex confertifolia (C4) 155 30 19
Ceraloides lanata (C3) 154 2.2 1.4
y>ij3bnb 39566m
iifnq)mh6n59n (588)
Feracactus acanthodes 17 0.35 2
9x 3i 6 nî) 369n. hm&mhu 3ihi'b0l|)0'b30 b y330 i 6300^333560, 3bhi'bn^)n 61536:5(13

3691 0ijbn0b03hsg 63353(116 8ib3o6d30b, hioii mho33 55951163^ L oran th aceae -b
mjibnb 65(18(08595360360, hm8036oi5 ^ 6 3 3 3 6 63360b bnb^38i9n pi ojn-
Qi6 np>3&36 Byipb, bijôiih Bypnb (133066 3650553636 8 ib 3o6 d30 o b3 8(5365(1-
nbib (ym330 038016333^60 ghmob 5 ^ 3 3 3 3 0 33homg6o 8^0615). 330n3ihi'bn^)n 653-
6ih3 03001(10 8 ib 3o6 d30lb 3 3gjh(’i 3oghm0 i 6 o03 ho 369^ nyrnb, hioii 033d0fnb 0nbn
6yiQ3b3n 653590956 ^ 3 3 3 6 0 60301031136360b biôhm himg36m6ooi 6301306365.
339o3ihi'bo,jo 3qjhm 63^ 6yx)b smhoij036b, 309(13 8 i 63 o6d30o 95 8obo 6553360
3<ghm (gihcnmg ihob 5ibb6o0o. 3ihymqjoono 6i0i6bob 3ohm&366o 338o3ihi‘bo^nb
6553360 303(11*1 53056 063(136056 (65b. 4 .4 4 ). 338n3ih5lbo^36o, hm0c36oi5
9b553b5g Rhincmthus-ob5 95 Striga-bn 8ihi530m 3636obi 95 6 in h 6 i0 ib 3 6 o b
0)3633680 bib0936oi6, 3qghm 909 6i9o0hm36i6 6ih6nii953636 6ib3o6d0ob 6y0ob
h3gn8obi0i30b: obo6o 633(^0 83^ 6y50b 65(135336, 309(13 6 i 63 o6d 30 o g i ih
b3hi336 655336b 6i6n6ii5 30, hng3bi(5 0301010360 ^36m6ib o6y3636 [222].
33ngn^3hn ^)h^0ni6n 01536^3360

330030^360 6i60g3&Oi6 06301 h3501^163600, hm830oiioi3o65i5 gi6ibiboia»3-
&30Oi 3nj)o63nh30 n 6308360 (06060 i6 ghm6 i5hm33636 6y0o6 8ihi5b) g i 630-
836b 0mhob d0n3ho 63i03i. 3mo3o0m3ogh3co 330030,5360, hm5nhni5 ihoi6 00^3-
6360, bi3b36o, Hymenophyllaceae-nb 6ih6m8ig536036o gi '6(^1503(1010 53o8hi
bBhicjig 6h36o56, 6i5hi8 63o8obi 013 60600b ghmb 66(15035933 o6m336 6yi0b.
- 261 -
ffioaoao
1.0 - C
/%
'<n
I 0*8 ' L
s 0.6 ■
50
i
3
| ° .4- \
jO . 2 *
\ Qu
A *\
—y
0 30 60 90 120 150
Time of day |h] Time (min)
Time of day [h]
6 ib . 4 .44a-c Loranthus europaeus-nb 91 9obn 3>bin 6d x f' 0(jo6shnb ( Quercus robur) Byoob 6^ci6-
bn. n3pnbnb 31*116 i3n6(x)n b(p6mbi3C30i i 3b^hni3n a (jhi6b3nhi(yu> g i b Jbnp]8nl> Byonb 3 ^ 3 6 -
Ijn io n 3eoehm6nain ( L 9 ) g i 3i 3hm6ncnn (L c J ) 6u36:>h33&nb, ^ 3 0 1 3 3 3]bnb (Q u ), Sjgihjdncn
Jii3m33hi(vi 6riigijyib 3nhmbQ63n. Byonb ei&sen 3fHi35l5nicnb 3nhmd363n Sihitxitsn 6(536^3
fcibujSb 83^ Byôb (8ip:>oni ^hiSbSnhiyni), 3ngh3 3nbn 3ib3n6d3cn 9(5361(13. c Sshitn^n 91
0ib3nBd3pn 3)536ihnb 6i{,33&nb j>sbb6iX'f'i&nb bihnbbn 6yoncn bb3iQibb3i 6inhn 0rt8ihi53dnb
3nhmi>363n (n53onl)b83d i Q^bnb P 3hm3&3n 6ypnb 6i>>i9nb 3jb33hn935^xn 939l5nh3&i) (Glatzel.
1983)
61b. 4.45 Byonb qjihprninavi 836^330016^

(RWC), qjfTKnonb 6yonb 31^36(511100 (etT0^
9n5n930n) 9 1 Byonb mhcnJcobcnsnb & i& 33& n b
6i 9l>ih36ihfU)6m6> (ee'fOhn SiJbnQjSn) 33nojn-
(53h i 3 h ^ 0 m 3 i 6 3(536:>h33681 2 5 - p c m 6 n
&3i03nl> 33893a (Sinclair, 1983)
- 262 -
80365 ^ 07)^0 m .b e ^ s n a ^ r m o & o u p a c m rtoaoan
&33hn gSncgn^n (fciBbiigcnhg&nai 3m9 mnm3nghgcn £)h^cn56n bibgm&gbn) 030

ouhgiib nbgoi o^nbgbgbb, hm9cg&ni5 bgpb gBym&gB 6bQ3j3bnbi g i Ri3m6ig36n
6ypnb SmSihi^gbib: 6;)gnb 9i53ihn d)mh9 i g,3n9hg&0n; 9 ncdi6 hnb qjmh9 nb 0jm63
gmcnpnb dnhgbn Bromeliaceae-b mjibnb 6ih9m9ig5360369n; Araceae- nb
6ih9m9i9536p369n pi mhjng^^bSn 33Qi936ghn 0 3^ 336 ^ Bypnb bi9ihi2>m jbm-
3nQ3&n pghmgbbi g i q}mmc3&9 n gi bb3- ^hmSn^x 63n9 nb ^yggbSn Ri9 m6 ig 36n
9^^33630° Byonb (g.n. 6nigi5nb b x^ n h iQ g g ih3n(j>633on Bycnb) 50 %-bg 9 g^n
gSncgn^g&bg (nb. 01130 4 .4 ), 9 i 9 n6 hmggbiu 9 icnn cgnli)m3 ib i i 9 ^ynb
bi^hcnm ajn^mSibnb 9bmcmg 7 %-ni [ 184 ]. Bycnb 9m9ihij>36nb Sipipn g6ihn
iSunhg&b Bycnb 3 g0 U330m&nbi gi Sm^gS^niQnb 3ihg6nb bihnbbb Sgîbmoghig
35n0363om3i5 ;jbm3nog&9n, 9ij>., 93bmqjno9n (nb. 6ib. 4 .4 ). Orchidaceae-dn,
Bromeliaceae-dn g i bb3i mj:>b3&3n, bigi^ ginqgn^ghn bibgrnbgbni g i hm9 cg&nu
6 ib3 nh6ignb ibnSnQi^nnb CAM ShnBijnSb nyg6g&g6, fci^ggbnb 3 mdhim6 nb
gpnghn gn6idn>,nb SgSggm&ncn Bycnb imhovJcgî gi6ip gmSgbg 93061^1636361.
i;)3gi6 J)i3 m3 gn6 ihp, ibgcn 9u36ih33&b Sggdcnicn bi6g)hdcn3n ]6io3;jm 3ghnmgg-
6nb gigbm63&cig îgi^pBi (6ib. 4 .4 5 ).
4 .4 a i3 36^ h D 3 C n O l ^ L O 'b n & ^ e m D b D b f l b 6 y C f lb
hoanan
4.4.1 ai5DB^hDXn b ^ h f lb ByCHb 6 ^ 6 b ( l
4 .4 .1.1 eycnb 6 ^ 6 b n b 6^6i!»nCD6i
0U35ih33cn biqjihnbi g i 6mgi&nb giqji^niBgbgon bm6nb Bycnb hggn9 n 9 gnd-

035)1 {>i9mnbi0)fnb Bycnb 6ici6bnb fci^mcgdnoi, hmSgcnij 3bj>i3bni g^mbnb^^nb
6 ib3 nh&ignbi (2 .21 ) g i 9 n6 ghicghn 6n3ainghg&36nb hggn93&nb (3 .2 ) 6ici6bnb 516-
linQ3i>36nbi. SugBihgxn biojihnb Bycnb hggn9 nb y j x ^ bngng3 g)i3moi3QnQni q)ih-
oimbnb ]hcn3x'bg. hmj,mh(3 6ic3jgbnb 3J3n3icg6^n 99 H20-b3 (g.n. Qn^hn 9 2-bg).
013 513 19 ^ ^ 3 3 6 0 1 3nhm&3&b gi gi3g93g6cn, hm9 9u36ih3x n biqjihn gbhg6-
330ymq3ncni Bycnoi 0 brnQmg 6ic3J36nb bihjbg gi ih ihbg&mbb 915361)13011 Bycncn
3bh363xyf’iqjnb bb3i Byihmgbn, Bcnghn 013 ioiBc3X n^ 6 i 63 i 3cm6 i 9 n Sgcrv
3nb3&3cn Byijvi (6spjjjdnb b^dy>crt }±dn Pr) gmbihsg&i 6mgi£>nbi g i ©ijgBihgg-
Qn bicghnb 3 ngh imhoijcg&ibg LE) gi 6nigiftnb bggi 3nhng6
Bycnb 5 ^ 1 ) 1 3 ^ 3 ^ 3 g i 6nigij,nb q+i9 i 0336g63 n 9 nb Ridnh3ib3 (L^p); nb. 61b.
4 .46 . bi59 niC 3 yn63nb ghmb 3*)mbnb^30nb Bycnb 9 ihij,n (+AW),
higgle gqjhm Sg^n Bicojn 9 mgnb, 3nghg mhcnJcegbi g i Rmhgubg&i, i6 i c g b x ^ ^
(-AW), higj>i5 6icgjg6nb himg35m6 i 3gh ojihi3b imhcnJc36nb nB^gBbnsm&ib.
ij X 56 jp 3 m9gn6 ih3, 6ici6bnb 5i6^mQ3?)i j)i3 mnygh36i 3g9 gg56 inhig:
- 263 -
rioaoao
fob. 4.46 a 6id 36ihn ^ynb

6ypnb 6ioi6bo ^m&iSn g i b
i(t<9mbi3C30> 33hm3nb 9]b6ih3n.
ET 33i3ryjhi6b3ohi|5ni; I n6&jh-
bQgiyib; R cgnono36'b3
81601619360 Byipn; P
ogmoipjJrt)^ S^hfia&ipn &p<vi
Age of stands [years] (Molchanov, 19 71)
6 ib . 4.47 dptfohn ^ynb Sycnb bicibb n BjqjmcnpnQ 91 933qjmoip:>3 33hom9369o. Pr 6sj>3jnb b ijh o m
[11019360161; Prc 311133630 51^1363(10 6i£3job 01936mbs; Prs p3hoi3&b Ri3oiyoioo0o 6103)0; Pr;
n6q3n0j)hn5ni (60191580 fisb3(yt 6yi0n); D 9(136130(136390 6^190; E T 33i3oy>hi6b3ohn5oi; Tr (J36m'bnb
OjhiBbSnhi^ni; l:.sc) 3313011111501 6(119150916: A W is36m'b3n Sypnb bijhcnm 938153300161; A W p 6ycnb
339(533Cfi&i gjn^mSibiSn; A W so 6ycob 333153300161 60191590. i 9 630061980 bid3i9m p 966
33 6iC3jo 30190b; 3obo 52.5% 3611369361 i^ o ib ^ h o ib i536oit>ob 33i30ihn500b. ^hiGbSnhujnnbi 91
60191501) 33i3mhn500b bi93iCD6ncn; 47% 513906361 60191580 91 5X3019019 0 .5% b3 ih e 3& i 600161-
bob 3m 3i^36ib (Schnock, 1971). ghi>ftn£Cn 63060160 iiy336obiort3nb nb. Brunig(1987)
- 264 -
9036 * 6 3 ^ 0 cn ^ u sn a^ rm o & o u A 33090
Pr = AW + L E + LRp (99 ) (4 .15 )
Snghrxyiftn^hn (VVjjhi^^hxn Byihmjbnb 8n b x 3no'. AW 5>6nbn036;> hm^mhu

abmQmg 6nigi5nb 6yQnb h]‘b3h3n, y n. 9 ^ 3 3 6 3 0 0 9^ Ridnhxn 6i9my3636ign
6y0nb hbmp36m6i. 6nigi50n 6ypnb ySX^'bO 8b(p:>on 33815330^6^ ifn6n8630ni bm-
603(1 bih^tftcdn igh3 &>b$q]DyA>3 - oim3onb p6m6nb 938935. biqjbxSn 6y0db
939i53X fT'2>i. 6i03fo£>nb 8(13639^339, 8(5nhg36i 93 biqjb3cnb 6n0mb b(p633b 9 n6 n8 -
38b. 98 hi0 (1350(1163680 6ycnb h3b3(i336n 33fi3b36i 338(1195(119(101 - 63n9nb 33(10-
mg9 n, 0138(5* 6oigi5ob ^339} qj363&nb 6yonoi 5ij3h36nbioi3nb hi9g36n83 j^nhii
bi^nhm.
3 i|Tl0fT*6r,Dhn ai3i0bbbhnbnai AW ^ 3 3 3 369^ 8305^3936 Sypnb n9 8ihi5b, hi(5
8536^(13013 9ib i 8 n g i 93(^hnlj8ni. 6i0ibn3i6n i536mb36nb 63900 8ibnb 3/4-b3
03,5b gi ihgnn6nb 63gcn 8ibnb l/2-b3 9 ^ b 6yipn 8339536b. 9^36^33011 biqjihnb
Bypnb 939i5330fn6 >b ibibnioi36b ge>3-C's93 hn g i 6cn3hn 93hy3(n636 n. (15(1 9 ijbn-
838b i(p633b 93 <3 mai03 nb qjibi3 n. 6ib. 4.47-b3 811153830(13 bm9n3hn bih^ypnb
qjmoi0m3i6n ^ynb Bygnb 6i0i6bnb y33Qi 8^33636300 b iq jb x nta b i 9 oihob
qjib363n.
4.4.i.2 bocanw639nan (63ibn) e^cDjfi
9y36ih330m6nb5i6 Oii3db3qji0 6oigi5b3 9 m b xn 6i03j3&nb (oii3nb3q3ipn ig-

500360b 6303^360) himg36m6i 3qjhm 8 ^ (13^ 3ngh3 8i536ili33pm6noi 8^nghmg g>
qjihx 6oigi5b3. 8 153 6 ^ 3 x ^ 110 1 giq jih x 6niQi5b3 6yi0n b3g36i b3 9 y 36 ih30ii
33(1380 ihb3&30n oi33ob3q33Qn b(i3hi533&nb 5i3pnoi (di6*hji>x>n BysQn), 6i0ib6ih-
0936 fi33(116330136300 (fodm63jcnjdx'n Gtppri) g i 'i)m£)6ngi6 Ri9(H9C'36ono0n
(RidmypjBomQn GysQn) 6yQob bibnoi. 6303j3bnb n9 himg36m6ib, hpi93cn|5 8(53-
6ihob 5 i 6iih 53Q36i 9 ni1 36dig36i bjcdnbsBjgmdn s6 j 6j^m-6iQ3jjdn. b300r>-
bi63gm9o 6ip3Jn 5i6bii3oih36noi ihioii6i6ihni »jyo369n. 3ihj0(i 5i9i3i0n g i 3ih-
jnb 5^301 ihb3& xn 6iC3j36nb himg36(n6ib 539386933^1 36n0363om6 i ij3b b33-
60b 033b336nb 5i3hi53C36nbioi3nb gi j33^ynb 9j536ih330Pi6nb 5i63(ioiih36nbior»3nb
(6ib. 4 .48 ). p3hm36ngi6 g i lyn£)6n9i6 fii8m6i«)e'36oio 6y0nb himg36m6i 9 noi
3qjh(n gngni, hitf 3cghm gibhnc6n ihni6 nbn6n g i hiy 3qjhm 6C33(ii J3hjn; 6nq)0nb
bhmb dnhoii6 6(1393590 633906363 1.5-33(1 83^1 6yiQn, 309(13 0n 03X 6(iigi58n
[153).
815363(1330 biqjihSn 9^3363^0 6ypnb x n6D^jn 6i6o0o obih336i imhoij036ib3;
<3fnoip36nbi gi j3hjnb 9n3h 03noi3nb36i 0ioib3 3mb33gho0n 6y0db 8bmQmg
38(50(13^1 636000. i 0 nli)n9 8(5363(13300 bicghnb 803b gii>i33&x 6yi0b giîh530o
6yi0n 36ng36i (n6,t,jh(jjîjnjhn Q^sjfshftjdn). 6nb0Oi6 ig5(i0360o (9i(pi00oii,
b(j>3nb 1503 90636360 ^ ihb3630n bi6 i 3nhri36n) b308nbi63gfn8n 6 i 03 j 3&o 83^13.
3(igh3 bb3i 3i 6 oopo»360 o.
n6ii3h|53<35f*Dhn gi6it>ih536nb hifnp36m&i gi8(nj)(ie3630Oi 8(536^3300 biqjihnb
636^95360(116^3 g i bn0^ngh(i«33b3, i5h30i33 6i03j3&ob 8mb30ob ghmb ihb36x
93li3(Tihni0fn5n3h 3ohm&36b3. bnb 3^ 336 0 3i l|>ihi i93op ig gib3XD^ên <3(1*01036001
i6 606336001 3qjhm 8 3^ 6yi0b ^ 3 3 6 3 6 , 3ngh3 h6npn g i gngn gjmoiQnb 8^0163
3ih33&n. hibij)3nh33Qni, i8iboii6 3hou>g gng hmpb ibhxD^b 03qgmoi03nb (16,336-
- 265 -
e a ^ n u Aaaoao
bo3hm6*. 6063013*6 06^311153031503110 9*6*3*115360 b*63*9019 20 -35 %

63*9536^ dpnjh Q^nphm 6/1631^13*6 i»>y30£>9 n - 50 %, bon603(10 bih^ycnb
03/11019/113*6 i!»y33&9n - 15-30 %, b3&^hni3n))X 63fi;)6*h369o - 5- 15%, 3*9636ob
£>9*3n - 10-15 %, b 01901 ^hm^nix iiy03&9 n - 15-70 % [46 , 52 , 174 ]. 1)33^3
*3*33db 6*93j36nb b*63*90i9 10% (5-20 %), 8ôghoi 333s 636^6*11360 - 50 %, 6*9-
*b6*ho yi 8(5019019 3 -5 %. b*b0iq)90V'b*833h630i 03*110101636^3 0663)1133031503)10
9*6*3*115360 10%-b3 653936/1*. 13359365901 *8n6gnb 3ohoi63&0n o663hi53q3(5nob 60-
901 6i93j36nb b*3ham h*oig36oi6*9o 01539365 8*cm o6636bo3oi6ob* 9 5 ^nSnb
(63/165, 6oib0/i, 010139/1), 63633li*63hob* 95 j*hob bod9n3hob 806393/101. b*'boi5*9oig,
Rh RhV RhM 0
6sb. 4.48 Pinus cembra-b &*h9;)9m 6iC0j3i>rdj {^SiBnpjbi BscoJjbnb £^$1600365^3 J3jjynb
5530365. a 3h]pn - b>59mi03 dcrvjhn 6309nb ghmb (100% = 6 bav3n 35.5 99 6seoJb); b 9h}pn
- b^65hdQ03n dprjhn £>3n9nb ghmb (100% = SO bo*-3n 14 99); c Bh^pn - b ^ y i 63/18* 6nbpoii6
3hou>p (100% = 1.8 99 BbpoJb). ^33^ 3: Rh = Rhododendronferrugineum-, RhV = Rh. ferrugineum
p i Vaccinium myrtillus; RhM = Rh. ferrugineum + V. myrtillus + bi3t>3&n; O = 9063x 0 9nCb
(Aulitzky et al., 1982)
n6^3hu3qj(5nnb 6/19/1 9 noi 636/15, h*(5 6*3936*9 06^361*13)1/1* 6*93^360 g i 6150)13*

63060b 63301360 (6*b. 4 .49 ). 60139/1 8(536*113x0 b*q)h/ib 5*g(?36oi3*b 3b*^nhoi3&:>
6ygnb 55)1333390 h*oig36oi5*. 66019019 85b 638935, hatf 3b 8oibg36*, 6y590i n6y3&b
03/1101936/19*6, 6/0^36/19*6 g* 013)1013609*6 6*6016330136*^ 5*9P36oi3ob 6335Q<T>65.
(101501)1(5 5*6*h3&ob <be,3hX n 8*633636390, 6063/113*6 6^33660 01)133)1 83^0*, 309)13
0301010013*680 (630501019090: 1-2 86693; 6330301019*30: 0.5 86693). b*6*8 6*92)3-
60 6o*g*5ob “bogiôhb 60*0163336, 3*093360 9* 65)11539013*60 693901360 *3*33636
1-2 88 6y*cb, 601901 b*3b36o ^01)1030*63690 - 15 66693.
- 266 -
80066603=50 0i66ib68na6«?na&060ij ed ^ o b rtaaoao
Total precipitation [mm per event]

6ib. 4.49 ^ynb biqjihn 91 BiQjjg&n. Pinus cembra-b ijsb dpnjh Soihxn sihjn, idn^md 8nb&i6
3^j36non ^ynb biqjihn^ So^h^cm 91 0131101 dQnjhn 53ndnb ghmbnj yi 6niQb&nb b^giSnhidg;)
himg]6fT>6noi Bieojb s^h^db (Ovington, 1954; Aulitzky, 1968)
4.4.1.3 D3i 5HU)hi6b3nhil5ni 9l306ih33Cn b^hfie^
9U36bh3x f' biqjihnb 0n]h Bypob b>h33> 6nm8ibob ft^Q&nb 3hm3mh(yi39fU>

(61b. 4 .50 ). c n ^ S is ^ O iD X 0 (Bficnp^bnb vi)h36b!}0hn500b 06^36^3016^ 80 36^ 3-
X*1 3sbnb bnS^nghrôb 0i(j)36it>aii6 3(101*9 3e3&3cm&b, 30610916 0ib <bC'3Pi 3^
Onihmicnai^n (516^3301(136001 0bnb hign^nnb 3ohgi3oho 0mJ03936nb 830150(13-
61, li)36ni6rTi6nb ‘bhgi, jihnb 030150(1361). ^^bôhiijoob bi5hd6m6n 9^33001361
6306006361 dpn3h 0Ô9hm 0 (536ih3x b:>qj:>h3 n (61b. 4 .51 ).
ii(hb6 b3ohbunnb o6(i)36bo3ri6ob 030150(1361, 5 i 63 ohm636x n ©1536^3300 biqjhob
ob3nb363h363bnoi, 5100^1^361 sjhmgndsdnflhn dndm(jjQnb 6 n6 ^(pdpjg,mdnb
(Rae) bi03>Q36noi. 3 (536ih 3x n biqjihob 6o6inj’0935m6ib (R ^ 8319536b ^h>6b-
iohiyoob y33pi 03019536390 6o6ii(n0935oi6i, 33hdm9 91 qjmcncnb 90033^0-
3ho Bo6nh0935mb36nb (Rj) 3*80 [ 156 ]:
RC = R ,+ Rae (b0 63-1 ) (4 .16)
9l536ih3xn biqjihob (i536m(bob) 6o6iip0935m 6ob 5 16 ^ 1 9 3 6 1 4 .10 qjoih0x n b

9bj,i3boi, b ig n j AC-b 6 ^ 3 9 ^ 9 9nu30x n:> 6ycnb >mhoiJ93&nb 5(119036,1)0 0(J36>
h3bi 9 1 5ih30rri0jj339o 3^h n b 8ib36b 0mhnb. y33p i 3 ^ 1 0 3 ^ 0 0 mu30xr^> 6n>
9150b q3ihoim6nb 3hoi339<b3. 00165(530360 b b 3i9 ib b 3i u36pVbobiai3nb 0my3i6o-
Qni 61b. 4 .52.
05033 3160161301003(136360 0oij03g36b 311150b 91 03bi6i0obi9, 155933390 bnb
030oib333s0no. 033^(739130. 6153360b 001^039365^3 91801330936390, 3183(10 (jhi6-
binhnyii 3mhxMJn;>9 na qjoioi936nb 3103I1 lb39i3nhou>6 16 8ibibon6 (ybh. 4 .7 ).
9 ^ 3 5139361b 16936b sihjob 51933390 61609360b 3(10103(1015595(31(131 91
qjmcnpob b33unqj033h0 550^(10160161.
- 267 -
*
w
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4 .t 0 Alpin e herbs
4: . ' • Mea tows
A Dwarf-shrub heath
\ (
+■ Tree stands
O, O,
x Forest herb stratum

------------------1-----------------i—
0 0.5 1.0 1.5
Total foliage [kg FM-m5]
6sb. 4.50 963163 9 ib i (81160615351 ihi93hJ6oi6o 3ibnb 63900 6m6i) gi 6ypob biSîpm ptnn^hn
3 mb9ih36i (Pisek & Cartellieri, 1941; Polster, 1967)
-o
c(0 —.
Sw
1 Te
.1 P
2H
a- E
CO —
Leaf area index [m-m'1
61b. 4.51 6ho6job 3ci6îonnb gihgmdncnn ,jhi6b3nhi(jnnb (Tr/Ep) ei6myieod3co£>i 8303^10103(11;

bnbSnt^^ (LAI) 91 0690309310^0 8(536ihnb q j c n o i Q O b ^hi6b3ohi(5ni'b3 u36m'bob b ^ j i j j h n b
5130361. O 0690309310^0 qjmciricor • l336Wbo (Sugimoto, 1973)
- 268 -
3 (3366f?>a7)g;o m 66^68ng^^na6a6au g ^ n u rtadoao
Canopy conductance [cm s 1]
61b. 4.58 6ypnb mhorjQnb (1/Rc) U36m<bnb ^hi5b3nh^nnb 9i9myTp3&X'3&nb i3 b ib 3x ,n 9p'X'rrilxn3f>b nb. Kellinher
et al. (1993)
y ft^ ^jnnb & 6 dnQ<b3 (56 Bpob 5568530016580) 81536563300 l>5qj5hnb ljh56b3n-
h5(yinb 550rob5cr>3Q3Q5g b5^flhm5 501(101^03606 ghoi3 o 153593659016065 95 6990b
05hi5nt> (301965. 50nbcn3nb 5hb3&oi6b d0nQh 9 Dii50n<bnhQir)0 n bndx,MJn3hn 60193-
9360, b595(j 55oi350nb6n63630n5 y33Q5 dohnoi59o 309(10190150360 bngng3 95 <301-
oi0nb qphoiob n693^36(1. 03tb p39X n bo3hisxfl 6^63001563606 6^0163 3m8 m-
5363(10 I536m'b36nb5cn3nb 3353m^h56b!3nh5ijnnb 9003(10 56 6903(10 8089065(130165
5560100139365 363(153(1)03390 &5056bnb 80165(53836001 56 0nbn03ljh30n 55<bm033-
&ob 9535036001. 50 nb 0555900159, 65b. 4.53 60193560905 93h60 nb 6505b65hnb
iihi6b3nh5ijnnb 6903(10 90658035. 3b 015365113300 oi565b5l6 oi55goi365 5hob 6ycncn
bijfdihnbsg j'bhjG^jQymqjnpn. ^hiSbSntônb 3583(10 80165(530360 93365658365
}mhoij03&ob 900956 903693 153593659 3ohoi63&b. hoig3b5(5 6y0ob 65303601606
ghmb 069365 6553360b (135390(1365, ^h56b3nh5(jonb 6311930165 30933 303(101 dfyi-
3(19365. 50 ghmb b5ôhoi5 b50359Pi gpo3h ^hi6b3nhb|j0ibcnb6 3hoi5g 699001 153-
959 9 oi05h5536ob 3 ohoi63&6 o 6990b 6h363ob 80608593(10 ©583363603606 g 55&-
55h06365(5. 8 8 (159 63500163660 001*6560 6(53656301501306 9090 860636300165 5j3b
6y0ob bihj3ob 600360 65833636006 9595365b.
^933660 ^h56b3oh5(50ob 60o3ho 8089065630165 95601309363005 0901010360b 85-
bob 5563ooi5h365<b3, (301010360b goqj3h36(505<63 g5 5b53<b3, 55630133 <336336065
95 y0oih^,3bob ‘bhgob 3hoii53b3663. 501010198(5336 13933660 59^13 55lb5<3 b x ‘b3.
630901019306 956936560156 3hoi5g, 35(1306 501(101^9365 85^390166, 601901 <3010193606
'bhgob dohooogo 33(1001906 6oi0oiboi3ob 50633b 85jbo838b (65b. 4 .54 ). 05033
ooij0nb boi8o3ho bih^yQnb 65(159863563 i!>9336'63, 1101839015 601(18 606336b
65h056g3036,b3 309601 061336603(10 13(156630(151505 5^301. 33801050185*63 g5 *658-
oi5h9o ^15663065(505 00606335693 3(5385. 3b, dohooi5g 5g, 55801633x05 5vi6mbqj3-
hnb 5oihoij03&ob 956590 365(1001; 585b oboy 385^365, hold tadoihob b3363&ob
95 556653301(136001 30 90630b 33(1001960 6553360 5h 5hr>56 6565(^90359 95 6(13-
959 550363905.
- 269 -
(jbNxyi 4.7 63636(103 3nhm6363n bb3igibb3i Vi6nb b3 8036^30^ ^ s 5b3nhiuni
(a Kline et al. 1970; Jordan & Kline, 1977; Ogino et al., 1986; b - Jimenez, Morales, Kuceia, Cermak, &s9m3j33y636ocn 9m6ii33 6 3 6 11; c - Doley & Grive, 1966;
d -_Ladefoged, 1963; Braun, 1977; Kunstle & Mitscherlich, 1977; Cermak et al., 1984)
s
I
O
oi
o g
d o o'
oJ o o
*0
128
o
- 270 -
%
c
i*
in in
*8 -8
ifi to
-DOC
Cl OJ >1
4II
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0
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I §1
8 0 0 6 5 6 3 ^ 0 m ^ i^ s n & ^ n a & a io u rtaaoao
c
2
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Q.
O
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to
O
6ib. 4.53 pjh60nb ijhi5binhi(yinb gpn^hn g i 01303!™ 31836(1 LSoC^hiCjh 33hm3i9n b i3] 53^snm
3;]hnmgnb jp69i3CT'6i3n. gjnnjhn 1331c o6igm&i6o g i9mj,ng3&3pns Cmi i C!)h >9n6p‘b3 (Kiendl, 1953)
----- W ater consum ption of tree ------Num ber of leaves

— Transpiration per m 2 leaf area ----- Total leaf area
Bab. 4.54 3 3 bn9ie<pnb (jnhnqjnb Byonb SmWshj&i j,>gii6j|ihn93&3cn: 3hcno qjmcnonb qjihcnnb
3hoi3X^3' y3X3 ojPKJicnb qjihon'bj gi y3X3 <3f'0'Cn^ himgjSmbi'b]. Qhoin bnb qjmoi0;)&nb him-
936n6i'b] gi y33Qi qjmoipnb qjihcn'b] gi>66ihn33&xn ^hi6b3nhnjnnb 06(536^3^161 y3X^3
Sipipni tiqjbxHb 3]hnmg8n, hm(ji qjmoicnb dibi 1(^6336 3ijbn9]3b; jhcnn CTjpioipnb qjihoinb
Dho'dx't’Oen6{)ihn33d3cn (jhiBbSnhujni ysÔiV) Sipicni ,biqjbxn*3 gibi6ynb9nr hmî cgmoncjftn
n9c36rb5 (Braun, 1974). dbj^bn 3m6i(338360 6063^13360 Picea abies-nbsaryfo nb. Cienciala et al.
U992)
- 271 -
E a ^ o ii rtadoan
(jbhncn 4 3 bb3*eibb3i ^n3nb 3u;)6:>h33(V,t&ob 6003(10 gi gpnjhn ijhi6b3nhnjoi

(0hi3ipn i3ijmhnb 3ob3g3noi)
^ }06ih33om6nl> ijn3n
63cn6ie3n (39) ero9n (33)
83(1^60160 3u3&sh33&n
(jhm3ni30n b03&nb 6ih5i336n 2000 - 3000
(jhmSnixn 63n8ni6o oy33&n 1500 - 2000
bm3n]hn bihjycnb cgmoxvrgstjn dyOJ&n 500 - 800 4 - 5
3^59663*63 5n63m3>6n (jy33&n 300 - 600 2.5 - 4.8
biC3hmqjnC3hn 3u36ih33cm6i 400 - 500
200 - 400
Ericaceae-b&Jo 338951(10 5^^0636300 636^6^360 100 - 200 2 - 5
8Vx}3cm3s5n 9* 6&nhtaQibfn3i6n 8(536^ jjxn & i

nbpoi636n pi 03(169016360 1300 - 1600
3i(j>i06i0ib33Cm6i 800 - 1500
6m^nm 89300(360 110 0 4 - 5
3ih|j,3pm3i6n 893001360 400 - 500 2.5 - 4.8
5h3hn3&n g i ^316360
bicr>n6n 9g 30013611 pi bidm3h3&n 300 - 400
b,j3336n - 200 2 - 5
p n i l& & n ^ 3&nb 8(}3(tth3 xn ite
3i 0mq)n^)3hn cniSibi'bmîpm363611 2 - 5
3cnnb 6i3i036nb 8(j36ih330m6i 10 - 20 0.3 - 0.4
00^36(1^6(1 tjjBphi 80 - 100 0.01 - 0.4
ihngjon 39^66^1360
bb3iQibb3i 3000^3(1 bm6 i 3 n 8(536^3300 bicghnb 803(1 6y0nb bbhj3nb ^n3-

03(10 8016^30360 0my3i6nQni (jbh. 4 .8 . 3hcnbi g i 06^33 3008^3(1 bm6 i 3 n ^ 3 -
36b, 3 ib 3 î 63hn £>i63ncnbh3&nb 5 i 3 m, 3qjhm 3 i(T>i0 o J>hi6 b3nhii3ni ibibnioi3dcn,
3n@h3 3g30rri36b, bm0m i8 33i6ib363036b 3b 3^^336363011 3qjhm gngn ij3 d , 3ng-
h3 5^0330^013630 63RJ6ih36b. 6y0nb bih53ob 3nb3g3noi 30(1330 ig5n0b3 ihoi6
nb3Cio <330^01(3360^360, hm330 onioi3nbi(5 b300nbi63gm8ni 5(136,1^ 690360; bm^-
33(1 nbn6n 83,3 6yi0b 5^(53836, 30gh3 6i0Dj36b 801^301 (nb. ybh. 4 .9 ). 9 3 hip
(13500163630, 6y0ob @3030(50,^ 3i(!)36ibaii6 3hcnig, 3^36ih330n biqjiho 83^3(1-
g3&i. 6nigi5ob b3gi3ohob b3300t gnb^bôi 8(536ih336b 3 rnhnb nbhg3&i, 0136-
(ji 6 nigij,3 n qj3b3oni bnb(j38i dp03h 3001^936^. i8ho5ig, b33&b 3 mhnb 8i6do0b
5i6bib(j'3hi3b ihi 3bh33&nb 3hoi03hai5igiq3ih3i, ihi03g 1^03330 b30ii 033b300
bnb^30nb 5i3hu3036nb hig03b0. 53^030^6 (13500163630 60O3hn 9m6i(5303&rib 5 >
3 moi30ib ibho ih ifeb. ij 03od03&i gi3ig5o6mcn 630806 33homgoboi3ob gi0i-
bibnioi3630n bi93i0oi 9m6i(539n gi 53iC3nb ghmb ihb3&30n 0o6n0i03ho bngng3.
0(336ih330 biqjih3 n 6y0nb bih33ob bi33i0m g i 3 o6 o0 i 03hn 3m6i(53036nb 5>-
3 moi30i ghnig bibihjtfbgmi 33^y333m6 i 3 o g i pi 6 g3 iO)^3h gi>6363 ih36 i 3 o (0i6g-
3iqj^3h ih jn ^j^h iSo ), hmg3bi(5 i 3 8m6i(53836b 5i63obo0i3oi (135001660 6i03-
j36nb h3gn3oii6 Jji33nh3n. ib3, 3i5ioooiig, 5i8moi3036o 530(1336366, hm0 ^ynb
83663(1 biqjihb ym33060O3hig 3bi^nhm3&i 0o6n030 110 93 6i03jn ( 10-12 03
ym330oi3n3hig 3353<!>iljnnb 33homgnb 5 i 63 i 30 m6 i 3 o). 6y0nb (139080b 5i6 vjm036i
(4 .15 ) 530633636^ 013 big oj636i ^ (136 ^ 6 30 3(10 ^yob 5i33636i; g i 3ohojooi,
- 272 -
a o o e ^ a -a ^ o O ttB ^ s n & ^ r m o & n b p a^n b tfaaoao
S (^<CKicoQininaj
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(0%, >c3db b], 33l>i»Cn3 i!>n) pihj>3:> b^pb S^Symftb p^DÔ^DO^ Bypnb pm6nb
p36333b 93 3i 3hnb g:>^36 n:>63&3b.
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p a ^ o u 630090
4 .4 . 1.4 eycnb 6^hD l5b3^ 0 Q^hOiW
63031)3606 bibnoi 9o ib x n 69300 901003639 3h 01(101^09363 - 9obo 636000 53-

930(1356363 tb3g33nh303g, 636000 y i 9306(10,5363 6039350b (ph03 03363690, 833-
3(1019363 5(136,506 699366 93 09063(133606 5330001 63601001019 639363 <bp 336b3
93 01333633690. 6039350b '639330(10936 6y0ob 5393(135633 9393h36oon om0o 5363-
t>m0o3, 536b3i30ih3&ooi 939o6, (10193635 530mb3i 3033O3 9go63h39o 63906360b 63-
i 3<!>nCn (1350016360. 6039350b bo(nh030o 63(1356306 (936(13,506) 030336363 30 93-
b3d03630n3 9bm0frig 3h33ohg33oho 030101936001.
dsgshjubnpn GyQnb 113019360163, 330(133036 yoi30ob3, 930012)0936x^3 113003-
030b 93bho0fn63<b3, 95363(13300 6303(106 ,50363 93 6o9ô0hoi33b3. 9393(1360601306
56(1. 4.9 0mu303QO3 6633936633 ^nSob 9jj363h3x b3qj3h9o 63031)3606 (1301936m-
60b, 301(101^036063 03 6y0ob 936(13,506 00163530360. 99h30 h35om6369o 6y30n
qj3b336ob 9o3h 0993639 o6v536bo3h30 9306013363, hm9 5393(13(5633 9363003. 6393-
^36001 09093(1 3360,53,53680 536632)301(136x11 660936300163 3j3b 603935063 93
93iiho^,ob b3qj3hob 0O3h 6y0ob 9361*1306 363hb. 6y30O b6h3O)30 93 9090 9363-
j,3h536ob 53h393 9306013363 ^yob 6039350b 9o3h, (10193905 39333 Qhoib 93033(13-
003 93,5(10,506 bj30O b3qj3hooi ^30336001). 3^09(101 6303b63ho36 9930oi36b 3 6y3-
00 6303 0306013363, 06333 1101501(15 60oj3&ooi d0O3h 93^533600 63doi3h36b3. 363-
33 b3b^03 6y0ob 0360133 63yo6x 60393590. 39 ghmb 536963(10 6y30n 5(101393-
63 53633690 36, 013 b3033oho 93bho0o3, 53d0O3h3&X39 636013906363.
dpnjh gibhnpn qjjhpmdjdnps6 6y0ob 636333(^3 63,50 b 3g 33o h x ^>0 5393-
0(1356363. 0132)0 6393^360 90903, bm0Pi 65363(13x0 63qj3ho 96 oho, 6y0ob 2/3-936
3 /4 -0 Q3 53930(1356363. 6013660 50536m <33(19016360936 63601(1356000 6 y30 O d0O-
3ho 5(1330,5350300 363(15006 530m b9oh30 b39n9 6366,53636b 3C’633b 93 333^903
539mn63omb 3hmbo30o 9m303636o, 036y3hn 93 0133(150103360.
4 .4 . 1.5 9 l3D62)hDDCfl b ^ h f lb SyCflCJl J b h 3630 Cyn<3 flb b b 3i 6 b D 6 fl
05363h330O b3qjhob 6 y0 ooi 3 ^ 3 6 3 x 9 ^ * 3 ^ 69363 3h3 0bm0mg 6303j3&ob 039-

33016001, 013953 3b 5^3, b9 oh 939oib33339o, 390133(13603. 05363(1330 b3q33hb 693-
00 0036019363 6633 5'booi35: 5(136,506 69036001, 632>30XD&n°V 63901363(10 9 oih-
693001 93 663. 3(10930 (13500163600 6399030 95363(13x0 63033(10 b9o(i39 30013(1-
9363 9bmcm0 o9 395003690, 63935 5(136,506 690360 b3933ohoi3& 03931136001
3b0mb33. 339093(10 h35om636ob 65363(13x0 63033(105 3:1,503(139 093636b 5(136,506
69036b. 6039350b ("h93 <3363600 3(1636x1 6y0nb 036m3ob b3033036ooi 65363-
(13360 b30b 36901636 03 36j3h3636 6990b 6(136336 60016033(10100, h39536 6393^3-
60b b3booi 0 m bxn 6y0ob 63600b (b3630 050 'b(j>330o 63x^6363 93 6y0ob 6(136-
30b 9090 6hob b303303&ooi 53033 6036019363 95363h3x b3qj3hb) ob33 36(13-
63636 3^9oib<33hm0o.
- 274 -
8ff>^0b * &56300166360b 6367 ) ^ 51)05
5 tohOamb 6330363 9l5363hnb bhei-

6363fl013h3&3‘b3
9oixn bnumi5b0nb J>i69 i 30m6i 9 n 'bhgnb 36ihn 9jj36ihnb 306n93630m3:>63bn

oi3nb3&ii. 9tf36ih3b j>iifi6ni “bhgnb bm636n ycmh^ftbi gi <33b33&nb 633hm3&9n
(s 3 nj)bcjhn djhnb&jds), hm9036nu 3^363309^10336 ibi0n cgmcnQ3bnbi g i y3i3-
nc3&nb ‘bhgib g i qj3b30ii bnb(i>39nb 83g0n3 6i3hb303&ib 6 nigij)9 n. 03mhign
33hnb^3036n 5i6i3nhm63&36 j^jiih n bnb^39nb 33hnmg3p j,i63ncnih36ib g i j3h-
jnb j>ibj3C36ib. ndnbgi 3n3b3gi3ig, hm9 9(j36ihnb y33pi 6i6npb ih 033dgni bh-
gi g i nj,n ih bg3<bi ghmnb y r ^ o 9m6ii)330i9n1 bhgnb 36ihn 9(j36ih3b 9cn3Qn
bnumubgnb J>i69 i 30 m6 i 3 n 36ihfi36g36i. ygmh^nb gibni636nb 030oib333i9ni5 yi
9(336ih3b, ihiR33303&hn3n h3j,363hi(jn30n ai3nb3636nb 5i9m, 933dpni bhgnb 3hm-
U3bn 6i6mn6ymb 9mb33636nb i3nh^36ngi6, 93hnb^39nb gihfi36npn 3&636ngi6 gi
Ri9 myiQn6363Qn 35h3g3&nb 3i3gnqj3h36(5niunnb 039331^16001.
^>6 jncrshjds ihnb 9u36ihnbi gi 9 nbn 6i6np36nb b^h^j^hnbi g i cg36jbnnb
b30np36i, hm930 nu 9n9gn6ih3m6b bi0 i 3X n 9l536ihnb bnjjmijt^nb 5 i 69 i 30 m6 i 9 n
g i aiim636nb 9 m6ib 303rn&nb 3hmi53b3 n (gjnQm^j6jbn). 9 ^ 36ihnb
5,i63ncru.h36nb bb3igibb3i ib33jljb 5ih9migj)36b 3jh3g3bnb 5i9hi3P3&i (gi-
yncgncn bhgi), 9 mi530Pi6 nb 9m9i^36i gi mh&i6m3&nbi 013 Jbm3n03&nb gnqj3h36-
yniijni. bhgnb 3hmu3bn ihnb bfnybi0n 6i6n03&nb 93g9n3n b36bli>i6bn3hn g i bn3-
h(3Xn 9i(i36i. i 9 933ilJ33ign Shm^bnb 939331^16001, 5 ^ 3 3 3 0 ghmSn 6 ih 9 mnj9-
63&i 9 ibn3hn ogrih936 n (5n5i6(j3hn b33&n, b(T»3nb 6 yip9b36ih 33&n), hitf J>i6i-
3nhm63&b 6nmbq)3hm9n agn^mSibnb 3bih9ibihn 9 ihij,nb 93 j 96 ib.
5i63ncnih36nb qjnbnm0m5ni9 36gm{>363hn g i n6g3ijnh363pn bhgnbi g i Ri9 m-
yi0n63&nb 3hmb3b3&nb j)i3bi03hn 03ji6nb93&nb j>i9ni3036nb ai3i0bibhnbnoi
9 ni(p6 ni 96n03630m3i6 6 ih9 i(i3636b, hm903&ni5 3hm6039nb 3imcgnbnn0m5n3hn
ib33jlj36nbicn3nb qjhnig 96n9363pn3i6ni. oi39i5i 636361103 3nhm63&3n &i63n-
cnih36nb 3hnnj3b36nb 9n9gn6ih3m6nb 03bib3& ymî hi9 ihnb b^m"
6n0n. 9(536ih3b3 3haighm30ig 9mJ93gn 9 ngi gi 5^(13 qjiJii)mh36nb 3hoim60nm6i
gi 9)336ihnb 9hi3i0qg3hm3i6n h^j^ni 9xnb3 ihoi303&b bi3hoirn 6ih9mg536nb
93930 i 336ib. 0 i 6 mhi^mhni9 n 9 m030i 33 qjnbnmgrn^nbîB &i6bb3i336noi, hm930-
bili 933dpni gi£3£)9mb i 9 i 013 n9 gjij^mhnb b3CrlC36i. 3incgnbnmcmj,n ndj-
P363cni cghonbn0ig 003953b 9u36ihnb bi3hcnm ji533nb i6i0nbb 9m3036icni 03g-
3n3n u3^03&igfTi6 nb 3nhm6369n. n9g 36ig, hi9g36igi(j j>i63nou>h36nb cgnbnm0mj,n-
nb i3033nb 930img36n digni6 hoi30ni, 63636i3n i 9 3hm6039nb biq33d30ni6n
j)i3biC 3hn j>3033i gng bnhoi3033&b i6yg36i. i j i 9 g 3 ihb3&300 i 5 i 9 m>)3033369i
33h 5igi0ib3b cg36m0mj)n3hn i6i0nbnb bîgnnb bp3ihn. 3hm6039nb i3033>
dn3&nb i 9 gm63b3 bij)9 img gngn 5i9mi5gnQ36i gij,hm3gi i 30ii3h x n 9ii36ih33-
6nb 93 b6 i 30nb gih&bi g i 93^y333m6i9n, hiQ5i6 9 icn 5i6bij)3aih3&30ri 9603363-
0m6i ij3ai ig i 9 ni6 nb bi933h63m bij9 ni6 m6 i 9 n. 3303h 9u36ih3363n i 3033i fi>
(^>h36x r>i {>i6bij)3cnh36non i(j>9mi33636nb, bn3hbXn bhgnbi g i Ri9myi0n636nb
9n3ihai3036nat. 9bnb higni^nnb, ^3933hi(i)3hnb, 5hi3nîynnbi g i Jn9n3hn 5hign-
- 275 -
a ^ o a n b cW3 ^ 06 ^ a o o e ^ n b 8 A ^ 6-a 66300 )6 ^ 35 ^8 a
3615360b 5^3036^ 9(3Q5ihnb 5*6 30 0 1^ 3 6 ^ 3 qjihcnmg ihob 5i6boQXn 8(536^300

bmj>iQn qjnbomcn^nnb bib6i3Qm 60563600. 03895010 0^33680 5i6bo03cni ^ )rr*"
poig 'bmjjo^hcno 3hmu3bn, hoi03Qoii 32)oiqjn'bnoiQoi503h ibi^j^b 6i2iC3^n SD^êCPD"
6 i 3joi6 gi gicn8m6npo.
6sb. 5.1 5ih]8m gjiJ^mh^bn hmjmhu <gn^m3mh9m636nb 5iij^ 03h36nb bi03ic36i. I 3bnb hignnjn-
nb bi5 {|hd0n3m6i, SnQihcnxD^^. bihobbo 91 n6j,35bn]hrri6i; IP qjm^mômpo (Sm^p] PCO/d^dX11
QC3); T G &hi3o,jnjoi; St b^h^bn (bnuo33, brnjb], {>i0m9hm6i, 'b^pSj^n 6yi(yi);
qjni|)m3mh3m6]6n: IAA i]Jbn6n; CK |yi,jmin6n6n; G A 3n6;jhnQn6n; ABA i 6b[jn'bnb 0 3131; JA
sjib0o6nb 03131; ET 30*103611; 3^ 6^3^131101 6183363601 bn6ai3'bnb 19500360 (Matthysse & Scott.
1984; Parthier, 1991 0m6nj3036nb 0mpo<go(5oh363oo 3^ 016^0)
5 .1 ‘b h e n w 6 ^ 6 3 n a i^ h 3 6 n b hD bD C XSni
5 .1 .1 < 3 f li! ) T O n h a n 6 D 6 n b 9 6 n 3 3 6 3 C n 6 ^
0l536ihnb bhgnbi g i 5i63noiih36nb h35xnh3&i bmhô^Qg^bi 0ngi g i 5 ^ 3

qjijj)mh36ob 83833016001. 36901536^0 gjijôihô 5i3036ib ibg3636 0 oi032) X 3 h
- 276 -
93 J 5 h ] 9 x (536360b 5339363 6030103(1363013 153906 3hoi(53b3&b3 (4(136-
Ui)hn3i5nnb3 93 ,4(136693(5006 9333301600)) 9 * ^0101(19063(5(13939 - 901390 oih536ob-
dnb <33(1593680 (cgnljm3mh0fn636nb 93933016001). qjol4m3oih9oi636ob 3ynq|obooi-
901503110 960936390163 o939o3, hoi9 obo6o 3^ 139 36 36 "43(103(1360^ hoigb. 53(13-
9m 3ohm63&ob 30j9ob 8 39935 69363 3(1010 36 9h33390 G30lyri3oih9oi6nb bo6-
013'bflb 36 ])oi6iJ36vj)h3ijoob (53909360b 93833016001 9(5363hob y33p3 636090b 06-
<301(19011363 53h39mb 99501931130160b 9363636. <30(401301(19016360 3j^030hg36036
9(5363hob 536300i3h36ob cgsbob, 53113901 qj3j(4oi(1360b 9oij93g36ob 395090b, gho»-
ob 93 o6(536bo30i6ob 83636380639 (63b. 5 .1). 606330(10163606 93 93608639360b
01(1536016 606393001 obo6o 0633336 bo63h53li(oj,39 36 36,43501606(43(1 h33j(503&6.
cgnljm3oih3 m636o, 53(13901 <33jijm11360136 3hoi39, 360853636 8(5363(136 <bhgob3 93
53630oi3h36ob3i>36, 0633336 8(5363(106 393^33^3(10 9oij9]936ob bo6jhoi6ob3(503b
53h39oi 30I1016360136, 3(135390(13636 bhgob 06^3660301636 93 9o03hoi39363b, 603-
0103(1363013 15390b 3(101(3366, 9301 ,4(136630111,46, 80183(1353636 93 9oi6o90b3(503b.
nb30io 53h39m <33^(401(1360, hm^mhoij 3hob 9 bob (13903(503 (06,4366030163, b33-

dii^DOn 9393953690163, 9 mj93936ob 6365(^9030163), ,43933(13^3(13, 5(1330,431503, j 3-
hn, Bygob 906363, 0101390b b3q)3hob 9360193, 6039350b 6ygob 301^36(50390 93
9hi]3crt jn9o3ho ^jôiho bb33g3bb33533h39 90^9393636 9(j363hob bhg3b3 93
536300^363^3: nBpjjynjhsp - 53630oi3h36ob 3hoi(53bob g36y3&ob 36 9380133-
hj6nb 93933016001, 3.0. ghoiSo 03h3539oh3693g; hsmgj6mdhn3sp - bhgob 3(101(53-
bnb bnfij3h3'b3 93 06^366030163^3 901^93936001; dsqjmhdnhj&psp - 66339^6 bhg3-
b3 (001(103015363^) 93 9o93hoi39363<b3 (^hm3o'b936o) 533936001. 5339360b 3b
bb33g3bb33 qjoih93 3h0i93630i9n ob33 53g 3bg3hoi30o, hoi9 b36oi90ioi 839350
6(133390 03015360b 3(1010169030 90^9393600133 53901633390.
5.1.2.1 bn6^0ncnb 6^3C36^ d ^ n O D h D b flb 3hPH50b06‘bD
0b363hob 53630on3h36ob 3hoi(53b8o 9bnb h3903(503 bb3393bb33 hoig6 36(13-

936b. 3b 3hob: 6ooibo6oi3'bob <301,40^,408393(503 (9 ^6 -- Jgoihoiqjogob bo6oi3<bo
3hoi14mjgoihoiq}ogog36, 36bo83&ob3 93 36^01(50363606 bo6oi3‘bo); bhgob 8083(1-
0139360b h3539oh363 (qgoiôi(4hoi3ob9o); 9 oidh30i6 ob 3(101(53606 63(10133
(301^0(636^03) 93 6360(501(56901 3hoi(5368o 53630013113606 8019933601 6143900b
p36y36ob 939oidh3336390 d3g3 ( gjm^mnGgjjijn^). 8bob h3go3(503 53393636 369366
^D&33^39X e't'63b3 90033(1361501136063 93 qjoih9oh36ob 3hoi(53636b3 (835., jpoihoi-
393b^3&ob 90033(136(50(1363), 33(i39X 9 ^> oih536ob8 x 9016336*63 3hb3& x 3(101(53-
636133 (qjmtimdmhqynfrjdjbn, ob. 63b. 1.34 93 ybh. 2.9 ) 93 36goi5363ho ho^8ob
93(135390(1369360136 3hou»g 5333906363 h0i50ih(5 8(5363(106 9(?3-(p303ho 93 63bm-
- 277 -
a^fnoanb ^ 3 *^ 06 ^ auoG^fnOb 8 foe;6-&b6300 tt(o3 to 83
63(10 hn^Qnb 9*hcn3nb b*03*Q3&* (qjm^m^jhnmgnbdn).

qgm^mjinb^hBQ^njjXie ijiinjhni 9 bnb higniynnb unbqoh-x^hinnbqjohn g*
6ncn30-ibpm n6qjh*6noi30n b33jljhob 6(06*533013611 (ybh. 5 .1). qmijm hjuj 3li)mhj-
b n b hmpb * ^ 30 3 6 3 6 305036^360 - q3n^mjhm9n g i ^hn^mjhmSn. qjn ^ m jh m d j& n
6*h0m*g53636 <33(0(501501633(1,50(136*9 jh m d m lh m ^ jn B j& b , (1(08036015 jhm9n q jm h xn
353qjnb b*booi 93015*336 <33(15(06003601*6 *b0rob 0ymqj ("0*31^30*6 ^3^h*3nhm0b.
nbn6n 0(0(115*336 3hoio3hoi5*hg*j06*g roh qjmhQib: 60013011 b33j^hnb
99oi*6oij030o qjmh9 * Phyr *63 Phy660 , (1(00300(5 620-680 60 -ob cg*hj,0369n
*6bmh6(500b 03033(06001 5*9*0(101336* 6onjo0n3h*g ij^n^h 6001300 b33jl|)hnb
09oi*6oij030 Phyfr *63 Phy730 qjmh0 *g; 0 ^j 600130 b^j^hSo (700-800 6 0 )
*h*b^,*6o03ho Phy730 5*hg*oJ0636* Phy660 O|fnh0 *g. o0 ob 0nb3g3ooi, 013 (101501-
ho* po* g* 03JO 6001300 bo6*oipob 6o0o 9 bob h*go*yn*9 o, cgn^mjhm336nb cgmh-
0360b 330*g5360m6* 5*6bb3*33630O*. 5(103,5rojhoi0 n 9oi*6oij*3b i5oboj3h bn-
6*oi03b g* jn0O3hn b^hjj^hncn 3*b0ni3g36* qj0*3Oi636b - 050 ijô^ho* boi^mb
30033680 g * b*3b3&ob 3(101(501630*80.
<3301,501(131533,50111360 fi*jboi3o0o* 33hoqj3hn30 3hmj,m30*'b0*9o (60018306(1*-
6360). p3hm-q)moi0o*6n 0i536*h336ob bo6*oi0ob (131533,501(136^ 33nh3303b ym3-
0ob*, 6*h3m*g536b 03(0010360, *5(130133 ("3(101360, y3*3o0ob 6*600360, 6*ymq)nb
5*hbn g * oi3b036n. 1131533,501(10 0m0353Q36ob 01(1036(50(13630^ 5*6*6o036ob 633-
33016001 0(536*113 *pn j3*0b bn6*oi0ob g*i530ob 5301636 g*, *J3g*6 5*0oi0go6*h3,
*J3b <33(o^fo0 (og30*i 5nnb, 3.0. bo6*cnpob 3300b 15300036*^13 9o;jt533*gn (133;)(503606
36*ho. 0mdh*m6* bmhi5O30g36* qjmoipob y366ob 33(1393680 ^h^rnhnb 15301103606
6*03*036001. <301,501,5(1013300 9 mdh*m6 * 3‘b(i36330yoi<3b g*i5303pn bo6*oipnb*90n
qjoioipob fo3lio 0 * 03h 3jb3oi“bn(50*b - 000^633* j,m03hro0 nbn bo6*oi0nb 0*jb o 0 * 03 -
h*g 5*9roy363&*b* g * 0*fp*0O (1*90*1500^5*6 5*g*b3h36*b 9 mhob. <3(0010366 *b *-
bn*oi36oi *5h30i33 330om^hm33Qn 0rodh*oi6*. *hj^05*90 0nbo 03833016001
Ubhnpo 5.1 3bnb hipni^nnb 5130363 8(5363113013 5i63nonh36nb ^0^3636^3

(Salisbury, 1985; ICronenberg & Kendrick, 1986 8nbog3ncn)
a I = nSg^Jyii; Q = h3^1936mdi; F = 'bhgi
b B = ^nbqj3hn bn6ioiQ3; R/FR = (j>ni 91 03jn 6n o ix n bn630i(Yib <71363033(1pmd3
c P = qjmljni33hnmpnt0n; C = I5(ih>)ipxn ^3 ^nmQm^n^hn hn^dn
d n6(j3h3spn hipnitfnnb 3n(«36ibi pi hîjunib 3mhnb
ahmyjbfi “6331^^0393- b>vUhxn S^hnmpnb tvji& nnb 935-

6rib bib3 933395360*063 ,yi3n c 303^)63 g
oi3bpnb 5i)pn336i 93 3(^1(536363 I R / FR, B P ben -

P3hmb bn5hd30n thgi Q, F R/FR P 5cn
Qvjhmb bfi3h(5m6hn3n mhn36j>3(yi3 5cn
qjmonpib br>3h(5m&hn3n mhn36^3(5(13 Q.F B 6cn
y333ncn*i Q R/FR C ben - i3nhi
5163(1011(1361 gi3i5hpi33&3on mh536ri3&nb 1r R/FR c
8336361 I R/FR P
boi
0393bh3*61 I R/FR
•jSbr&ybrk bn6oi3'bn 1 R/FR 63
3®bn93bnb 5iiijrr]h3&3 I R/FR
8386(136(16 5r>j36(5nipn I R/FR
- 278 -
I
8 6 «?0U5 e?5 &5630 m5 6 3 5 0 b 6 0 5 7 )^ 5 0 0 5
0nn(p6Q3i y 3 i3 o 0 3 6 o b fcioi&m&i g i 3 io h i j,} b b 6 } (h iy , 0 1130 b 0bhn3, b 3o b 36-

ym6b Qi9v!)33h3ib), 15 (130 133 oi3b03&ob "bhgnb b ^ r^ Q n h ^ & i.
qyn(y T)8fTihc3m5363(yij)3hn 3hmi53b3&nb ghmb bg3&i 36‘bo036ob 5 11^ 0 3 3 6 1 gi
536360b h35300(136}. >,9 ghmb 0mj03g3&36 03gailjmh36n, hm0036nu 33h3gnb 51-
I13916 0m b x bnj>6i036b "oiih506ni6“ 33h3gnb 8051 b^h^j^h^bnbio^nb 5 1 ^ 5 3 6
qjmhSig. 6001300 bn6io>0nb bSjj^hnb 03mhign bibo56i0m 613110101 Ca2+ 918633-
0n (jfipi j)i00 m030 o6 o. 5i6ioi363Qm6nb 15 310 3 6 10 ^ 1 06333b Ca2+-nb Sm Si^bib
^ ^ 3 0 1 ^ 0 1 0 0 g i i0nb 518m, ii0 3 m g x n6nb 5iij^n336ib, hm030nij, ou>3nb
9bhn3, 11^03(1366 51(133330 36^00366.
32jm0m5n3hig 060036301*13160 0m30363&ni ip0mu3636nb qjmli)mo6g3;ji5ni gi
9nqj3h36unh36nb 3hm(53b36nb qgm(jm33hnmg30 n 0ihoi3i. pm, i6 3^y3m
3 ^ 0 ^ 3 6 8 0 6^3^15303^300 0hi3i0n O^Sihnb oi3b0n i(n0mu3636ib o6y3&b
9bm0mg 0 i 9 n6 , hmg3biu 9H530X bo6 ioi039 n gogoi (j>ni 6noi30n b33jljhnb 616-
npn (bo6io*0no* 1(^0^101536360360). i(j0mb<33hm0n pni/03jo 6001300 bo6ioi0ob
83031(10361 03ig536b 1 :2 -l :3 , bmgm 033(1300 (jyob ^338 bo6ioi0nb 83^0 6noi30n
gjjhnb b^j^hn 2-10-33(1 iC130i(i)36i pm 600130 b^j^hb [5 0 ]. bo6ioi0noi
i(p0m3 ij3636036 n i0 3nhm6368n 33h ib3hb3636 i(p0mu3636ib n;ji003 , bi6i0 ih
83^530361 b33jijh30 n 0301053601*161 (015., q3moimp(5336nb 030036)- n6n33
93nd036i ooiJ3ib d^nghm 3ih3nb j330 0mb33ghno 0 ^ 0 3 6 ^ 3 . ib30i 3ohm6368 o
i(^9mi33636nb 3hmu3bob 8363936} 5333006361 I0m6iyihnb h3530oh36ob 03;Ji6n'b-
859, i63 3m3xil5nob himg36m6ob 3ih35x<ih36p}g. i(?3iTH5363&nbicn3nb bi^nhm
bn6icn0nb 6ihj)33Xn him036m6nb 0moibm36n036i<b3 5130361b ibg36b vJ3033hi^3-
hi. 0i5„ lbm5n3hoin bn6ioi0ncn i(n0m0i536360ob ai3b0n bn(503nb 513036001 3qjhm
0j)lid6m6oih3 bg3&i g i ip 0 1^(5363&nbiOi3nb 613036 516101361b 0mnoibm3b.
goljmjhm036nb bnbli)30ib 033d0Oi Q(j>3-^183(1 (503000 5i6ioi3630ri6ob bi&-
5hd0n3m6nb (^n b ^h o h î. gpnb bi6j)hd0O3m6nb 33hnmgo(b0n 8(5361(1300 6 ih9 m-
19536b ib^hm6m0n30, I0n6gnb5i6 gi0m3j)ng3630 bo56i0b, hm0xni5 &i6 i 3nhm-
636b 03g0n3 36gm5363h hnÔnijib gi 0ihoii3b 0hi3i0 3hm(53bb, 0iO) 8mhnb: n?-
3^(536361^ yomh^nb i 0 mb30ib, qjmoi0ob 51801b, gi^m^nb 3hm(53bb, bi0ihi5m
mhj,}6m36ob 03j06ib g i 516^33011136001 y3i3O0m6nb gi6y36ib (^bh. 5 .2 ). y330i
3b q3mljm33homg30n oii3ob363h36i bi03i036ib id033b 3(536ih3b 0m30lbigmb 51-
h30m 3ohm&36ob (5300036360^0131-^. 3(5361(1336^ (11*180360(5 n(bhg36ni6 53ip3nbi
gi 63n0ni6o 33hnmgnb 0m6i(5303m6nb 3ohm6368o, j,iiR6nioi b^h3^j)3hxie e^>
qj36ji5n3hig i8j>ihig 5i 6 bb3i 336x n 03^10*0360 (bjbm Bjhn gndmhajnbdn). b030-
Oii03i'b(n33oi0n 0 m‘bihgo Phlomis fruticosa-h [ 167 ] gi Sarcopoterium spinosum-nb,
i2)li3o*33 RhgO0m3Oi i03hnj)nb 30hi0 h32,nm6360n 6 ;>3hlJ3P02>3Cn Prosopis glandu-
losa-b (bi0oihob 6 ^ 0 g i gogn qgmoi036o q)ml5mbo6oi3<b3hig 3<3hm 30}3ii3h36ni,
3nph3 “b ig b x ^ b 3qjhm Siîhi q3moi036o, hm0036o(j dnhooiigig 6yoob 03030150-
vjnb 3ohm6369o o0ymq336oi6 (nb. 6ib. 2 .26 ).
9l536ihnb 36ihb ‘b3bli)ig 83ihRnmb 5i0hi3036oboi3nb b30biyh30n ghm 5i6bi-
i3Oih3630O 3jjm0m5o3ho 36 0836x^16;, ijgb. h33hmg3jun30O qjibob g i‘b3b(j36ib
g i 3mffbig36ib, ib333 j)hn3lj|mj>i036ob b3mhi65O3036nb g i 5i03(5i65n3036nb fii-
bib3ib b30b 36ym6b yjsjncm bnb qjm^m^jhnmgjQn n6 gjj^nnb bij)0img hoi30n 83-
ji 6n'b3 n. y3i3n0m6nb gi6y36i g i S m y ^ & x ^ 0 8 ^ 3 , nJ536i 013 ih i j^n Si^h n
3nhm636n b30biyh30n, 300163331 013 ih i y3i3n036ob gi30l|)33hi336nb 1^030061
iy3i336nb 031'bnb gi6y36ib, ihnb 013 ih i bi>)0ihnbo ghm oi3b036ob 0m06nq33-
6nbioi3nb, 0^6361 013 ih i 6iymq336nb 6i3i3hi530363on tfbm330O g i bb3. &hd30n
91 3mi03 gpob "0my3ih3o" 3u36ih3368o ihb36m6b y3i3O0m6nb 0o0go6ih3m6nb
20-^3 03(jo himg36m6h030 g i bihnbbm6hn30 3ihoi6ô. gpob ih n ^ o ix 0 bi6-
- 279 -
a^rtoanb a^ 3 ^ 3 G^ 9 ua 6 ir t 0 b 8(f^d-&> 63 om,Xna&<>8 a
5 hdcn3m6nb Snb^ggncn 3hcn9 i 63cnnb5i 6 &i6bb3i3g3&ns6 bibQmdj&n, hib3&n g i

[206]. ibQcnn 0hi3igqj3hfn363&i 6ndg,b idp33b gnqjQhQSuniynnb 3 hm-
U3b]6b bibombnb 3 nj)6 noi g i bjQb 36ym&b 0U36ih3Oii pi 3iOt Qi99j)33h i 3 cni
L>3Clll5nDh y ’iD3fie:Xjnib.
(jbhnpo 5.2 aujBihjoij gjfn^rnS^hnmgnWb]

(Salisbury, 1982 dnboegncn)
SD = SîCD etTJ LD = &hdxn 9C?3
&Syvyab$>rb yhrr^xpn 81536:^3 (Dpnb bi65hdpr>3m6i
n6gi)ur>i cnjbpob 5^63000(136 0b 33hnmg9 n Chenopodium album SD

o o b c n b {(iC’nsQbi Betula pubescens LD
Nemophila menziesii SD
C>3hnb bnjhdjBn “bhpi £)h^pm3i6 81536^3011 LD
Alstroemeria cv. Regina SD
Glycine max LD
qjmcncob jsSc* Kalanchoe blossfeldiana
•gmoncnb b ;jiX 3 6 iy n&> SD
Hordeum vulgare SD
9 ^X 53CfT'3iGor'5 Oryza saliva LD
Oenothera biennis SD
g b 0 i5 h m 3 3 6 x n mh&i6m3&nb 5 ^ 6 3 0 0 1^ 36i Solanum tuberosum SD
Allium cepa LD
y3i 3ncm 6 s Rudbeckia hirta LD
Cosmos sulphureus SD
9g3ghm6nofi y3i3fiQnb b3h6x" Cannabis sativa SD
Spinacia oleracea LD
hQShmgjjuns Bryophyllum-is saxeobebi LD
q jw m c u a a e i <gihamqjmcn0m3i 5n b] 3^36^3360
bi93icm gi 0s(nipn bn3i(ncngs6 SD
i 3nl\inb j,*9mb30i ojbhonmgmoxyiSiSn b3 8(336^1336(1
biS^m gi 8i(Tiico bn8iciTngi6 LD
5.1.2.2 iWaiJDhiiMhnb 6i3CD6^ 6^63DCJ1^hD6nb 3hni50bD6‘bD

bncnbmb bi^Sihnbn himQ36rnbi ihnb brujmybcnb ihb^bribnb Qhovjhonn dnhn-
ODign 3nhm6i. ym33Q U^CiDX bibrujmijbpm Shm^bb S^bibiS^&i ^QS^hi^^hnb
5 i 6 bi'bc 3hxn gniSibmBn, bmpm 6mh3iQ3hn I3bm33c3mj33p36nbi0i3nb bi^nhmi,
hm0 y330i SQîfemQ^hn 013 2,:>63ncnih3&nb Shmijjbn 3ih3 m6 n x ^6 nymb ^hcn-
0i63cnou>6 S^hByd^pn. bhgnb 3hmu3bnb ij^nsmbibi pi j)i63ncnih3&i‘b3
i(!>3hi ibg36b ih i 3nhgi3nh (h^mgj6mdhnjn £1*3036$ - SîbrnpnbSnbi g i 6nm-
bn6ot3<bnb pBpl^nnoi 3<bh3633cynwgnb 33333(^6001) g i 3nhgi3nh (h35X X 3nD^n
3hnnj3b3&nb 33333m6ncn - oijhdmn6pjjijnx oijhdmljhnmgnbdn, cnjhjdmdmhqjm-
t>jbn) 6i3C36ib.
5i3huxD&nli 3ynpmj,nnb ibSjj^nb 5icn3ipnb6n63&ncn, (J3033hi^3hi jg n g jb
‘bDîSCOîb ibg] 5 b iC'3 mu363&nb ihmujb’bj. oi3bp3&nbi g i b3mh3&nb *(pdm-
Uj6jdnb iphgn6sQ3hn 36gi nymb SQbi&iSnbn n3 3nhm&3-
- 280 -
8rt«;oM a^ 63om^fnaini>
6 n b i, hm 00Q 6fH j b 3 Q b ;j6 y in 6 3 6 i b i O f c i t i h g i 9 u 3 6 i h n b g y ^ n g h ^ & i b i g b 3 3 0 g 5 m 3

2 > i6 3 0 0 iih 3 6 ib . q jih c n m & i 3h(5X D & nb 9 j m 6 3 9 ^ 3 6 ^ 3 3 6 9 0 g i n 9 ^ 6 3 0 1 6 3 6 6 0 , hm 0-
036(5 o 'b h g 3 & o i6 gngn 03933^ ( 1^300 (5 3 ^ 0 3 6 ^ 9 0 16 ^ 3 n h o i& 3 6 9 n . i p 9 o i i5 3 6 3 6 n b
(!)3 0 3 3 h i^ 3 h 3 c n n 6 o 3 h 3 i o o b i d i n g q ji h c n o i i ( y b h . 5. 3) . 0 13 9 n 6o 9i 03h n ib D d ^ h i-
iK^X0 'bgîhn g,igin0ibb, bp9onj3636nb 36ihn (J3033hid>3hnb 9 i^ 36 ibcni6 3houg
3 jb 3 n i6 3 6 u n i0 3 h ig n " b h g 3 6 i. i p 9 o n 5 3 6 3 6 n b b n f i j i h 3 b d n h ig 9 n 3o m o » 36 b io n 9 i^ 3 h
3 n h m 6 3 6 o ii6 9^ n g h m ^^dnN bj - i 9n b 3j)O iofn & n 3h n o 6 lj3 h 3 h 3 ( j(i i j n i 9 3 b i d o o 6 3 o n i .
lb iq jb 3 o 9 n 6^9^3360360 ( d n h n o ii g i g , R h g o o r ^ o io b 6i h 9m 9o i 6 n b b ib 3 o » 6 3 6 n ) .
b i 9o iih 3n J > i 9 c « 3 3 6 c 3 6 n b ( h 6 n c n <b i 9cn h n b 0 JO 16 3 h 3 5 o m 6 3 6 o g i 6 ) b i 3o h o b 3nhcng.
Ubhnpo 53 8(536*1130x1 cn^bcjbnbi 9 1 W mhjbnb 5 15 10 3 3 6 ^ 9o6o8*03ho, M3<in9sC 3ho p* 8i J jn 9*C3ho

(1393]h i^ 3h36n (°C ). 3b 8m6:>(53836o h33h3 h i3 8h i3iC O 6 * h 36>6o (5 ^ 10 13 6 1,
Srugijnb (536ns6pn5», lj3h8m33hompo) p i 9n6ij>i6n (6m96nqj36nb b ijip ni. ib iy i. 5i(nn336oboi3ob
oi3bcnb 3 bseymq)6i ) qjiJ^mhnb &>8m 03b id c 32>3Ons 830153100^. ( 8h>3*t>o *3(jmhob 8ob3p3ocn)
9tS35ih3 8(160838(1 m^naodn 8y)bo83an
bmynb b3mh35n
0U3&ih3Oii ^>0101^36360 0 5 15-30 3040
6fbei6"^ bmynj&nb 38hi3C3bm 6i ~5 -25 “35
Oi3h8mgjn03hn 5m p i5,nb bmin3&n “25 45-55 “60
8ih(j3(XTi3636n
Sgjpnib 8^153(11113636(1 S4 -2 0 -30
lbn0n3hn bih^ypob bmhbc33pn ( 0) 2-5 2025 ( 30) 30-37
6hn6jn 10-12 30-37 ( 35) 4042
(jhminil36nbb p* b36(5hni5n i3i>nb €4 ^ 536^ 3360 (8)10-20 32-40 ( 40) 45-50
mhc3&6ni6fl 6ioh6i0:>b36o
l535ghnbs p i 8ic>ic 9cnob 8(536^13360 ( 3) 5-10 -2 0
8p3Cmb 6inh6bcib36n 0 ) 2-5 15-20 35-45
lbm9fi3h bih^y3p3n •>3Qyl3nh363Cn 8(5361113360 1-3 (6 ) 15-25( 30) 3040
ihmioi3&b> p i b36^hm5oi369o i30iJ03oh3&3eo 81536^3360 10-20 3040 45-50
3pi66mb 8(536^3360
bi<3b3Q3 n-5 i(no 335igo 10 20-30( 35)
bi8o\ih0n-5ic>o336igo 0 10-20 -30
viH ^D & o 10-20 20-30 3040
bni9n3ho bihjypnb b3 8(536^3360
60631^36360 4-10 15-25 35-40
g itimmajfrioiQoi3636*1 <10 20-30
gi6iQn ^ 3 9 3 3 h iv 4 3 h n b 3nhoi&363n *("901(33636^ 0 6 1 ) 3 6 3 6 don3h 6x^>. 9boi0o\g

9i 3 n6, hoi(5i ,j3933hi03 hi 1CPC bp39i036b (6ib. 5 .2 ). ip9on53636obioi3nb b30-
biyhjQn ghmnb ihR33nb ;j6ihn b3cb 369016b 9tf36ih336b 3cghm 6ih9i^36nai
5i3h|53cg636 i 9 i 0 1 3 n9 i 9 iboii6 3hatig, hoixo cnjhdmhj-
dJC^nhjcn d jp 6 nbdjbn ih ido3336 bi93id)6ib cn3bob 6MI'n30D^ ihib 3g b iyh x
3ohoi&369n. Rosaceae-b, Primulaceae-b, Iridaceae-b 9hi3X?n bib3m6i g i ‘bro&n-
3hom b3 9|536ihnb (ihyn, iC3ob b3, (5i(jb3n, noji6n, 6id3n, <30^30, (jpni) oi3bQ3-
6n (nn3936ni6 3qjhoi i> Q 3n 0 ig , 0 13 nbn6n 9mb33636nb 33hnmg9n gngn b6nb &i6-
- 281 -
&*6 0 9 [% &*3 C?36 * 91)06 * 6 % 8 6 e;*-&*63 na)*63& *83
9 i 3Cm6 i 3 n 5^6013909636 gibipn ^ ,39 3^ ^ ^ 36 0 1? i6 bi68miC0 yn&3nb 5 i X 3&>^

(601503001 b ^ h ^ n q jo ^ n i 0 -8 °C-ob 3nhm&3&9n). <bm5n3hoi 9(536ih38n ic?801(5363-
6ib b3cb 36ym6b 9 i{nicn 63933hi^3hi: 6hn6jobi g i lb3onnb 3ig 9 nb 0136036^3
40 °C j,3033hil!)3hob 9mj93g3&i 06333b 9 ioi ipamu3636ib.
3353»ji»jn3h0 5^63001^3606 ghmb, 8n6nb“b x i mh5i6ri36nb giymojobi g i bn3h-
153po “bhgnb Shm^bo bb3igibb3i 81536^380 o6y3&i 6i 6bb3i 33& x i S O ^ ^ i ^
hi“b3: lbm9 n3ho bih^yjvib 91536^33680 - 10 °C-‘b3 gi6gi; ^m Sojjx 81536^33680
- 12 - 15°C [ 17 ]; ihjôjjob, 8ipie8orinbi igh3 5i'b}qjb3C<bD 8my3i3O03 9j536>
(133680 0°C-ob q3ihj,p368o [ 116 ]. o6^36bo3h giymqjooi “bhgib 6^0(1936} 633(10
booi6rri. 3jh3g36nb giymqjob m3lio9 i 03ho ,53933(1:^3^, hmip giymqjnb bi6-
5hd0O3m6i 9 n6 n9 i 03 hoi, &33ho 6iQibn3i6n i 3Q33h x n 9b36ihnbiOi3n6 30 °C ip -
633b g i ibgmb ihob “bhgnb 3hmu3bnb lj 3933hili 3h x 9 iJbn9] 9oii6 [6 7 ]. ^hm3n^3-
6nbi g i b36j)hn3o2)36 nb 9 (536ih 3368n bn3h(53Qn “bhgi y s x ^ O n6l|)36bn3hig 9 n9 -
go6ih3n6b 30 -40 °C-“b3 ; gi6ihfi36 8(536^33680 30 - 15-30 °C-nb q}ihj,c368n. <336-
3360b bnj,hd38n “bhgnb ^3833(1:^3(1x0 n6^3h3ign bidding qphcnmi. Ibm0n3hn
bih^ycnb b3 9y36ih3368n qg3b33&nb “bhgnb 9n6n9ig3hn ^3033hi^3hi 2 -5 °C-ni,
hi(5 bij)9 img gibipni. i S n ^ g ih ihnb fcibi^nhn nb qjij^n, hm8 qj3b33&n “bhgib
n693636 33h 2,ng33 ycmh^36 ob i 9 m bx ^>993 i& hd x 3635 63ni6 838 mg5 m8 i 9 g 3
(nb. 6ib. 5 . 16 ). 30}hm 016000 (1350(1163606 8(536ih336b 3bi^nhm36ioi 3cghrTi
9iCfi0o (J3933hi^3hi; 9i5-, Citrus-nb <3363360 “bhgib n6y3636 abmgmg 10°C-'b 3
“b33noi. (jhrOnijXH pi b3&vjhm3nj)Xn 9l536ih33&nb 3qjhf*i [503 h35nm6368n
5i3h(53Q36nb 83“b(T<393i, dnhooiigig, qj3b336ob “bhgobioi3nb bi^nhm ihibi2)8ihnbn
l53933hi^3h36noi 36gi oymb 5 i 9 m633x n . i 9 ob b i 3ohnb3nhmg, 3 3 (^ 3 6 0 6 goq}3h3-
6i50i(5ni 83080361 9o9go6(i3m6g3b i5h3d33 (oi39(ji bidding 6300 ^303noi) gi6i-
Cn (j 3833hivj,3hnb 3nhm&368n. “bi8 oiih8 n, ^1^1*15363(1 ig5nQ369n ygrihlj36nb 83-
a b
Germination temperature [°C]

fo b . 5.2 a,b b b ^ ib b ^ 0 ^ 3 81536^33630 orjbcD&nb 50%-nb 5i63ncniH36nb}Oi3ob
bi^nhm ghmnb {»0<,>3<'e3&OCO&:> i53053hi>53hi'b3. a Rhgnpm30inb 6ih 9(n9m6nb Cariophyllaceae-rh
mjsbnb 6ih 0m0 ig 536o 3J>n (S biqjb30n, W “b:>0oi}hn). 0}<nn 5*63nou>h3&nb bnfijih3 OOÔ-
hi^3hnb 0^36^^006 3huu>g 08^^366; bs3bh3omb 6ih 9m3m6nb 81536:^1301^ cn3b(vj&nb 5^63001^3606
bnfijih3 3qjhm 0sr>cns 0(jnh3 153053^3*136133; b gjhn5>6nb36hn 0(536ih33cm6nb (e> 3 ibibrocn3630*1
bi0bh3Oi b03co>s03s'b(p33omboT3nb) B^h0m8ie536o36nb oi3bpnb 5^63001^36* jqjhm b 6 h ^ > p 9o3g-
o6ih3m6b 839^36001 e^b^Q 0)3a:33hi1i3hil53, 0i 3 o6 hm iji bi03hmqjO03h b3 0 ^ 3 6 ^ 3 0 ^ 013^00 35301
3noibhg36i 0 ^ 0 (Thompson cit. Bannister (1976) gi Mitrakos (1981) Snb^noi)
- 282 -
hnb,j33nbi g* y3*309m3*6n 3hn3pihgo3&ob fi*b*b3* 6311936* 8bm0mg 5*66*i)30v-
I136001 503 33hnmg369n.
¥3*3093606 R*0my*oo636o6 069350(136* 6936* Obmpmg 5 *h$33x t!3333h*63-
h x (i663h3*p3 n, 8*511*8 *3 3 ^ 5 3 6 ^ 3 5*3936*6 *693636 6b3* 63333(1*63(1390 3o-
hm5>36oij. b i 3 cnhnb 8*11530013636^ mh60o*6 h m ^ d X 81536*113366 9* *51130133,
<bmj)n3hcnn b3 8536*1106 ypm h^b (8*5., *6*8n) 6 mh3 * 03ho 93*3091*1606*01306
6^011936*01 63906*906 5113(1 ^hnmgn. nbn6o y3*309m6o6*oi3o6 9 b*g *hn*6 3b m-
Qmg 3*8 n6 , hmu* 8*010 *3nj)i93ho 33hnb(j33* (1*0936033 j)3(ihnb 5*68*39016*80
n3yfnq336* -3°C-9*6 +13°C -893 (6*83*9019, 3 -5°C ) 6383;)h*63hn6 3ohm6360o [86;
165], *8 3111*15366 361*1936* 33h6*90lb*50* (6050306 3mcnbm36np3&i). 013 9(10106
Ungn 33(100190 dpoah 0 mi03*, *6 050 9*0(10333* 15°C-'b3 0*0*90 63333 h*63hooi,
3b 3q)3j(i,n *0*h 3mj03936b. 6n6* *"boob 53mqj(i636ob* 9* 3 *cno 6393^503110
31193606 (8*5., 6*0ob 6386300) 6m0j33b3n q)moipo6* 9* 93*3003606 3hn-
0(^(190333606 ^*6*63* bg3&* 20°C-‘b3 b330cn, boipm y3*3n0m3*6n 33&3^tj>n3hn
j)3663hni6 b*6m0nim 6*8019*90636*6 6^011936* 60*9*5,06 jcghm 9*6*90 63033h*63-
h* (10°C-o6 <3*11503680), h*5 *3 8536*1133606 5 * 3 ^ x 3 6 0 6 *9500*6 b * 8 oi*h3 o*
5*6*^^1636x0 (Sib. 5 .3 ). 93*3003906 603(1530 <bh0*6 93*30901606 93895(113 33-
hnngSn ob33 8*0*90 63833(1*63113611 6^0(1936*. bm5*g*g, 03od036* 001^3*6, hm3
3536*1106 5*63001*113606*01306 361133630013*6(1* *hs 5*^333911 63333 h*63 h x n
m36<i3D3 n, *(1*839 63333 h*63ho6 m36(i3*03hn 3o0go6*h3ri&i.
6*ymq)36ob* 9 * 013693606 5*6300 1^36*6, hm^mhu 63bo, 36*^0(10136* 3qjhm
0*C*>9n 63833h*lj)3h*, 309(13 8536*1106 3353ib*6n3h 6*609366. *00^010 801393 9*
15030 33536*bnnb 63^101606 8^0163 3*6o6*6363n 8536*ho6*oi3o6 3qjhm b30b*yh3-
Cn* 33j,36* 6nI)hn 5*811*3036* (03363(13606 83^86*, 0n6ob'bxi 9^ 8 o6o 6j333* 6*6-
11036*9 9*ymqg*).
lijdDjh^jhnb g(?j-(psdjhn yjc* (oijhdmijhnmgn) oiootjdo6 ym330oi3ob 6*-
b*h{,360fn* 8536*1106 ^ 9 0 6 * 9* 5*63001*113606*01306, bm0m 035336* 3*606*6(16
63933h*63h39 n *8390639(16*980 6*38*019 3*115*9 *hob 5*8^133301090 [2 7 4 ], 3016-
6d63663hn 3908*606 8jm63 h35om636o6 (110183901*01306*5 9 * 0 *b*6 o*oi36 x n*
6j 333h*63ho6 d903ho 903-0*03ho 83^3016*) 8536*113360 9330*b3 3qjhm *j 6 n3-
Flower Develop- Elongation

initiation ment
„ 30
O
0
®u 25
Hyacinthus
1 20
n.
E
a 15-
i 10 ■
E
Q. Tulipa
6sb. 53 Q i 3nnjn6&)&nb O 5■
y3»3nQfnbnb* pi &s63notoh3&nb*ai3nb
- 1 ----- 1------1----- 1----- 1----- 1----- 1----- r-
m3^n3i(vjhn (Hartsema
et al, 1930; Luyten et al., 1932) Months
- 283 -
^ 3^ 06* 3 (3 0 G 6 t n 0 b 8A ^ * - a * 6 3 0 0 ) * A 3 M 83
hip 3non>hQ3&ni6 0i 0 n6, hmg^biu piQob 10-15 ®C-ncn 6>vn6ni QC'n-

bib3- Di^j^boabi g i 3gi66mb bbji 0 |£)6 ih33 &0 n 3b 5i6bb3i33&i 2CrC-b 3 ;)iq-
536b. <bn0n3hn bih,i>y(y>b 0(5361(13011 30>j3bpi6nbion3nb g(P3-(ni03hn 013(1©™-
33hnmgnb m^^nd^dni 5-1 oPc.
5.2 0l3O6^hnb tobnbFlbbeFl 6^63fl(JDh36flb

w b $ > i\
y330i 9 (536ih 3x n mh&i6nb0nb brujm^bQ^ n6y3&i fti0hi303&nb 3hm(53bnQi6;

0ib 0m3y33di 3D6Di!>i l503ho &i63nou>h3di - bhgnbi g i mh&iCmaii Ri0myiQn636nb
3hmi53bn. fii0 myiQn&3& x fT'h5i6n36b3 r>b33 30011(19361 5 i 0 hi303&nb mhîSm^bn,
hfn0036oi5 odc33i 0m0g336m cnsm&ib. i0nai nj)3h3&i ihb3&mdnb 6h36(13631. 516-
I IP T T P
\ ; u
6bbi 5.4 3hoi6(vn3>6

0Uj6ih3<7u> biUujmoboni bfiico
p i dlbt)} jihjQm qjiJj,frih36nb
5130361. I 3 bnb hignujnnb
bihnbbn, n6&)6bn3hm&i g i
bi65hd(yi3D6i; IP gm^mSphnm-
gn; T T
ai3h0pO3hnmgn; MIN 9063(1-
iC^hn 33361; MO Snjhrnmhî-
6n'b936nb >Jir>3<n4>i; Y ( 8^3-
6shnb 6ycnb >vj36unien;
tfjh^rnhnb 3m^36(jnico;
6mgi5nb Bypob iiTV336ynb(>i,
A ibndnQi&jbnb Shme^Jiyii 91
dieiCujSCDî (Evenari, 1984
h 2o mo \ 0mpnqjn(}nh363cn emfthiSi)
- 284 -
8G36^ o b w u o o r m ^ n &d6300 tt 6 0 &0ii ^ 583i 0
3notohQ6nb y330i 36 qp'bi 3 n9gn5 ih 3rn6b 6363on.b3hig 9303(1306(^3630 6 mh336-

3 n, 3(1010 gibngiS 03(nh30o 5ig:>b30nb immh906iU0i 69365 3mh9m63&nb 03033P>-
6ncn, bmgm n6g3unh36i 95 cimgnajn^nhQ&i - g,ih30m 3nhrn636nb 0n3h (6ib. 5 .4 ).
5^63001^3606 01001(11330 cgibib 033bi6i036i 0(j36ihob boi5(nij60o6 ^3-
hnmgn (ybh. 5 .4 ; 65b. 5 .5 ) gi ij36 gi0i6i6nioi3630o 0^306363(136360 013
gj36jun3hn ^153306 6mh336o, hm0036o(j h3&30ohg36i 536360b 90033(136150*03110
iJiyi3fn6nCTu> 95 3ohm&36nb 5^3036001. 5563000(13606 yri330 aphiSo 9^3-
6 ih 3 b j,b ifi6 o i 0 ^ 3 0 6 3 6 3 (10 3m cn bm 36n 036 36n 5 ih 3 0 p i 3 n h m 6 3 6 o b ig 3 n gi g,i-
9 m n h R 33i 3i o i n hm 6n cn. y 330i 9d 9o v
6 3 3 3 i3 n , i 0 a p b ;) d n b 9 0 ^ ( 1 3 0 3 0 * 9 f c i 6 b o 0 3 i i h 0 3 n d 0 3 6 i, h i9 5 i6 6 n 6 i q p fc d o
ih b^xn 3 n h m 6 3 6 n & i 6 i 3 o h m & 3 6 3 6 030951*100 q ji 'b n b 0o0go6ih3m6ob oii3ii-
b363)13636b. ib3, 0i5iQncnig, oi3b0363n bi0ihij,m 6030103(136360b (1363(130b bo-
3633 9i0mj(n93630ni 03r630o 0^36ihnb 333606 h330063; 0063(^3(10 603010-
3^6361101 0m0ihi536i g i y3i3o0m6o6 qjibob 6 mh0 i 03ho 3o0go6ih3(*i6i 3 mj-
33936b 3309g,m3 o o*im636ob boi$pnjb0ob36ihoi6(n6ilb3 gi 663.
Ubhnpi 5.4 bb3igibb3i 9|j36shnb brujmijbpnb bib^hdQnsmbi g i bn^n^bpnb j^>69i3<vo6*3o y3i3nom-

6nb bi5{,hdQn3m6i (Altman & Dittmer, 1973; Van Valen, 1975; Haiper, 1977; Knuner & Kozlowski,
1979; Wareing & Phillips, 1981; Tomlinson, 1990; Brunstein & Yamaguchi, 1992; Lyr et al., 1992))
915365(13360 bntyrybprib J>i58i3em6i3n yy>3ficm- bniyn{5b(Vib

6nb bi55hden3m6i bi65hdQfl3fn6i
(630(16^0) W lm Slm M m (63cy*ief>)
3hofipnS5n 6i>p:>bm36360 hi9p36o93 ijyfos i-ae3
( 1) 2-10 1040
9hi3jc6pnj6n 610^6(^36360
333* 6^j36o 5-10 50 e i 83,40
6-eo 50-100
qjncmp0i5336n b3 8(536^3360 t>3Jl53bon(b
5nh3X bjpen^ ~ 10 60-150
Alnus 5-15 60-150
Populus, Salix 5-20 100-120
Betula 1020
100-200
Robinia 1040
15-30 100250
Fraxinus 200400
Ulmus 15-30
3050 ( 70) 150-500
Acer 300900
15-25
Tilia 2040 ( 75) 700-1200
Quercus 500-1400
60631^3636(1
Juniperus 1020 300-2000
Cupressaceae 1020 300-2000
Sequoiadendron giganteum 15-50 20004000
Taxus >10 20009e )
Larix 1015 200400
Picea abies 2040 200-500
Pseudotsuga menziesii 15-20 500-1500
Pinus sp. (b36^hm3ni3C(V 5-8 100-300
1020 ( 40) 300-500
Pinus sp. fbro9o3hn tah^yQnb) ? 20004000
Pinus aristata ( i c e ’s1® 0 Benj^n) 7 >4000
Pinus longaeva (&tQn 6:>bn6n) 5060 50100
3^0360
- 285 -
aÂ oan b 2^ 3 ^ 06 ^ auaeafnOb 8 6 ^ ^ e ^ 63 nmd6 3 M 83
A dult p h a se
•5-50 y e a r s
Vegetative p h a se Senescence after

3-30 years 200-300 ye a rs
F low e r form ation

and fructification
ct regular intervals v (5)
Growing W inter
Deriod d o rm a n c y
r f (\ l Y
®
Seed dorm ancy

months
Ju venile p h a s e G e rm inaô n
1 -3 y e a rs w eeks
D e sc h a m p sia c e s p ito s a
4 I
f»b. 5 5 gjmcr»Tif6(3336n pi 6iQibo6nb bibniimybpm union P i(j>8m6n5o6n, J 03336003^ ,

IM n a ^ X " - v WiXê 6 ^ 3 ncnih0{,X n SbDSih]. hm 3x°b SO*1 *h *O i33a3Cn. G l-G 3 636-
jhnsnxo. SS b3&b36nc3hn, S bo&icohn (Hess, 1991; Uranov & Serebriakova cit. Rychnovska (1993b)
3 nb393001; £ 3 ( 5 ^ 0 3 ^ Rabotnov, 1978)
- 286 -
3 i)06 *(n0i> u ^ n u n o b ^ n &*63 om*6 a&ob 3*8350
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mh5i6m36nb 900)3(136(5011361, 6ib0oh6y5o65g36nb, ybnQpdnb, (500360b! g i 0063(1-
103(10 6030103(1363606 0^01(115361 Ri6ibibob bbjxQn g i 36gmb!33h00o. i 9 3hm-
1536360b (135x1(50180 gng hmpb ibhx3^>06 q5n^m3mh0m636n - o6gm0d0ih03i336o
(IES), 5063(1000606 8913360 (GA), (50(3^1306060 (CK) g i idb(jo'bo6ob 8313360
(ABA).
fii6ibibob j,i63oaiih3&ob 3hm(53bo 83080361 gi3ibibfiicnmcn 15(130133, (10151*1(1(5
99m63QO mh5i6o'b9ob ShjGs^Qjhn gi jQ jSnb 5 jhnmgn. 81*180336™ onr*i6ob 639b
6y3D0»b 89m630o mh5i6o<b8ob 895^1(130161 3333 y3i3o0m6ob q}i(bi0o, 51661-
3301I136001 djdsb 3 mhmfrj6jb n b ghmb. 516330011(136300 i6 559563(136300 9(j36ih3
6ih9 m3 m6b 556330015(13630 i6 61303619 bo(jm(5b0ob36ihoi6 0136036b. ib > 8151-
Onong, 30obo3&ocn (0i363 5i8m6i6m0j336o) 5^1,330(101300 6 ^ 3 0 03001111366
&nhR3&b, hm00360o(j Ri6ibibob 31113360b himg36m6i 1(1161*1(185031119 9161905.
93bid03630Oi 0mbg3b i5h3cn33, 0 o3hmb3mhm5363<bob gih(j>33336o, hmg3bi(5 yi0O-
69361 0^,3hoi63&ob ihibi30ihobo (111*19361*161, i6 obo6o 8^386003363001; 815.,
6(106306 bo(jo3obig0o 0j>hd6m6oih3 3033600 ‘biOjbx 9 n 0^,3(10163606 <301(100(1361
63(19361 ,33,3(1593606 6^190580 [245 ] (fjn^n ‘bsqjbjQnb b^jhnQ-jhmdnb q )j6 m dj6 ri).
8^3(10163606 600^09(10133 g i b10^,3(13 31(1306 "bhgi 61(18011953636 8600363001316
<31^01(1366 ihi 3ih|3m 6iymqj36nb 5i63oonh36obi0i3ob, ihi03g 306homj,363'bnb
338950100 0o3go6ihpm6oboi3obi(5. 61901033606 51630015(13606 ihib30biyh30O
I0n6gnai 51801633300 03<b(*>3g3i 31(1519 ihob (56016090 036sp3oi6i3o, bmggob d y
3h63m6ibi gi 83^9333016100. 151^33600, 85010 6i3h(5303&ob fihgogmocn bi’bp-
31(10116, gi6i0O ^ O ^ h i^ h o b 3ohm63&9o, 038663x0^ bi0^,3h3 3ih336ob ihibi3-
9ihobo bhgob 030cnb33336o [ 180 ]. 30o0ij,3ho 3oho>636ob 5 1 3 0 3 6 ^ 3 01(1036^0(13-
6 x n 306(1001001503110 5100133033360 od033oi6 qjhoig 06o03630m3i6 00161(530366
8(5361(13011 510(11303606 03bid0360m6ibi g i bohoiXDO&'bD-
61901030b Ri0my1006360b 838935, ft>6ibibnb ^s 6 jn o tih jdib 0ihoni3b '60501-
(3i0n ihb3630o 5363,3033(10 iSihiô. bb3i 0bho3, 306hom6o ob33 08016300 01(151-
6nt8nb 5130361b 51601590b. 6i6i6ibo 5 ih330 mh^,y0 x nJ> ,3(103001003(10 36901b-
33h0noi (hm00ob 53601000 0^,3(10163606 6o0o l/ 3 -oi) g i 151130133, 06,3353036^,001
91 6iymcgob 33g0non, hm0036o(5 6ih0mig53636 03m630O mh5i6o'b0ob Jboi3o036b.
386hnm5363(bob 3hoi(53b3o 3b 5ih30m(53i 516530(1016366 qjo^mjhm036ob bob^,30nb
801690161b oi3b0369o. 510103360b 83^56033(10 g i jo8o3hn 838601^53360 01360b
55^30133 83m630o 01(15160^00916 51913(5301. 88016390 01(151601380 556613301-
h36noi 3,bh36330ymq3b OT3bgob bb3X nb 03636060130b biôhm 6o30io3h3636ocn gi
bb3i h3'b3h33&oon, hm0036o(5 0ib gibôhg36i 0m53oi636ocn - i(n0 rnj3636 obib.
i(n0rrH536360b 3 ^ 5 0 36ihobi0i30b gogn 0 6 o 8 36 x (T'^;> ij3b cnjbgnb bmdsb,
enqjjhj6 (jnyjnnb g n 6 jb g i bi0ihi5m 6o3cno3h3636ocn 3'bh36330ymq)ib. 10 ghmb
386n036x(T1 5130361(5 30 01130b hmgb cni0i0m6b. 3mh3m636ob 5i0mymq3ob 030-
33m6noi 306hom6o 0^g0o3 o6 qjmh0 i(5o30 0 o0 m(530i 3 oi 85m630 mh5i6o<b0on6,
hm8pnb 03O,i6m0o'b0o 033bi6i036i oi3b0ob 5i630ou»h36obicn3ob biôhm 0mo>
bm36n03636b. 3mh3 m6 i 03 ho 3mmhgo6n5oi 3<bh3633cymq3b oi3b03&ob bxodisoib
0m06n<336ob 33homg9o - 5163300111136300 cr>3bo3&ob Ri0miJ336ob 03933016001
0(536ih3 3cghm 3 ^ 5 1 9 I0ihi536b gihR36o0, 6mh0i03hig 5163001^3630 oiQb036b.
- 287 -
&6 ff>09 (% <V>3 c? 06 a 9 0 3 6 ^ 0 1 8 6 ^ 6- ^ 6300 ) ^ 06^83
3 r*i36 nq33&nb QibhxD&nbib, 99n630 mh6* 6 nb9 oi*6 g*9*3*39oli363cn ihb^ftnb 5*6-
y33^noi, oi^bpn nd]5 b g*9r*i33ng360f*i6*b. *9 ghmnbo^nb cnQbtyib 9 ^ 3 3 6 3 6 * 9 0
yn<g6nb 013 *c'3m(5363&nb*oi3nb b*^nhm y330* n6qjf*ih3*i50* g*6hpi3O0n*.
5.2.2 :xrami50606:> Q) Q)b)bcx>>. yrnq}6Whyn<36^
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,30336001. 53I1 bg36* Q63h5nnb g*6hm336* ^Qninpnbnb b*83*036001; h^gjj^nob
3d3H3*036lj36b 9* bo6oi3bob*oi3nb b*^nhr*i 9 *b*0 *b 3<bh36330ymq}b 336 li)m'bm-
cgmbcg^nb mjbogi^ho (50300. 36bo03&ob de novo bn6cn;)(bb 3&od63636 qjn^m-
3mh9m636n (9 * 6 -, G A j^hnb 3*hnni30*b9ob *03ohm6ob 8h38n), *9 ghmb 6936*
b*h3b3h3m 6o30io3h3636ob 9f*i6n0nb36* 36gmb33h99n. 030936 Omgob g*yr*iq)nbi
9* bn3hu3Cn bhgnb 3mh0m63&nb bn6cn3bn (CK, IES), 3hm^m3pib 3 nb b^h^j^^hnb
h3mh5i6nb36i, Oo^mjmBghnxn b36oij3ob* 9* 3hi*i,3306361^ bn5 oi3bnb 6 **Jiinjh 3-
6* 9* 6n0rib o6y36* 013001 bhgnb 3hm(533bn, hm030o(5 6^ 36630*9 6*0mnb*l33-
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bih3b3h3m 60301113113632^01 333&ng*6 j>>gsgnb ^^m ^h m gx 3 3 3 6 ^ 3 . *(T»9 m6 *-
U36n 9^0m3ing3&0m&:>b nd36b 0*b 030936 , h*u qybb 0mrt3np36b 0 ^ 3 0 93 5 *r»3 -
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361*1936* i(p0rTH536360b b*66hd0O3r*i6*. 01536*11301* bm6n3hcnn b*b3i*i6ob cn3b0n
Cin3936i oi*69 n9 g 33h x *9 9 * g**bpi*i36ooi 3hoighr*i30*g. *0 ^n3b 33301360*6 6 * 5 -
bi330ih36n0i b6h*cg*p 6*3(I'n3O^306o (0h*3*0n 6*nh6*p*bn g i 0*h(530f*i3*6n; b3
0l336*h3ou>6*6 - ôhoqjo, *03nb b3 9* bf*i603hoin bb3* 3om63hn 63360). *b30in
Jl333* b30b 36ym6b *(j’3ri|5363&nb*oi3ob b30b*yh30o 3ohr*i636ob b6h*oj 6 * 9 m-
y363&ib.
bm603hoio 0i536*hob (3*p3*hnb36h6n, 3*00360) R*6*b*b36o 6*633f>oi*h 36300*.
*b0*g 6*^663x0 oi3 b03 &n 330(51336 6*633000113630 gi 9 mb*96 oqj3630 fi>6*-
b*b3&b (Anemone, Caltha, Ficaria, Heracleum, Gentiana, Fraxinus-ob 63*h36ob 6*h-
0m3*9636036n). 9h*3*0 oi3bp3n *(*'81*1(53636* obpxD^* 6 *C'n33^>n*J 8306m^*336nb
(033hn3o 6*hbn, 330601^*30 6*3hoi36o) g* b9 oh*g 6*h30m qjijii)mh36ob (0*6-. 03-
jo 6001390 qyhob b33j^hnb 0jm63 fihgopn) 0n3h. y330* b30moibb363630n qj*j-
^mho 06333b dmb3j6jdnb qjsbsdn 6*g*b30*b. *b3ain b*b3m636nb oi3b036n CP03-
g36o*6 d0O3h 6 x * g* *h*oi*6*6h*g. b*6m0mmp, 0*oio 0bm0mg 0uoh3 6i6o0n
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b036nb ghm9 n 6 * 5 x 3 * b 9* *h*b30biyh30O 6*h30m 3ohm636ob 01*309*6 *15003-
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d n b jj 6jdnb aj^bngib 6 *3 fr'b30*b bôhg36i 6*h])333Cn 3ohm636o, hf*i6i*ihni5*i:
bo6*oi0ob 830633* *j*3g3 60*9*69 ° R*qj0 X *6 RhgogSo 0 ymg Oi3b036*3g3
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bog*6 d0O3hn 6308360b 03333^6001 (3 * 6 -. x*^>6mb 3(j36*h336ob oi3b036ng*6).
- 288 -
aooG ôb w u o u n u b ^ n a^Baom^fnaanu âa& o
capbpnb 3^3(1030 fcihbnb g5005 (3 i&-, 0o,i)hmrih556n'b036ob 803(1), 5i(j'0336ob 3 q3 -

6m^3n 603ai03h3636nb giSpb ^jbm3303&nL> b5^0Qob 9 m063 Q3& x o^joô (36pm-
'bmmjmhxn 6i3h(j3Q36nb ghmb) 95 bb3. ^01503(101 3360^333^311 0 ^ 0 3 6 0 (^13-
936056 b5m5v!)i63h5g, ghmnb Jp h i33X 0603(13503600, (15(5, 506501, 55901633x0*
dnd^nb 9m0u30o 9 ri03 ix:)ho 653hcn36nb 9n3h. 63636(103 3nhm6363o a^bpnb
Xp3 mu3636ib 3 3 mj3 3936b 0 hi350O qj5;jomho, nb3, hm3 3hcnn 95 05033 b5b3m6 ob
a b
6sb. 5.6a,b Scabiosa columbria-b jy^noiihQbnb unyvi 63636*103 Hibn^jSn. a 6niQij>nb

hi 25 b9 b n p h ^ ^ ; b {>i63nCTiih;)&nb bnlvjihj. J>i63ncn:>h3&nb:>ai3nb (^ ^ h i^ jh n b 8n6n930n ihnb
8®C. <biqjb3pnb QibsSynbSn &i63ncnsh36nb bnfijih] 3qjhm 9i{picni (a3 > aj) (Rorison & Sutton,
1976)
0^0360 153^03659 5ih33m3n bb35g5bb35 ghmb 0693636 5p3mu3636ib (65b. 5 .6 ).

i(pdm(jj6jdnb gj5bi 95 ^(?dm(jj6jdnb bfisgns 65h0m5g536b 3u36ihob bo|j-
mijbpob 5i6bij)3cnh36oai 95 hd6 m&05h3 3m6ii33cnb. 50 qp'biSo 3m0ioo630n 363(1-
6>!>ni X n g^5ibbh336nb 95^035(15359 95 (P3hmb b6hiqjo 55630ou>h36ob5ai3ob
bi^nhm (i^ftrnhnb 03b5;J36305Q, 5(n9m65U36b bôhg3<bi gngo himg36m6ooi b5i33-
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8(153500 bb35gibb35 bi3o0hpi36ob ^338 (535035, 3^0(138503(10 ^ 3833(15,2)3(15, 60m-
0)3hn Goh3bmh36n 95 bb3.). 6059550b ^giSo h o b i 95 9 obn 8009365(13 ^ h o b 033-
5nb b5j,m6li)5jijm lbm65 5hob 55(i33m 3ohm63&nb y3X^'t|3 9090 l535C365gm6ob 5h3,
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m0 3m0535co ai5m6 ob h5mg36m6 h030 g 565j)5h536 ob 93395^3 gogo Boqo 9 mgob
3U36ihob bo^m^bonb 50 33(101*19^3 (65b. 5 .7 ). (550^3300 8u36ihob 55630-
oiih36nb5oi3nb 95 3m3x5l500b 6i3hL53C36ob5cn3ob 5p3mij3636nb 035^5 306003-
630^13563^5, h5g556 815365(13 0bm0mg 050 o6 3(1(5309365 505 013 o9 356005^311,
hm93b5ij 0 obo 5563oai5(i36ob y3305t 3 05 hd6 m6 o5h3 qjmh95b - cn3bQb 033d0 Oi
53 3nhm6360o 5595(16365.
- 289 -
a*tf>aanb 8 0 3 6 6 6 0 b 8(f)t?*-a*630 m*(fi35*80
Weeks after sowing Years after emergence
6ib. 5.7 ibipkiWigi b] 9(536*11301* 5*63000^ 36*. 9*9^30^ 36* £>* 5*9^^365 (Lawrence & Rediske
cit. Kozlowski (1971) 0nb3g 3noi; Tagawa, 1979)
5 .2 .3 3363i!)^n3hn <33b>. 0dC^3hfl 'bhPflb SOhfiriPn
95 Ri9rrty>pn636nb 3hmu3b9o 9 ymqjo 0u36}hobiai3ob, b i 6 i 9 nb

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6096365 hin33 3i33cnhn (3390036360, 3(3365hnb 9 n6 nbb30i 95 3n6obj3335 05 bib
9 mhnb 3y5h93&5 Qm55 hncn030 -6 h03030 ^ (1395505 {sQmdj^hjpn https). 9 o6 nbb 3 g i
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03j>56m03ho 5^0311165 (cgm^mbnGo^bobi 95 b36atj3ob 9i(j>50o 5j^n3m65, 0n63h5-
03hn 6o3ain3h3636ob 5^03(10 330>30b365). 9^365)13015 ai565b5bm55prri3653n 603)1-
(5300 im6i3h36(500b 3ohm6363o 06903090b 839950190 &39obiai3ob 55958^)1300
960836301*165 ij3b 9o6obb395 95 6060^3385 65600360b 95 3353^ i5n3ho 559(153-
0360b mh556n 36nb ( i 3 m65 yh36n, 0^ 31136 0 95 bb3_) b6 h5qjo 5563001511360b
36ihb. 3353tiMjn3^ <3 ib 50o 69365 <33601,1,033)10 3o5b^nj,3hm&nb5 95 55h30m 3o-
hm636ob599n 9m90qj02)5U03h0 5g53^5(5nob 6o356cn3ob3636nb R50my5006365.
93bc>3930fl bhgnb 365)106 9jm63 mh556m36ob (hmftmhoî qjmoi036o, 93530-
0360 95 65ymqj36o) R50my50o636ob5oi3ob, 33oh3X0b ym30ob5, b5ôhm5: 9 o-
(jmbob 5î(03 ri 6 nb h3 5 X nh32>5 ypnh(j36ob 3)1001*1)190393680; 311019563010b 5 3 3 h gb 3
55695536300 03m010360b (305b^njhm636o) fi5bib35b 3 nhob ihb36xn ghmob
- 290 -
3l)36^ 0b l ^ u o o n u b ^ n &>6300to A 350b 968360
n6i!,3h3iQn; ^h^gnb giyoiqjnb* g i gnqj3h36i50iuoob 3hr»nj3b36nb bBoiho 9 o0go6 i-

(13016*. hm9xn(5 gi8ri3og36xoi bihjQm qjiJ^mh^ftnb 3nhgi3nh (8 i5 „ &i6i-
01363001b*, hi|j b x b 36yo>bb ]3h]gnb iDgCD^nb ^Qib^n^hmbnb “bhgib) g i ihiSnh-
p* 3nh 5*30 36 ^ 3. qjmoi036ob 331139360b h*01936016* 9*80130936300* qjoioi036ob
3hn3mhgn]036nb bngng^'b] gi giymqjnb 36*^33. cgrtoionb j,*9 oob Shmu^bn
^ 3 9 9 3 6 * 3 i 3 n6 , hmg^biy 6*h9oioj3636* ^hndmhgn^ddn ih b ]& x n DShjgaftnb h*oi-
9 36 0 16 ^ 3 10-15-3 ]h 9 ]^n 33*1390. n3 q)mono363n. hnQp^bQiy 6 3 ^ 9 gibipn ^36-
^DîiiDh^bnb i6 8398030 9*‘bo*636ob (83^8*, jihnoi 9i6C333i - QuoBihnb 5*3my-
3 i6 i 6oi6‘b*ob*oi3ob) &*8oi 33b 3oi(r*6n3b 6nh3ip3h txndib, 33h393&ob bngog3
^6gi nymb nb3cnn33, hmj,mh(j 05033 bib3fn&ob 3i$36ihob 6oih0*93h <33010193690
(6ib. 5 .8 ). 331139360b ^315003033(10 8ijbo8i03ho <bm9 nb 3oi6li)hoi0n boihuo3093bi
Zelkova serrata Acer campestre
$>
A 1/32 A 1/44
TH -14% TH -12%
PL +3% PL -1%
PD +19% PD +16%
fob. 5.8 q}mo!(vj6nb 83p*h3&*: 6m5'binb 8u36*h3b* g i (18433 l»b3mbnb 9u36*hnb:> 6363^03
3nhm63b9n. A qjmOTQnb qjihoin; TH qjmcncnb bnbfo; PL 93bhnb3&hn 3sh36jn9nb 3}h3e3&nb bnf,hd3;
PD 03bhnb3&hn 3>h36Jn8nb 3jh 3e36nb 90*83^10 (Komer et al , 1989)
536>ini3hi9 - g68-ob 303*101 9*tj>i90 83815339016* DS^jgnb 6nhoi33n (gi 3 ibi-

bn*0136300 6oi,yioi gi ^hogo 3ibnî^36nb 8oiy3*h3go 0l536ih336nb*ai3ob) 3^(136-
33Cymqjb 3qjhm gogn 3jh3g36nb qjoih8nh36ib [100].
jjh jg n b e^>6jC3$bj 5 * 3 9 3 6 ^ ibg3636: ^3(1501(1390 6533i, qjn^mjhmSn g i 50 b-
oj3ho bo6xngnb *131533,1,01*1360, 3oih0oi6*O3h*g *135390(136*90 (IES, C K , G A ) 33*13-
gnb 33900b 930301*180(136*9016* g i 3obo 830 ^6 0 9 36 30 0 03*^01*1360 (Sift., H+
0016360; 6ib. 5.9 ). 35h3gnb 90115390160b 03**190160010 bhgi - (dV/dt)l/V - 3hm-
301*1(50390* 35h3gob 33900b 391(^3033*10160b 6 019300b (m), 3311390b 803b Bygob
5*dôig*gm6ob g i bb3ifri6nb, *1018390(5 6ih8moJ0636i 5*6*^1)0163630 vlDbj/nhob
3oi(!)36ijoicbi (T g ) p i ^hôihnb bib*<b(P3hm bogog3b (*101830015 bi^nhoii 33900b
5*6303013013 ob; X p ™ 1) dmhnb (3*6 -, 3b3b38bohi3n, boio*3 o, 6ho630o g i
bn3o6g3o 050 3ghob 0 .4-0.7 MPa; [2 4 8]). 2 ^ 0 ^ 0 3 3 6 3 9 0 bibnoi 3b 9*8013093-
- 291 -
s^fnoanb ^ 3 ^ 06 ^ 9 i)aG6 ff)0 b 8(f>^,wv>630 Otortato3 a
-CP
-cw
6sb. 5.9 9u]6ihnb bnfihdjSn 'bhgiSn fiihorjpn bb3igib b 3i Shm^jbnb bjâ^^hn {>i9mbib3C36i
(Cleland, 1986)
630365 5^9mnb^36i 630935650(159 [ 144 ]:
(dV/dt) 1/V = mCYp- T plim) (s’ 1) (5 . 1 )
6990b 6ij)036m6nbib, oD^^hnb boO^nhob 550 m, 33(13911 33b 0630365, 5 6 5 5 -

63030b 3(101(5360 ghmb3 5gh3 699365. 50oom0 535930b 33hnmp9n 33(139360 oi53nb
301036(503(1 boi05b 33(1 5(^63336 (nb. 65b. 6 .6 5 ). 50 3hoi(53b6o qjn^mbn5cn3'bnb
3 6 5 ( 16 5 95 5bn0nQil53bnb 5565600360b 113300b (nb. 0 15 30 2 .3 .3 ) 0093659 5j3oi
66063639^65, (156936595(5 obo6o 5hn56 cbh9oboi3nb b5^nhm 65(560(169506593606
6066^19360360.
5.2.4 6DB3h^l3flXfl y3^3ncn6^ 0} 6^yn<3n6i

3363 il)5i5n3ho bibnym^bom qj5bog56 yj^npm dnb cgsbidn (6mbhgn0ob 56 bo6-
60030b g j5 b 5 ) 5 5 9 5 6 3 9 5 bmhi5O30036i 9001(103660 5hb3&xn 5303503(10 03hobo30nb
hOMOohO&ob 03633016001. 935300360b qj(Tih8oh3&ob 365(10 5580169065(130166 njo-
956, (1018 013001050693(50113630 0(5365(133660 3b 3hm(53bn 069365: 53630033(159
030^0(136300 5b53nb 6o(p633ob5b (3690103(10 o^^D^0): 305bli)rTijhm63dob 5 5 (13 -
33300 h5m936m6ob R56my50o636ob 636935; 3353^51503(10 bb330ob 55(133330 bm-
05093 6i<b(i9fib 636935 (556b5330ih36ooi b505h55oi 01(1556013606) 95 3hmo30-
63bob5 95 65b6 oh65gnb 55601656601(136300 (133000b 30(10163660. 6(15350 0(5365(1360
9353O0m6nb 0693(50(1365 3695 oyoib g 50oi3og3&30 o 55(13001 qj5jl!)Oih36b3 - 5 5 6 5 -
Oi3630m65b3, (33833(15,33(^5 9 5 6y0ob g3qjo(5nonb g5b569nbb3. y353n0m6ob 0io6-
93(50(136300 536360b 555^0^336560 56 933h3bo(i3650o 00165600301636 qjoorn^mh-
0016360 (0501 6oihob, 33(1-33(1016001 3(560160 653(101360) 95 6330301009360. 93530-
00160b 65(105036001 o6g3(5oh36ob 636935 3b 3(101(5360 0009065(130166 66(150359 5 6
55(133330(1 630331163636001 (855., 535030b ghoib; 65b. 5 .10). 935300360b 55690b
630935 93530901606 3hm(53bo 6(1309365.
9353000160b bnb6nh3b3 frshjdm qjsj&mhjbn, 36goi5363h 05h3530oh3630 83^5-
6 ob0 360156 3hoi5g, 0 mj3 393636 6o3oin3h36ioi5 15300b 63633016001. 50 3hm(53-
bob50i3nb b5*fohm 363h500b5 g5 b5036o 05b50ob 006019365 (501(1(50399365 qjoiooi-
bo6oi3bnb, 0n63hi03ho 6030103(1363606 6301306360b, 60105(15536300 6030103(13-
- 292 -
80066 fn0i> b.iboonub^n aa 63 om.bfn3 &nb ? m 3 &o
Bab. 5.10 1536^*03(1 33hm3*9n b3bn0ri3*6 01536*113360(1 y3*3npnb jjmjihnb 5*63001*1136* (Zeller,
1958)
6360b 9 m6 npn‘b ] 6 nb g i giiiQh^b^cn qjmoi036nb gi90nb Shmg^pdob bQOibQi

5i 9 my3636nb bbSîQ^dncn. i0hnj,iQ, ij^n^hn y3i3npm6;> g* 6iymq}36nb 5^63(101*-
h]6* in 6 i 3h35bni9 nb 3363iiibn3h 'bhgiboii6 9} 6n30in3h36icni 0 m9 }hij,36 ibai:>6
(n9 339cnb333nboi3fib, 013 9 mbgi 6 nm9 ibnb gijjihjtfi, 9*5., ubm33036nb Qnph
93^9nb i6 git>ni63630n 6i6np;)6nb Ri9miJ336nb 9 og36ig). 9hi3i06oni6 8153-
6*h3369n *9 nb 3393511* hpShmgjjunnb 36*hnb 9^3300136*. y 3 X i't)3 3<3hm bb3Cn
gi i5n3n h3j>nm636n 9bm0mg j)hn3l!)mj>i936nai ihnb gibib03&30o - 9 icn h33hmg3;j-
Ijnnb mhj)!6m3&nb Ri0myion636nbig9n 9 n6 n9 i 03 hn 9^01601360036360 ij3cn.
jhcn6pns6n d y j 6shjjdn h33hmg3jbnnboi3nb bi^nhm 6 *b9 nh6 ignb 6iohcn3&b,
393^3660^*9, 9mn3m33636 9n9gn6ih3 ibn9 npiunnb ShnuQbSn (3hcn5cn*6
9ih^3pm36369n - 963i63 mh&iBmp&nb q3m^mbn6cn3<bnb 65 %-9 q3; i'bm^nb 6i3h-
cn36nbi gi qjmbqjmhnb 50 -90 % 9mn3m336i ihbQ&3cn 3363i!Pbn3hn 6iBnp36ngi6)
[34 ]. higîB 9hi3iQ phcnBpniB 9(536ih39n onpbgnb Ri9myion636nb gi6y36nb 836-
g36 30£>00)il5n3)hn bhgnb Shm^bo R3hg36i, cn3bp36nb h*mg36m6i g i bihnbbn
gi9 rnjjn93i ) x ni o9 <y>h39 m qjij^mh^bnb 9 mj93g 36 *'b3, hm9036 nu 5 ibi^m 6353Qn
nym 6363hiunx qji,b i3g3 9^nh3 b6ncn igh;j g i 9 nb 5i 69 i 30m6 i 8 n. 93'b(j>3g3cn
i33&nb 3nhm&368o, 0U36ihnb ^n3nb 9nb]Q3ncn, 93ndQ3&i biphoimg ih Ri9 m-
y*pn6g3b y3*3npnb 3hn9 mhgn303?)n, i6 hpShrrig^Jun]^ mh5i6m36b i^ Q g ^ i b*936n
9ibi0ncn 9m9ihij>36i 3363v!Piy''3hn 'bhgnb bibih^jdcmg - 'bm j^h ig^npn y)3b
Ri9 nyiQnb36nb 3hrnj3b8n 9 ymqjn 6iymqj36ob Ri0mu3o6ib (6ib. 5 . 11 ).
mh6on*6 dsQsbj&b (hm9036nu 30(1330 630b 0300^113636 hm'bp^pbb g i 9 n6 nb-
j339i bi9ihi5m mh5i6m36b - 9m9g336m 630b bBhiqjn 5if>3noiih36nbioi3nb)
y3*3nQmi)nbi g i 6iymcgnb j>i63naiih36nbioi3nb bi^nhm 363(15006} g i bi936n
- 293 -
aartoanb cv>3 ^ 0 5 6 aoo66fssou
0ibipnb ^gngpbn 6i6npn 3533 0m0ihi53&3pn i i3°'- 6i0m‘bi0oih363pn hm‘b3^3&nb:>

g i b i 3 bhij,m 0n6ob;)339i mhj,i6m3bnb ‘brnQi imh30i(jnob 0i(pip bihobb9 oi y3i>3-
np3&obi gi 6iymqj3&nb himg36m&iboii6. i3g i53ih ig 33 nju33ni6 dh^j^Q6cn^ 6n
dspsbjdnij. 33<b(j'3g30 n bi33j,3^jnm 33homgob 3Jm53 Sibnî^bSn 6030103(13&;>cru>
Shmg^junib, h33hmQ3;Ji5nnb 36:1 hb gi 0m3i3ipn oiin6nb 3'bh36330ymcgib 9 mhnb
ihb3&fTi6b im03 hm9 nbn: cn^Siji ymî y3i3O0n qjmh0nhg3£>i, 0 ij)hi3 cn^nbcn^g
0150(13 ibn0n0ii3fi3hn 0m53&nb 303(101 gngn Bnpn b0ihg3&i h33hoig3ji5n3po b^hjj-
^3h3&nb fii0oiyipo&3&ib (6ib. 5 .12 ). y3i30Q3Qnb 6i6np36ob 330i5oh3&ob ^ ( 1 3 ^ 3
(nb. 6ib. 2 .70 ) bg3&i 363(150^ gi“boi53i (0oi536i), hiy 3bh3633Qymq)b 0 m-
lbi0oih3 mhj,i6m3i)b h3t>3h336nb bi^0ihnbn 01036016001, h35363(1 iônb
36ihb gi 33&3i!>i1503(1 5i0hi3p3&ib. 3oi0ih3p g i 0i(pip0ainb 633b 0hi3ip60Oi6
6iQibm3i6 0u36ih30n y3i3nQm6i nbp3g36i b i^ im g gngn b6ncn - 0m0g335m
y3i3nprn6i0g3 03ndp3&i 3hcnn i6 mhn 6300(5 ^n &i3ng3b (nb. 6ib. 5 .17 ). ib3cn
3nhm&3&3o ibnOnpiijoob 06 nhn 6 i 0 i^n bij)0 ihnbni 0bmpoig 3363îl5nDhn i6
h33hmg3J(5n30n bhgnboi3nb.
b j d(jj6shjjbdn y3i3np3&nb, 6iymqj3dnb, ai3bp36ob Ribib3i g i 5i630cnih3&i
h353pnhQ36i 36goi5363h bi0ihoii3 03jb6nb036oii6 3hoiig, bii33&n 6030103(13-
6360b g i ibn0O0il536nb 5i6i6np3&nb 03333016001. i0nb 83935^9, dpo3ho 0bb0m-
nihm&i imSjQlîjniSn 03gnb bn3h|jxn bhgnb 3hon53bo>i6 - ibnOnpi^&nb 6 i y
0360160b ghmb 3309350 6pobicn3nb fiinbib3bi 303(101 03i5ig 336D6 i UnD^ln- 300(13
h33hmg3Jun30n ypmh^&o.
h j 3hmpjjtfn5b j i bndnps^jdnb g i 6ib^hyjdn 03nd03&i bi*)0img 0 60 33630 0 ^ 60
oymb. qjn^b i0nbioi3ob b^nhg3&i 00hi0o 0 ibob 3hmg3j^nnb 5- 15 %, b03pcni-
0 3i‘b(p33cnnb 0ihig003i63 03bib - 12 % [ 139 ], 600330b - 20%-b3 03^0, 3i0 0 i
Corypha elata-b 16 % [2 5 0 ], 3i3pb 35 %, boi0oi ô^f^bnb bb3igibb3i bib3m&ib
5O%-0g3 [ 138 ]. mhbib0ni6 0u36ih33&0n h33hmg3^ n x n 3hmi53b3&nbicn3nb
bn0n0i^3&nb 0oioiboi36o03&i j>i6bb3i33630Oi. bh30n 0bb9 mnihm6 nb 30(10163600
i;'ihig063i63 63ftjob Simmondasia chinensis-vfo 0g3ghm6ncnn o6gn3ng36n y3i3-
O03 bnbi gi 6iymqj36nb 5i630cnih36obion3ob 09363636 ibnOopi^bob 30 -4 0 %,
0 i 9 n6 , hmg3bi|j 0i0hoi6noin o6gn3ng3&n 0mnb0ih36 0bmpmg 10-15 %. 3gi66m0n
0i0hm6ncn gi 0g3ghoi6noi n6go3ng36b Smhob 5i6bb3i33&i 5i9oinbi^36i 0n6obb3-
g i 6i6np36ob J,i6bb3i333p bhgi 3 n (bj3bm6h030 gn0mhqjnb0n) [2 62 ].
64b. 5. 11 6iymqjnb idmh^nb 9^3 ^1^093-

Bypob 3m^36|yiic'b3 (Pyke,
1989). 63 9uo6ih]3b3n g^gjmpi^nnb
&i 3CD®i 6iymqjnb ii)fnh(j'b3 nb. Obseo
Water potential [MPa] (1993)
- 294
8U0 6 6 6 0 U l^ u o o n o b ^ n &6630 m6 6 0 5 0 b 368060
6sh. 5.1S h^Shmgjjijnftb g iS m y ip ^ d x j

6i b n d ^ v j'b j (Kaw ano & Masuda,
1980; 9ib i 036n nb.
Manhall & Watson ( 1992)
il)hm3nt)36nbb g i bj&^hmirti^nb nb^cnn 8|j36ih33&n, hmjymhnuii yi3X iii^ n ,

3]hnb bQ, 6gb3nb bg, jmjmbnb 3i09i g i yryjlvjbQ&n ibnSrî^Q&ncn ^b3ig 3rn-
9bhi2)3&nb ^nhmbgbSn, 3bb3mnihm&36 3on]Qn 6pnb 5i 63i 3pm&i9 n. b^'bmB^hn ipn-
a^^nb 0jm6o h35nm6360nu ihni6 3(j36ih 33&n, hpi0036n(j ym33Q 630b b h ^ ig y3s"
300^636 g i nbbi036 6iymcgb: i03nb b3, ^nhnqjn, 33hyi6n, h^bn0i, (jiyb3n, 63-
i3hfibi0n g i 0hi3i0n bb3. ‘bmSr^hn bih^yonb bb3i b3 9^36ih 336b -fi>3$Q 0bb-
3pinshm&i 933d0fiiOi 3bm0mg 3hi3i060r>i6 n6^3h3i0360n ("ic ^ jh S ^ n jc n dbb-
dmnshmb*"): cgmoiQfTi3i6 b336b, hm^mhij 63bn, ym330n 2-3 ( 5 ) 60nb, bm0m BnB-
3m363&b - y<Ti330n 2-6(10) 60nb 938936- 3m0 i h x h35nm63&bi (6 i 3h(j3036 nb ^n-
Q^h^b bi'bc,3h36oii6) g i 3i(r>s08cnian ghmnb 3b n6l|>3h3i0n &33hig 3oghm gngni.
jhm6nv|30n b^h3bnb 3nhm&363n oi3b03&n 9ljnh3 himg36m6nai Ri3myi0n6Q3&i gi
3ioin 5ipn336nb 36ihn 333|jnh3i)30ni.
5 .2.5 boencnhn <syb>. w 6 n 6 <bnanDhn
bn&3h39n 0U36ihnb 33 î 6 m03hn ij^n3m&i 3(330i, iSnj^^hn g i bn b ^rt tjhgi

3unhg36i, qjmoi036n 3qjhm 3|jnh3 bm3nb bg36i, 6ii03&ni y3i3fi036nbi g i 013b-
- 295 -
8 l3366fn0 l> 8( ^ 5-&6630 m6 ffiaM 8 a
p3&r»b (hm9c36biu >5hQOi33, 8150(13 5i(po33&ob ^6ihn ij3cn) himg^Smbi, i6nm-

^ h o b^h^bmh^bnbi g i Sihibn^bob 3 o3 ihoi 5i8dp3m6i b^b^^bi. 3hoi6pni6n
g i ^hcnb^p 3my3i3np3 81536^3360 (9 rn6 mîh3x n) gi&3h36ib 0693636 y3i3npm-
6nb 033 g 36. 9hi3ip6poi6 8(536^336^ hm8p3&bii5 ibibnicr^boi “bhgobi g i y3i3-
ncmbnb 33(100193606 9 rn6 i(53p3 m6 i, 3353^>i5nnb 33hnmgnb 6mpmb 6^6^115303^ 33-
hnmg^p^g £>i6ibp3&ign 6i6op3&o. 8hi3ip6pni6n &ipib3&o, 63^360 g i 63360 bo-
&3hob dohnoiig qjibiao 93b3pib 0693636 3 bmpmg 3 i 9 n6 , hmi5i 8hi3ipn ^n^pob
930gj)fri0 g3j)hignhg36i iSojpp^ho 33hob^30i. b3 8(536^3366 933dpnicn bo^m-
Ijbpob bi6j)hdpo3m6ob 3hoio 33bi33go i 3 q)i'6 i 9 n j,iiîhm6 - dpo3h b6n3h ^ 3 -
363n gi&3h363pn g i 6i6opm6ho3 9i3giho 63360b himg36m6i bij>3 img gogoi. 63
81536^33606 ibij>ob bibfîho 3oop633i n3 938cnb333i8n, hmg3biu 0^1(^3936^
i 9 m6 iyihob 5i63noiih36o6 g i 2>i06o30nb 9 hnb giymqjnb 36iho. 3hmg3j^ n x n
qjmcnp36nb 3 ib ib i g i bi3hcnm 3 ibib 9 mhnb ihib3p6iyh3pn oii6iqgihgm6i 803-
01001366 0 8 ^ 3, hm8 6 ib 9 oh6 igob 6ipi6bn 3hmj,h3b x ^ e 3^h3bg3&i g i 6ypnb
^hi 6 b3mh^n b3 3(536ihob bb33pob 9nh33p 63hôp369n ihibij)3 ihoboi.
gi63h3&i g i bo^gnpn, hm^mh^ 63bo, ihob bnymubpabscnjnb psd^bibnscnj-
djpn 0336^)8360. y33pi 3hm5h3b3po “bhgnb 36ihob 8^0163 8(536^390 ihb36m6b
6i6op36o, hm9p369oi5 jbm30p36ob goq}3h36i50iuoi ohp333i gi o6y3&i gi63h3&ob
3hm(53bo. 31,33 i(p9mi53636nb < 3 ^ 9 0 bi 69 mj)P3 bo^mybpob 9jm63 3 3 1 ^ 3 6 0 gi
Jbm3op36n (8i&., cg3b3ob 63b3b3&o, 3oh33pigo jbm3op36o) 3 ip 333 j)3g36oi6. pi-
CfD^Dpo o6oi50ip3ho 3 3 ( ^ 3 6 0 5 ihgioj3636 oi6 8j)3gih j,i 3 îh g i bi0ij,h 3P3336-
j)36ig. 3nh33pigo ogmcnp36n ib333 3ip3 y3ncnpg36i g i Ri8nn5303g36i. bb3i cgmo
P36n, ob333, hmj,mhi5 y3i30p3&n gi 6iymq336o, 66hiqjig 63hg36oi6 g i 6yg36oi6
p>3hm36b. ijô^ho 83^»6mpo'b3ob 8jm63 6i6op36ob 0336^03(10 93<b (p x 3i 6n-
6ib6ih gi3hri5hi336xn gi&3h36o6 ihm^bob 83933^16001 ihob 3i536ihnb 8hi3ip-
(5pni6 n ihb36m6obig8o 8353360b 3(101-3(1010 <gmh8i 8mj,p3 bi3353î^nm b3'bm-
nb 9 jm63 11350(1163690.
qjmcnpnb gsbjhjdnb 3hm(jjbn 93odp36i 6ihn8ihoimb bb3igibb3i6inhig (6ib.
>.13 ): 3hmj,h3b3pn gi&3h36obib (b3j^3gi63ho gi&3h3&i) qjmoip36o y3naip93S>oi6
g i 8ioin &i9 pnb ghmob 93bi6i8obig. 3haighm3po gi&3h36ob 938oib333i9o 663^
gib b 3i 33homg9o 5i9popo qjmoimpo 3hoighm3pig y3ncnpg36y 93g8o3ig 9 n'bih-
's a t f
%r 0 *
0 $
m
b
e
6ib. 5.13a-e (jnSobn (ipiS^nhjbxoi Wareing & Phillips, 1981; Longman & Jenik, 1987
3nb3e3no1)
- 296 -
9 0 36660 b w u o o n o b ^ n &66300 )6 6 0 5 0 b 368050
gn 8(336^3360 (9hi3iQn 6ioh6i0ibo, 'brôghcno ^hm3nj)Xf' qji63hoiq30ô), hoi-

j,mh(3 63bn, 303^30136360^6 b3Ju3Qi63h ^n3b, boigoi 53^1030^3690 0301010360
3hoighrrixê I53n3i - ^3 6^36^3360boi3ob gi8ibiboiai3630oi gi&3h36ob i8 mh
^n3b 6oihob ihb3&30n y330i 93bid0m 3>hoi6ô.
psdjhj&nb Zhmujbnb bmhi5O30g36i 6363(|)Oi3hig (9 *6 - *JiCDh36jn-
3nb 3jh3936ob goqj3h36(5oh3&nai; ihb36oi6b ^5^30133, gb63h36ob 5*001663330 53-
6360). am6 mj)bh3x 81536:4133660 bo&3hob gi6y36ob bO56i0O Qmgob 8016600)363-
0n oi3b036opi6; 013 *8 ghoib 8(536ih3b 030101036b 8m3i6oih36oi, 6obo boôiuboob
bi65hd0o3oi6s 0 ^ 9 36 *. 8hi3i060Oi6 8(536^33660 *8 3hm(53bnb gi6y3&ib 6od5b
x)033b 5^3801 oji^yn tô - 801503 9^3360, b e^h xn i53833hi,j)3h3&o1 b^h3b30n
10^3^150360 (tfbh. 5 .5 ). bo&3hob 2,0i0ihg06i(50ib mh6 i 6 o‘b0 ob 90163^3 6ih-
9ihou>336 q3n^)0i3rih8m6360, 5i6bij)30ih36ooi i6b(5n‘bn6nb 6^3360 , 30100360 gi
bb3- gi&3h36ob 938iR3h3&30O 801^83936* ij3b 03010103660 (50^01^06061^ 0XPX?
i)m6i536l!)hii50ib. 6ibgi i6nbi, b0i503hai 81536^36(1 - i]jbn636b g i 3o63ho0o636b.
bndjhnb qj^b^dn 8ymq}0 32^39360 oii3ob3&3hig o[5303&oi6: j0oihoi30ib^368o
o303&i bj)hoi8ob ^01^306360 (RuBP-j)ih6mjbo0i'bi) gi 538g35 - Jhoigrnqjogo,
jh3&i (i)o0iioiog3ho b^h3j(5^3611 g i 53hoi6^m30ibj)368o 6ih3fnoj9636i 30ibôi5-
0016010360. i5O^jfTi30i‘b 8i gi 5*6^530^36001, 36goi8386hi636ob bob^3di n ^ jb -
63&i; <3mbqjm0o3ng3&ob qjn'bn^hn g i joSn^hn oi3nb3&3&ob 6315300b 5i8m 600183-
86(1*6360 bg3&i 5i8ôi0*go 0016360b, bb6igo 6*b6oh6yi06ig36obi g i i8o6oi8-
g>336obioi3ob; 6 i^>30 oi6 b Snghmpib^nb, 33hoijbogib36ob, 30100033601001^09^3-
6nb gi 3(101^3*^360(5 *^03016*; jji^ ^ o ^ h o 36‘bo836ob gi5hoi33&ob 5*801 3^ 3-
mpo od36b 0obmbm8nb oi3ob3636b. y330i b ^ f^ t o l^ o 303836,50b gihe»333i893
qjmoi036bi g i 6iymqj3&0o 9i^ 30m6 b b36oij3ob 06,536^3016^
3b 8013036360 0633336 yncnb dmcnbdnb gihp333ib. hoig3bi(5 yngob
gi90ob 3hoii53b36o 3qjhiri o6^36bo3hoi, 3ogh3 8nbn bo6oi3<bo, 63nd03bi 8oibg3b
Uhhnpn 5.5 giiynhj&nb ^>35^3036^ bnb ajmcnpnl) Qib^hQb^V)
9i3ifijsh3bxo 3^6x3^
9bnb h a ^ n y ii
<3bh-nt> ejiynyxyi
jhdx° ojr'yy’ iSînmgn X
v
A
qjryyriSjhnmgn X
x
X
arjirjjn *1631
dcrrihn bntfb], s6 yn&3^ Y
A
6«n X
bnSGfoo X
1*160^(133
arOju(<jin gn jw njhj&j&n
i'bf'yyib bibj)^6n X
X x
ane^hicjbnb K)qgrxyvyi
Zn, Fe, Cl-rtb bn^bhb] X
6niQif^> cWi ■jhn g sd cxh fis
x X
b*rfn
X
OomcjBn
amaBsascx0
- 297 -
&6 6 3 8 ni> &63 CJ366 9 0 3 6 6 6 0 1 8 6 ^ 6- 566300)6635680
bbBbgo b9o6m09i336ob pb5hm336b 9 bom 3moibm36o036ob (536^3600 (oi3b03&n,

b9 m6iyihn). b3 fcboai bp36i qjmcnQnb i$o0ob oincnjdnb 60%-b cru>3npb6 5^801^363-
6b. nbjom 06 o3 3 6 3 0 *^ 6 0 303036^360^, hm&mhoi5bb qjmbqjmho, A m ^ n
gb 5(T)2,ohpo. 0mo»0ob 3hm£)o636ob cnb6gbcnb6m6ocnn pbSph b3pb ]6yfn6b 6(33-
6bhob bb3b 6b6o03&ob dm3bhb536bb, hbbb(j gngo 06o036x't'&^> ^>J3b 9u36i»hnb
603cn03h3&bcnb U30 ob 3 im6 m3 o3hfn6 obbOi3nb (2^6^330*36001 ihbb3pL>iyh3 0 3 rv-
hm63&0o - 63b03b, yo633&o). b3 oi3b0bb'bhnboOT 5bbb&36o bg3&b qjROi036ob g>by-
3001036b, hmjjfTihu 3m3036b (g3gb3o6b‘b3 ym33060O3hbg jh^bb 1.2 . 109 ^ j0m-
hmqjogo [92 ]). bbj03 3636b bhb 3o5036l536ob yri6b3h3bi50bbI bhb03p jgmhmqjno-
ob 3hm^3o6363o 030b3b0 bbm^b. 3hm^3o636ob gbOgob 93p 36bp &i3mCTu>3<v
b3q3093&b q)m,jmgo6b3o3ho j0mhmqjo036o. b0bb 3m3y336fnpb qjfn^mmjbogbuni gb
33h3go gbO(j>3336mgb 8nbj>b6 bhbbh5360m 60301031136360b J^6fn^b6b9p3. b6mh3p
b0o,i)(n0, 3o5036 (j36 nbb gb 3hm(j 3o636 ob 3R 0 ^03 jbo 36gb gbO0b0mb 36bo03&nb
33033^6001, oib6b(5 6bh.J3330o oii60o3p33hpi6 oo* (6bb. 5. 14). 3fnhqjoho6 ob h^m-
0360 o3bb3h33b g i gb00ob 33636x0 3hmp3j<j36o (0bj,., 6hogo30o ,j36hb3ohm03 &n,
0o3mqj3'b(io6ob ^nSnb 6b3ho»3&o) bjô^hbg (j,hb6b3rnhlyihg36b 3bi,3mp3n. 3b 6yb0-
3 o bb6bpo ib^b6m0o,i>36o 3q}3hmb. b6mh3p b0 o,yn3 Jh3&b qjmaipob 363b63 cg^hn
333b0mp. pb00ob '6^503(10*0 ih m p ^ n 5h6oupob 003^3^1536(^6 ijn b ^ h gb
363 b63 b33^jh0 o, hby 63 ^ 033^ bb03b03&ib j 0 (nhm30 bbôb pb63h3&ob 3hm(33bb
gb3ij)3nhp30) q ^ m h r ^ ^ h o b bb03b03&ooi (6ib. 5 . 15).
Phytol chlorophyll In
ihylakotde
Porphyrin
Chlorophyllid
in stroma
Pheophorbidd
OOOO RCC
RCC ■Reductase
V
Proteotysis
\
oooo FCC
Chloroplasl envelope
ATP P Cytoplasm
Tonoplasl
Catabotites Vacuole
(NCCs)
6sb, 5.14 qynoiQnb fc^noioo&bbobC Qhcn^g b^jb nnb

(Matile et al., 1989; Matile, 1991)
- 298 -
a 663 n m ^ o 5 ou i 38n 67)ffin aoa^nG^fnonw
Wavelength [ nm ]
Bbb. 5.15 Jcmhmqjnpnb gbSpnbib gjp]mh]b(j36j5nnb b S^ jîfjh ri b30n03^D^n. (1) 6mh9ip]hn 863163
(gmoimQn (aisfoqphgro&a: jQmhrnqjngo/Jbiejjrnqjngn+jjihrn^nSn = 4.6); (2) pm 863163 gmaimpn
(cni6ig)ihgm6i: J(>iihm<3 nen/fl5i 6^rn<^pn+iihm,jn6n - 3.5); (3) 0my3nou>0m qjmcnmQn (01161-
gjihgmi)!: Jomhmgncn/Jbie^niqjnoo+jjihni^nBn = 1.9); y3ncn3<yi gjmonmcn (<jii5io)ihpiV)i: jpfnhmqjn-
cn/jbiejifngocn+iihnxjnen = 0.4). U3Cf'COiO&n &a9n633xni Jcrihmojnenb 333(jnh3&noi g i iijjiftm-
pnj-jbnb ojp?T'h3b^36(3nnb 51'bhgncn (Lang & Lichtenthaler, 1991)
5 . 3 'bhQ flbl) 6 ^ 6 3 n a i^ h D 6 n b bO’b F lB D h n

an a0 n 6^h 3rn 6^
5p63notih36nb 3hmu3b0o 8(336^3 353361 5161013630^61^, gpob bi65hd0O3m-

6nb, vj)3933hi^3hnb g i 6i03j36nb 60O3h 33hnmgn<b9b. 10 ghmb, 51630011(1360b
bb3igibb3i qjit^bb 0mhob 5igib30i 5333006361 hmf,mh(i b30bsyh30o 013 ihib-
30b>yh3cn 5^381*1 3nhm636nbig3n 0353360b gi6y36ob 195000. bSohig, 16 1 013
00 ojs’bob gi9oii3h36obib, 16 gi6y36ob 6n6 195000 ij3b 0 ^ 3 3 6 360b ^homgb
(hfnj,nhi5 &>6< 013(^03600 xp0m^3636ob 6061 0mb33636ob ^homgn, 16 93130001-
6nb gi6y36oboi3ob bi^nhm 016^536300 cgij^ml^nb (jgi). 60691606 gi3hm5hi03-
&30n 0mb33636ob ^homgob 5 ^ 3 8 3 ihb36m6i 033d0oiai 0bm0mg 00 8(536:^336^
hm0036ou od303630n ihni6 bujoijmijbom 50500 gii0oii3hm6 8 mi03 ghm0n.
5 .3 .1 6 i6 3 flC T D h 3 6 n b 0 n 0 9 n 6 ih D n 6 n b 3 ^ h n ^ D 6 f l
5.3.1.1 a n e a r^ e arrw hen ai3D6ih036nb bubnisrasbom union
5181^53301000 bjbm 6jhn ^Qnds^nb ( ^1036300 - “bi0oiihn; 5310 31 - 63n0ob

33homgn) 0jni63 h35om63&9o djgdnjsg dmbshg d(jj 6shjjbb 3691 3jni69301 bo^-
- 299 -
a ^ o a n b &63 c?aG6 8 Q066 fn0 b 8 6 t? 6-&663 nai6 ff>05680
mubprib 801303 bi65hd0O3m6i. ^3013601, 0151000119, 3gi66mb OôSjh^&n. y33Ci

3(10160016 0(3361(1380 bibouoiybcoi cgi'b36o 3(1018163016 36933*510 0oy33&i. i(p-
001(33636060116133 yiQo6p36i 30(13300 0301010360, bmpn 931300360 080361 38m y
03b 3igi8o. i0 30(10163680 00606^391 0ibob "bhgnb Shm^bn 361(330361 h363 &-
3(11(50390 01(1516013606 Ri0oi910063616. '601503(101 bib3m6i8o y3i 3n03^,r,b Ri0myi-
006361 bg3&i 0U36ihnb 0o6ob'b3pi 61600360b 1^03(10 51630011(13606 330935,. 61-
ymcg36nb 0m06nqj36nb 33(1001g3 o ‘601533(1 9306096361 gi63h36ob 3hoi(53bob pi&-
9360b 60363611. bi6oi0moig, 0oi30n 0(3361(13 530361. (16361 ©bmgmg 0131)03611,
hm0036nu 009010336016 60163363606 cgit-iSo g i 30016 51(^1133606101306 b3obiy-
h30 30(1016366.
n0 (13500163680, bigi(5 8oixn 60nb 5i60i30m6i8o b30biyh30n 3ohm636oi (*53-
6ni6 li)hn3nj)363n, lboi0o3ho bih^yoob cn6n0 (135001636311), 53639361 0hi3i060ni6n
0(5361(13360, (10100360(3 0300n3ig oc6hg36oi6 g i 909 bo0np036 ic ’63336 (015., 63-
5300(1360, 150^9360, 3i003&o, 9 ip i0 6 i0 ib 3x (T'£)nb 61(10016^5360360 g i bb3-).
5 .3 .1.2 33hnnG3W G afTb^hQfl ai5D6 ^h3 06n
633(10 0U36ihoboi3nb gi0ibibniOi3630Oi bhgnb sj^njhn gi dm bjj 6jd n b 3jhn-

mgjbnb 3(10103(101(5301 (36pm03ho (10*50300161). 3b 00130361 51001061*5361 0(3361-
hnb bb3xnb bn5hd3-bo5i639n 33(1001930 'bhgiSn, qjmoi036nb 33hooipx (530180
gi bi0ihi5Oi mh5i6m36nb 33(1001930 91(501-513636130. 013108061(53001 33(10019300
“bhgob Shoi^bo ^01503(101 Fagaceae-9 o (6ib. 5 . 16; 0i5., 03bob *5o3o) g i 61163-
01363680 (6id30, 030^30, 601^0). i0 0336;)150016103(1 *5033680 igh3 5 i<biqjb30,b3
i0m6iyihob 55630011 (1360b 830935 605(1^380 “bhgi 6yg36i gi 5(^399361 8015301-
6361101 - 03mh3 6od5nb 830935. t>01533(1 3 3 1 <biqj63o3n 195090 ij3b 61901033606
9101*536001 5i63noiih36ib. i03nb b3, ihyn, (3113630 g i (5(13 iii( 3ni 3353ii)iiin3hn
bb330ob "bhgib 100113(13636 3i3bnb 51(1333 (61b. 5 . 16; 1030b bnb ô3n). b3'bm-
63hn ^0o0i(iob 0joi63 (13500163630 51630011(13606 33hooip36o6 ghm3 o 00161(530301-
61 (135390(19361 qjo*5oijhm03&o6 606*53006 (0301*501(131333*501(1360) gi 030,5oi3mh9 m-
6360b (bibo56i0m bi93i036i) 03333016001. ib3, hm0 3hoio gi 00133 h35nm6ob
0l336ih336ob 3353*51(503(10 1^030161 0b5i3bm. <33363360, 1101501(115 63bo, 'bhgib 06-
93636 y0mh(j36nb I0nb30i0g3 gi 3b 3hoi(53bo 53016 330rng5m0i0g3 5hd30936i.
050 9i0inj)n9363poi 6019150b *53033hi*53hi'b3, 6y0ob 0ihi5bi 91 0 ib 9 o 6^3360
6n3ain3h3636nb 5161600361^3.
j)hoi3oj)36bi 91 b36(5hm3oj)363o *53033hi*53hob 16 6i03j36ob ©150(139 00101*53-
61^313 j)0 0(5361(13360 h3i5o(i3&36 1^03(10 'bhgooi 16 5i63oonh36ob bb3i 3hm(33b-
ob 511^*50336001. 013001 03g0O3ig *536016 ^hoi3oj)363oi5 yi 0ihig363i63 63360b
0bm0mg 20% obhg36i 03g0O3ig, pi6ihfi36obcn3ob 91016160101363001 51630011-
(1360b 313^360. 0036391319 n0obi, hm0 *5(101305300 *593360 01(1190631631, i j 536-
39361 (301010100(53360 63360(5. 303153619 63360, 16 0ioio (5iCi33Cn ^3^01360 01130-
916 0030101036016 1110936003 ptpnb 5 i 60 i 30 m6 i 9 o. 0ihig ^36016 ^hoi3o2)363o
6gob 5i 60i 3pm6 i 9 o ‘bhgob 33hooig36o 3(101^33 83^01 g i 3(1010 30133011300b 031(15-
93680 (<boi553h 30 3(1010 06903090b 031(1503680) “bhgob 3hoi(53bo bij)3ioi9 b3 ohig
3hoighm3Cip ih 0n0go6ih3oi6b. ^3010 6y33ô0o 'bhgobi gi I0m6iyihob 33hom93-
00 5i630onh36ob 03833016001, 0(5361(13 01130916 0150036b b33(5oi0 o'boh36x n 063-
(1360b 16 3ihi'bo(j36ob 510m qj(Tiai036ob 001901619 9iiih53ob bi8o8hoi36ib.
- 300 -
8 6 ^ 6 &66300166050 U b08n 6 D6 0 808 eJ0 6 6 6 0 n &6
Quercus-Type Populus-Type
Populus trichocarpa
Larix decidua
A Bud opening begins ▼ Leaf yellowing begins
Bib. 5.16 Bpnb 5>i69b3(xn6b3n pjhmj&nbs g i <3 Db336nb 'bhgi uofj^hic^hn ^hmSnb bbsigibb^
b] 9(536sh33n (Hoffmann, 1972)
5 .3 .1.3 hoshm enjunxn u rw obn
hmg^biij 8(536^3 y3i3nQm6nb qjibi 9 g 3 9 ni(p633 b, ^ h n m g x 33636 ^15° ^

bhgnb 3hm^3bb 3 8 ^ 3 6 ^ h^Shm^jijnxri (5nj)Qn. 9(j36ihnb ^nSnb 0nb393ncn 3353-
Ûf^hn bhgi pi h33hm93J^n3Qn ^n^pn 9 n00 n6 ih 3m5b 3hcnghmx^Q. ^6 3hcnn
S^mhnb 930935. 3hoighm30i9 3b 3hm(J3b3&n 9n9gn6ili3m&36 3hcn6pnb6 g i &33I1
9399030 bhgnb 9jm63 ^hmSn^Q 9^36^380. 98 hi0 n h3j,nm63&nb, ^ ^ 30 133 bi83>
0m Qi> 9 i(T>iQ 5i63Q3&b3 6i 3h(}3036x11 9 (536ih 336nbicn3 nb Qi9 ibibnicri3&30n:>
y3b3ncm6nbi g i 6 iymaj0 bb9 mnihm&nb 9m6i(j3p3fn6i 3363i!)i>l5n3h bhgiboii6. ib3-
cnn JiS33i 9306119636^ i {^130133, ^hm3nj)369fii5. mh60n:>6 hmbQ^Q 6i0ib3&bi gi
j,3mq)nj)369n 3353^(503110 g i h33hmg3J y n x n <3ib3&ob 9 m6 i(5303m&i 5 ^ 5 * 9 ihnb
5 i 9 mi33cnn0n.
93 hip ^hmSniX 9 ^ l^^h m Sn iX h3j>nfn6369 n ihb3&m&36 9(536^3360, hm9-
03&n(5 y3i3O03&b 03001^3636 crjmoim0(5336nb 839935, 3-n. 9 i 8 n6 , hm(y> (P3hm3&n
3q3mcncmg hPi365. iQnb 9i5i0ncn36ni: Erythrina-b, Bombax-nb, Tabebuia-b gi
CaesalpinaceaeAi 9hi3ipn bib3m6i. 013 3333 gnqj3h36unh3i> x n <3moi0ni6n ypmh-
^3&n 339g35 9mb33636nb 33hnmg8n(5 hR36ni6, 9 i 8 n6 biJ93 63ij3b igh39my3i3-
nC3 t!>n3nb 9(536ih3boii6. ijitf y3i3nc3&n 3noiihg36ni6 8 n33 x (PD^^D- ^ 3"
cgmcn30 i 9 g3. i 9 ^n3b 3i3cn36nb bm9n3hn bih^ypnb 633I™ qjRoim09(5336n b3 9(53-
6ih3, b^bnon g i t,36j)hm3i6n 63^360. igh3 5 ib iq jb x nb 3qj303hmng36n (bb3i-
gibb3i Ranunculaceae g i Liliaceae) obi9onli3636 5i80nboi3nb 3b i biy3i3nQ3
yQnh^36ncn. nbn6n 6iymqj36nb 5i63noiih36nbioi3ob ny363&36 b i 3 cnhnb &mQmbi
gi b33&nb b h x 830^0130^993 ihb3i>30 9 mj)03 33hnmgb. ihj^nôbi g i 3ipi0-
9cnnb 3i3hi3n 6ipibm3i6o 8(536^3 oim30nb gig6m6nbcni6i33 bb6nb 6n6i 630b
3h3qjmh0 nh3i>x y3i3O036b (6ib. 5 .17 ). ib30in b^hs^&ni b30b 36ym6b 6iym-
qj3&nb ghmx 6 i 53 notih36 ib gi bi9 ihi 5 m 6n3cnn3h3?>36nb bBhicg gi5hm336ib
bi69 m>)03 3363iii(innb 33hnmg9 n.
- 301 -
{Wrtoanb <y>3 ^ 36 .> aiiae^fnou 3 rto^-&563 om6ri 3 to 83
Sbb. 5 .17 (536j)hacDhn ic3]6nb (2600-3200 3 bn3i(PC0t>3) S>iC™k 0^ hm'b3l) x n bibfujm^bpm

qjmh3nb 3u;)6ih33&3n y3i3npnb isnh^jinb ojnihSnhj&s pi yj^nonds (Moser & Zachuber cit. Larcher
(1980) 3nb3e3<ioi)
5 .3 . 2 3 D6 D ^ l 5 n n W 9 ^ iC fia ^ flb hflibanb bfl6 JhfT16 n(bnhD 6 ^
3D6D6X5nDhn bj^ri3m6nb phmSn 0n0gn6ih3mbi gi0ming3&xn:> ^ o'!) n9 3 ^5 n-

j)i^ 3 n b x b s y h x n S^hnmgnb bi65hdQn3m6ib3. ^hmSn^bbi g i bjb^hm^n^&Sn
bi3353î(5nm S^hnmgn nb£>X36i 03 hip S^hnmg^bSn 5yonb 3 bihgn g^qjn^n^nb
5i0m. 'bmQn^h bih^y^pbi g i 1503 ^QnSi^h bm63 &3 n 0U36ih336nb bi 30£,3 lî^nm
Sjhnmgnb bn6jhm6rfbnh3&i j)Cn0i^nb bjtmB^h U3ip3&igm&ibcni6 bmhyn^Qg^bi
Son^hn cgm^m- g i cnjhdmSjhnmgnbdnb bi93iC]6noi. bij)0img b3 nhig 5^6^01363-
pm&nb b30 nC3^:> 63330n6ibi i 9 i 013 n9 ihmu^bnb gi3cni3h36nbi 013 giBy^&nb
0i0mdhi33&3g dipig, hibi|j, ^1130133, bjpb ^Bymbb ^ O ^h i^ h n b USCnOD^^-
93-40 6 i 63 g“b 3 9 oi3Qn b i 3353îijnm ^hnmgnb 5 i 69 i 3pm6 i 3 n gp 33&n ("* 9 3 ^ 3
j)hd30ni. 50 °-gi 6 3b 3m3p36i ^0933 3qjhfn dpn3hg3&i (nb. 6ib. 1.43 ). 9(536ih3cni
bib3m&3&n, hm03pcni j^Bm^n^hn 1536^0 Siîpn gi bi33ipro 5i63g3&ncn ihnb
6ih9mg536nQn, b6m hx s0 3m3Q36ib ihni6 035336306^ i 3 m6 iyihnbi g i y 3 i3 /v
0m6nb 33hnmgnb 3bhn3 nbnbn, 303^3bBnQig, j>hd3pn gj^nb 0 my3i h x 3^ni- bmpm
336Diiibn3hn bhgnb hnÔx^Jxib 0nb3g3ncn - 0mxD gpnbi. |j6m6npni ^ (13 0 13 3 ,
5i6bb3i33&i 3iR lin33&b 3 mhnb: b3&ihj^>ni3pn 6 ih0 m3 n 6 nb 6id33b3n ^3h9n6i-
03hn ypmh^nb fiibib3i (gi bnj>hd33n bhgnb gi3cni3h3&i) n6y3&i 3i 3 n6 , hmg3-
bi(3 gpinb bi65hd0n3m6i g>igiiîh63i>b ^hn^njjx bc3ihb - 20 ben (Sî 33hm3-
30n 6 ih0 m3 m6 nb 6id33&3n 3b bp3ihn 14 b o b (impni) [4 9 ]; fihgngr^cn ig^hn-
*(i9 n, Liquidambar styraciflua-b fihgnpFQcnnb hn^3m3im^n336n cgm^m33hnmg>xo(?
3qjhm 3m x 3 'J>^3D£»3ii»i LSnrn 33hnmgb ihniB 9353363p6n, 3ngh3 bi3bh3cnnb [ 2 7 5 ].
0ih6y3nb iX iM ^ X 5 n3363 n 3X3iJiibf'3hn ( 36 iipnoi) j>i3 hi3P3&nb 5igihoi3i h33 -
hmg3ji3nx 't |3 (y3i3n03bncn) h353pnhg3&i gpnb bi65hdQn3m&nb (j30n03&nbi g i
^3033hiji3hnb 33333m6noi; bi0bh3Oi30n 3n93&n fihgnprôinb h32,nm63&9n 5i-
gihj^nb 333oib333i3n gngn b6nb 5 i 63 i 3pm6 i 9 n 0hi30g3&ni6 33&30>i bnDhi€? Q*
0bm0mg 0m53ni636ncn n3ncnih3&36 0 ^ 1 1 3 ^ 3 6 y3i3npo6b; Rhgnpm3cnnb 3 n9 3 &n
in, 3nhnjnoi, igh3 y3i3ncm&36 gi 336Di!îinDhig n33nicnig 3hi3cg3i>ni6 [ 2 2 9 ].
53gig 03533630n 0^36ih33<bn 53ni6 5i9mn(p36g536 x ,T|hli36b gi digni6 63Qi
5i63noiihg36mg636, hnb J>i3mi5 bi0oihnb 3nh33gn yn633&nbi5i6 gibni6g36mg-
- 302 -
86«?0b6 a66300)66060b b08063(f>0 908^06660066
636, >6 3nhnjnoi - dpn^h b6hiqgn 5>63rKnih3&nb &:>dm bj'bmSb

W^cyrngnQig 33^ 6^9^0936369636 (6ib. 5 .18). bjOQb6 &:>9m9gn6ih3, 335 3^ 0 -
nbi g> jond^yib hn(j 9 x ft1?)n^ bn6jhm6n'b:>unnb ihihb^bRbi ihnb 9u36ih3at> 553-
h(j3p36nb 3hcr>-3hoin 339 b(p39i 311 apjii^hn, hm03prnj 33mC3 jnnl3 3hfn(53b9 n, SQbbd-
03&Xn^, gnq)3h36unh3&nb 03933 n 6 ncn nymb 5>p:>p:>b3pn.
6ib. 5.18 3|j36ih3CTo thgnbs gi i^rdi^nb hiyi036ftb ynmhpnBiijni. 6ib>'bnb 6ihi}b36i 6i 6n<yi
ibibisb o ^ jh ^ ^ h n b s g i qjrfbnmeni&n^hn iiyiamBob jjj9nhb. b^hionb 8^ 336^ 6i 6ncn 36336360
Sjbiiidnbn ghmnb g3qjnumib (Burning, 1953). 8m6i(5383&n gpnb bi6j>hd(Yi3m6nbi p i |£)833hi63hnb
<'5D6i 30D®r*3 93bib3& RhgnQmjcnnb q)rocnmc9i5336 b3 81536111336^3 nb. Heide (1993), 9mgX36n
iC3hn g i tm8n3hn bih^ypnb b3 0|536ih3oiioi3nb nb. Hanninen (1990)
5.3.3 3l536ihnb i« n 3 n 6 n b i K> b3363&flb <3tt>3&nb 1530} 1503(1

âcnhnb a jn 6 3 h3&nm636an
b3 61336^3360 i0n6pnb b3<bm63h i53iC36igm6ib 35336^6 3hrn^m30i'b9nb

9gj,m0ih3m6 nb 33hnmpxn l53CnC3&nb, 03»j>6fn0n'b0nb yj^n3m6nb 93 U30 nb, &i6-
3nov>h36nb gi h3’bnbli36l>)3C‘rr*6nb (33ig36ipm6nb 03933m6non (6ib. 5 .19 ).
'bicgbxnb bmpmbo^nb qjnoip3bnb 936636011.6 R:>nbib3&:> &33hgnom ygpih^j-
6n. biSâ qjmcnp36n 5593001036*6 950693636, 9330* 9CfT»h(yi 5mh8m636nb 638-
33 m&ncn ^ g ig n b 9 mb3363&nb <3 i"bi9 n (yomh&jbnb ^ m h jc^n jh n d m b jj6 jd i),
- 303 -
(WfoOanb <V>3 c ?366 9 (3066601) 8 6 ^ 6-a 663001663 &68 a
063, hm3 b*8oih*3g3 8(5/1(13 b6ooi 330016303001 9 * 8 ^ 3 6 3 0 i*h5 *8n6pb nbo6n

0mb33636nb 33hnmgng*6 ih 5*3my*3b. b*8oihob 3mb3363dob 0g5m3*h3rn6*8o 5*-
g*en*6 0U35ihnb bb3* 6*600360(5 - 3*360380 9* bb3* Jb111300360, hm3 c 3611(5 3b-
h3633cymg36 3(j36*hnb bb33cnb 5*3 d03m6 *b yo63ob* 9b g ^ o g h ^ ^ n n b b*6n6:>-
*CêD6 m9 - 50631100(163606 h*nig36ni&s 303630^66, bmcn bhgob 030bc>39*3n
6*3(101360^ (0 *&., >&b(jo'bo6ob 83*3360) - 3i^3pm6b (6*b. 5 .20 ). *801(16331101>p
n<b(?393î 636360b *^03016*. (j)h*6b0*|5O3hn 3hm(53b36o 63(1936* g i 03hnb^30nb
33h3g36ob gsymogob 3 6 ^ 0 85330111*9 3(533*. 32j(i3936ob 6ohoi336o n8ymcg3&ni6
5*63oai*h36ob (jnjjQnb Gj qj*b*8o, 3.0. 8m3b*gg3&* g60 -ob 11393600^*1511*, h*b*u
*95090 3J63&* b*8oihob 8(^3363606 b^*goob 6mcf,ib. *b30i 3gj,n3*h3m6i3n 53-
6m8o 9*15300 069* nymb 9*6*00 ^383311*^3(13&nb 635*3036(^*5*6.
b*8oihob 80163363606 33hnmg3o 5*g*b30* 330608636* 33h3gob Snihmb^mSx
9* b363o])hrib>)ni33o 901633663: 36gm3o*b03ho 0306h*6nb* 9* 30*b8mg3b036ob
8 3 0 ^ ^ 0 1 3 3 6 * ( 3hm^m3o*b^ 36nb mboi0oh36*); 0 o^mjm6Qho36ob g * 3*^ *h * 33&*;
lbmj)n3hain 0(j36*hnb jooih oôib^b* g* ^0130036811 (5^*5301(136001 j3hjnb
jpmh36jn3*3n) ôo*ifTioQ36nb bob^30nb b^h3j^3hnb (1393:1150* 9* 3*530103606 9*-
6*633h36* (6*b. 5 .21 ). 33^* 6 monb8 ob 06^36^3016* 503630016b 9 * 36 bn 036 ob
838*95360016* 0(53036* (6*b. 5 .22 ). <b*3anhobo>3ob 3b*g3&nb y330* 3b 3hm(53bn
3o89o6*h3fn6b *h* 33(5(1*9, *(1*839 Oi*6 *3 o3g 33h30 *g, boi503hoi 81536^380 3qjhm
*9(13, b 01580 - 303(101 530*6; 9(536*hnb y330* 6*60080 050 3(1019(101300 *h *hnb.
3Jd3(i*Unjh n dm bjjSjdnb Zjhnmpn *hnb bhgob Shm^bnb ghm36oono 8363-
(136*, hm33on(5 o8*hoi3&* 369015363(10 33j*6nb336nb 83833016001. o9 ob 806393001,
013 hmfcmho* 3(536*hob bhgob <301(18*, 9^*530(13360*9 *(163300 01(15*601 *6 jboi-
Days
shorter
Draught
Temp.
drops
6*b. 5.19 8yiJ6ni5 ajja&ihjcni bjjrgm&nbi pi 9mb3363&nb qji'bj&'b] fph]9m qjiijmhj&nb V)9m-
j33K)&nb ySh&yxp &=>3A yi 3D&X n e"^h*a>
- 304 -
86^016 *56 a6630(J)66350b b08D63(f>0 903^06663056
Auxin
C yto kin ins
G ib bereliins
A bscisic acid
fob. 5.20 'bmdnyin
bihlyj(V>b b3bnpni3i6
3\j36>h33&3n qjo^mSmh-
3n636nb 6c<Thn pr>5i 9n-
V (Luckwill et al. iyvi-
Seeley (1990) 6* 3931101)
3D0n (383^3b 838016333580, ypmh^pbn 95 8 3 )^ ^ 3 8 5 ) 95 55301063330 ogyj^mho

^ 3 9 8 0 5 5 h35 305(505, 3mh8m630O 30101)19065(505, 55(13801 5560)1^335
9mb33636ob ^hnmgnb 3 h53500 bb35g5bb35 cgmh3 i 95 b^goi. ^nin^hn 0 b303Qm-
6nb 8380^333580 3b 3hoi(53bn 0B1J365 dmb3j6jbnb 6n6i Zjhnmgncn, hmSpob
phnbxj 9P336nb b565(^0030165 8(50119365 95 c»50ob ,53833)15^3(1360 (+ 5 °C 95
6550360) 2)03630m6b. b59350m 95 61190001301 5563936^3 bhjpn dm bjj6jbnb Ujhn-
mgn 000(^6335 6013863)^5 95 93i30&3h3o (65b. 5 .23 ). 50 ghoiob50 i3nb 016090
5dn6g 36ob 95950105b 5h 333d0O5 8(5365)106 553 oiy3565 3016336360b gj5<b0956 . h3*
'bnb(536l530m^fta 31,01001500b 5b33jjj3 o 8(5365)106 50 oi3ob365b 9090 060836300165
5j3b; 013 15030 <b50oihob 0 joi63 h3500163630. 835 <b50oi5h8 o o*6o0n 5dn6 gob 959-
5ni0ob ghmb 8(5365)13 5580135 80^3363606 0g5Oi05h3Oi6fi956, 0511 95^30569365
330950180 uo633&nb 8393559.
t>50cnhob dm bjj6jbnb SjhnmgagsS 558 mb30flb5oi3ob 633b b3 8(5365)136 bôh-
9365 65h])3330n 6565)1800301606 15030 33)100190. 383153b 838oib33358o 2 -7 *^C-ob
<35(1593680 j53833h5t53h5 3695 3oy)33036p3b 8(153500 330110b 5568530016580 (855.,
638b b^nh03&5 5 330(15893, 3580b5 95 8bb50b - 8 330(15893, 593650b - 10 530-
hi3g3 [210]). 80^3363606 Sjhnmgnb 601901b (8oib33636ob 838950180 331100190)
5^503oi6ob 55'bhgob 55801863330 050,301301)180163606 (33I1 G A 95 CK, boigoi 838-
935 IES) 3016(536,3)15(505 85l5x rT^,lj; 53636nb5 95 36*6083606 55^03(13606 838-
33016001 n'bhp365 6 o3om3h365ou> 15390b 06,3366030165; 8 oi6 o0 o‘bg3&5 b5h3<b3h30i
6n3cnn3h3636o 95 85^30016b giyroojob 365hn. h0i50ih(5 30 y3X^> 0oib58b59363*
0n 3hoii53bo 95bh309365, yooihôb 556300151-1365 95 b53h01019, <bhenl> Shoi^bo
83^0365 33n‘b(n3Qmb 8boi0mg 5h5b30b5yh30 O 55I13801 <35:1,501(13606, 556653301-
h36001 ,53833)15^3(106, 803b. 9^33606 b565hd0o3oi6nb 95 ,53833(15^311116 80185,53-
65 6nd5b 5d933b 5j^03hn "61^5-55630015)13606 3hoi(53b36ob g56y365b. b50350Oi 55-
63936^3 3 (5365h336nb bh30O 80^3363606 33hnrigo 01539365 'b53oihnb 636050161^
P5g6oi3ob5b, 58 801836^1956 8(5365hob 55630015)1365^3 001^839366 580690b 808-
en65h30i65 .
- 305 -
a ^ o a n b a 6 3 ^ 0 6 0 aiio e^n b
Dormant Active
(October) (March-April)
vftsiceJ emission
Storage of reserve Lysis of excess tonoplasi
Change in
nuclear prolems Excess of
plasmafemma
Partial loosening
of cell wall matnx
acrdic pectins
Less acidic Endocytotic Large central
wail pectins transport of sugar vacuoie
Lower activity of Membrane renewal
ion pumps and witb IAA earners
channels
H-lAA
Reduced amount of membrane-
located carriers and receptors
(no response to additions of IAA) Electrogenic pump carriers
and receptors activated
fob. 551 tn8nahn bih^ycnb 3yiJ5ni5 0^36^33630 a^jn^hn g* 9mb3j 636nb 9pam3ihom6:>9 n Symqj
iiSftn^Snb 33^3P3<^> 9mhnb jhb3& xn (jn^mtvn^n^hn 5^6bb3S336i (Lachaud, 1989)
1.0
>>
u
0
at
1 0.5
N
a
OJ
ai
>
^5
<u
QC
0
M onths
Bib. 5.22a,b 36bn9j6nb ©ujSihQoo s iy i3hn gjiV igiS 9mb3o536nb bvjpeoi3n jpgibspnb
ghmb. a I5nimjhm9 m jb n e ^ ^ 6^ 3mongj36mo mjbngi'bsb gjihgrninom iijnsm dj sC36onb lyvi^ftnb
iiii&njMn 839mg5>m9i'b31 'bbdoiihan 91 fci'biajbxfoo (Meyer, 1968); b 6sd3nb 6o63369n ijhrn^ o
9i'b36nb b]bri63hn 15300036360 (Esterbauer et al., 1978)
0 ^ 3 3 6 3 6 nb ^Qhnmgn g iS ib b b n ic n jb Q Q n b ^ 2 ^ 130 133 , 'bm^nQhoin 6 s o ib ri3 i6 n

9 l536ih n b so i 3nb, &:>6b:>j)30ih36ncn fr jm q jn & jb n b c n jn b , hm003&3 nu lb s 9 o ih n b 9 mb-
33636nb 33hnmgn 5hd3QQ36i 0h;>3bQn ^nhnb J,^50 i 3(vn6 i 3 n. a h ^ sc j^majn^b
a b i^ n h m jb s b n|jn3nb 6 ih i)3 3 3 0 n ^ h n m g n , hnb 8 3 8 9 3 5 5 9 ^ nbn6n 5 3 6 ^ 9 ^ 6 0m b-
- 306 -
86^016 ^6 &66300)66060b 10806360 909^06660066
Predor
mancy True dormancy Imposed dormancy
K /
Correlative Exogenous
July Aug. Sept. Oct. Nov. Dec. Jan. Feb. March

Months
fob. 553 'bmSn^hn bah^ypnb h3j>nm6]6nb 83hJ6fti5 0U35ih3369n 9mb3;)6;)&nb qji'bnb bj^Si^hn
S^^bsvjxoki (Saure, 1985). 9mb336;)&nb qjsVib iC^bnqjnjjiunnb (3ih>, 369m- 9* 3ynamb336o&nb
qp'bj&n) 93bib]6 nb. Lang et al. (1987)
3363&nb 0p5m0ih3m6npi6. 0hi3ip6cni6 6 inh6 ipib 369n pi 0ihij3Qm36360n 9 mb-

3363600 ^hnropb nB333b ihi 36pm{,363hip piShm&hiS^bxn Qn’b^'b^bn, ihi03p
dpr^ho bn^n3nb &i3C36i, hitf 363^30^9 j,i0mnbi^3di bhgob 93fi3h3&i9n (5^65/1-
cnihjdnb j^nm 6mdjhn djfijhjds). hm^mhu i,n j>ih30mb ^ 3 3 3 3 ( 1^ 3 ^ 0mn0i,!)3&b,
3b 0(336^3360 nbQ3 3^03(1036036.
5 .3.4 <3 D6 mcm&n:): a b ^ h f l b 6 ^ 6 3 naiihD 6 i , hPldFlhb ^ 0 f160 n b i

ic n a ^ n b i53c n c 36 fib n e e r u w i h n
0U36ihnb j>i63noiih3&nb q jit^ n b piByobi pi bi6j)hdpn3m6i p30mt|ng363Qn3

b3 n6pnb 0n0pn6ih3m&i'b3, hm03pni5 6pnpi6 6Qi0p3 n(j3P3&i. 3533 pnpn b6nb
5n6, 63636 ibai36 pijji33nh363pn igi9o36o :>i3nhp3&mp;> 95 09363693 3i536ih3cni
j(533nb nb3<ji q336mpmj)n3h 1530003636b, hm&mhnuii 93051*101033, y333O0 m6 i pi
<gmcnQ36nb j,3y3ooi036i. 503bnb 6331m B3b-fi33X36i Q^nphmp ihnb pij)i30nh3-
6300 i0n6pnb 0n0pn6ih3m6ibi pi 9i5363h33pm&3b 3 mhnb ihb36x n ii39nhnb
5mg6ibcni6. 3333 2000 630^3 03^0 b6nb 6n6icn R0630*3n ihb3&m&Qi 0336m-
(xnj^ho j)3036pihn; R3. 6.-nb 705 6pnpi6 ôm^mSo (oi3m6 oi) qjnjbohpo&mpi
ip36cnb iy3i33&nb phm; 1736 6pnpi6 n6j>0nb3n g>hm3p3&mpi q}36mpmg)n3hn 0 m-
6iU3036n [ 136 ], d33po q}36mpm2>03hn 9^3(^336360 b x 3<3hm 03^ 06083-
&30m6 ib 003636 cni6i039hm33 j)pr>d;>\!>,TiCm&nDh &i3 mi3P33363o.
imh3pi(5n3hig i b i b x n O936mpmg)oi ii>i33nh3&b j>i0my3636noin 6mlî6nj)nbi pi
03v53mhmpm5 nnb n6^)3h3b36nb bqj3hm9 n ihb36x biiooib3&b. 050 3yhg6m6i
0(336ih3Oii 5>i63noiih3&nb 150^000 ihb3&3p 93bi0R633 bibn^m^bcm [5300036360
pi nj,3033b 0icn b^ôbô^h ^ S n h b 30o3 i£)h qji;J^mh3&an36. hm^mhy Ri6b,
363010 ^ (130 ^ 0 36 0 ihb3&m6b 3bm0mg 'bmfcoohcn h3&nm63&3n.
- 307 -
(Wfoaanb 663*^066 90 3 6 6 6 0 b stfe ^ a ^ o o to fiia to so
hn^mhu ^■jbiQ jh-sdspn'bjhn 03(560311365, cg^mcmj/ii ou>3nbn j>563rKn5 h3&nb

bbBynb <35^5005. qg^BmQmfco^ho 3m30363&ob 6odj>ob 9n9(j30n qjbj^mh^ftnb 5 5 -
3 m3036 i 33*15(5 b i^ im g hoix*1 50ri(55655. bodfcob 0(10(5300 &33hn q jij^ h n ^hcrv
356301b 55gb<3ih53b: nbQain 06083630013560 n03x^<i, hmj>mho(5 ihnb
hi, Qmpngnunhgâi 0 hi350o 55*13 013 0055 qjij^mhnb 0 n3h, 0501 Smhnbii 6 n6 i
60nb 50o6gob 3n9gn6bh3m65 gi 9i5o6ihnb 5bo0o05(5oob n6^36bn3mbi (65b. 5 .2 4 ).
J F M A M J J A S O N D
64b. 5.24 Rhgn(yn bSocouS^stip^jainb h3j,nm59n Pinus halepensis-rfo bhpnb 3hm(j3b3&‘b3 yyidb^^hn
qji^mh^bnb 1336130061. T ,j393ohi(!oHi; Pr 6ic 3jn; Dr 531031 (Serre, 1976a,b)
5 . 3 .4 . 1 <3D6n°3i'b06n 9} TOPlOm&fDhn 3^6^

q)36mQmj,n3hn 550(11300636560 65(100^53636 55630oi5h3&ob {ph^33X ShmuQ-
b3&b (q jj6mo]5bjdn ): ypmh^Qdnb 50myh5, qjmoio^bnbi gb y353O03 &nb 55005,
<goioi036ob qyhnb 15305. g5j)3nh33636n b5{,hd6m&05g 5i05hlyi3g365, 013 50 0m3-
036360b 35h50XD^5g y3h5gc>36i 0n3ju335 bhgnb 35h503lj>h3&b1 565^m0n^h (15*13-
gnb gnqj3h36unh]65), 306hnm{,363^033*1, Snb^jnOn^h (055., b505h55m 6030103-
h363&ob g5j)hm336nb 8 o8go65*i3Pii>5) g5 60010(11503*1 (36bo03&ob 5^030165) j,hn-
(!)3ho3336b (65b. 5 .2 5 ) .
015365*13015 oi565b5lbm55gm3636nb J,56300i5h36ob ho^3 o 03od0365 556^01550-
93b g5 (<33601001503*10 ib^j^nb J,5oi350ob6o636noi) 95353039 3b qjjGmQm^njhn
- 308 -
8 fn m a ^ a^ 63 nm^(na60 b uosnsDtnO 809 ^ 06 atf 0 m.b
Leaf exDansion L . ■....■

■
Shoot elongation ::: r.•■ .e
Root qrowth cru. rL.y.-.i’vsi.a
Stem diam eter ^
Storage ........rMB.-SM
Cambial activity* Phloem Bn ^-J ft*
Xylem
i i £ i— i— i i— i— i— r—t— r
1.5
1/ Seed
i-l-K *
E.
c / / - I ..........
t5>
i
- 1 0.5
/ ' Endosperm y Embryo
-i \y ''
—o-* — _________
--- 1------1------1"---- 1----- 1------1------1------1-----
0 10 20 30 40 50 60 70 80 90
Days after opening of flow er buds
6ab. 5 5 5 9(j36jh3oni tihpirj^^nauihQ&nb ShmuobSn fiihoi3pn 3oJi6nb9o6nb ghm3n {p6:>6nc3b:>.

bogi 6*b*%n: 'bmSn^hn bih^ypnb b]j63n bhgnbi pi pi5hm33&nb i^hnmgjbn (Wardlaw, 1990); J33&1
Bbbi'bn; ih iy iix 9iC 'i o3cnr>'J 6^cn3i 9u36ih33n (Saxifraga oppositifolia) otjbcnb 5^63001^361:
A bnj,m^nb b^sgni; B iphjjpn 336hnm6xi63noiih3&nb pi 36pmb33h8nb b6hiqjn bhpnb b^>-
gns; C Q3&636nb qjmhSnhQii, 36pmb33h3 nb bi6mcmme fi:>3 m<£>pn&3&s; D 386hnm63cn ypmhiij&nb
fii9mip>Cn636i (Wagner & Tengg, 1993)
M jjijhjbnb (6ib. 5 .26 ) 03033mbncn. gngn 3 ib 9 li)i 6 nb 3m3C3636n, hmfcmhnuii

q3Roi036nb j,:>63nau>h3&b, o^bgnb 3tTi36ncg3&i, b6 q3maig3&nb qj3hnb 33b3Ci ,
03ndp36i 5i6nbi‘b(j'3hmb ^hmcgm^mjpgitjvj&nb b^33bQ36noi (qjm^rr^nb qyhnb
9nb3g3ncn). i 3 nb 93g3&:>g, 3hoi6inhn cg36mQm5n3hn 3m30363&nb bn3h(53gn 3ihi-
03(jh3{>n 33 ndo3bs giî6nQ nJ63b (6ib. 5 .27 ), >6 5 i 3 mnbibmb bnSijngnb
ShmqjnQnb (65b. 5 .28 ) bibnon.
q)36fnq)i<b3&nb 9n0gn6ih3mbi lbri0n3h bjly>ifTp9 n. Bgnb 3nh33c 6ib33ih3n <33-
6mq3b'b36nb gi6y36i b9 nhig gi3ming3630ni b33unqjnj)3hn ‘bp 3h x n t^ ^ h i^ jh n b
J)igi0ib3i‘b3. i 9 nb gigjflBs 33ndQ36i 3i 6 nj)S^3 n ( i^ ^ h i^ jl ^ n b g>i6:>6nQ3&nb:>
gi q335mQR5 n3hn 3^gnb 33gsh36nb bi93 iC 3&nai. i3nh^36nbi g i qjmoiQ3&nb 5^6-
C\ b3 3u36bh336 nbi g i &3fij3i>nb j^^nem&nb gi6y3&i g i o^bgnb 3 n6 i‘b3 fi>
3nnj336i 33bidc3&3oni 9 bmQmg 3 s9 n6 , hmg3bii5 S^hnb gi 6nigig,nb ib3933h^3-
h3&n bibmgmmg {>igiQibi336 <bc’3h3p 36n9363em63&b. <bmj)ig ig I i3nh£)&nb ftibb-
- 309 -
cW6 3 8 C% a^3^ 36^ 90366601 8 6 c?6-&6630a)6635683
Meadow Forest
Beflis peremts Carpinus betulus
Primula elatior
Taraxacum officinale ... s e n . ......

Quercus robur
Anemone nemorosa
jf||
a I
Ranunculus acris Ranunculus ftcana

Western Europe
Arrhenslhenjm eJal. ,J .£ k Ranunculus ficaria

England
Chrysanthemum leuc. U hk. 1 Ranunodusficaria

Ukraine
Knautia arvensis ... s / * / . _ Pulmonaria offlc.
:t T :T :i t - rn r
F M A^M J ' J A'S 'O N
Cold regions Dry regions
Pulsatilla violaeea
Caucasus 2000m
Saxifraga oppos.
Alps 3200m
Saxitraga oppos.
Arctic Canada
Minuartia sedoides
Alps 3200m
Lamium tomentosum
Caucasus 3200m
Potenillla pamlroal.
Pamir 3800m
m i I |2 0 3 H 4 E 3s
6ib . SS6 qj^Bmofrijin^hn gni&hiSobn: 9p3(vnb 6(j36ih3:)bnb (3mem63oin); 9 ]b 6 ihn ,jynb qs

6inh6i£>ib36nb (fihg. j y i 9i 6n i, n6j£nbn, h^bjowi) 9 i nbjcnn hj^nmSjinb, biQi(5 bi33£>!>iunm 3 -jhn-
mpn 93'b(Tr)e3c n i bn(5n3n<n ( 9ip i{x k n i, ih ^ y iii), i 6 63iC3noi (bv503n, 3pib6m) (Salisbury, 1916:
Schennikow, 1932; Ellenberg. 1939; Shteshenko, 1969; Jankowska, (jn^. Lieth (1970) mixedviT;
Ackerman & Bamberg, Lieth (1974) 9nbog3nai; Shalyt, lyij. Beideman (1974) 906393001;
Michelson & Togyzaev, |yi^. Voznesenskii (1977) Snbjejncrt; Moser et al., 1977; Svodova. 1977;
Nakhutsrishvili & Gamtsemlidze, 1984; Borissova, ury. Walter & Breckle (1986) 9nb3e3noi)
6nbi pi y3i3n0m6nb Qi6yo6nb b p 3h xn ^ © ^ h i ^ h i 6 -10 °C-nb qjihj>Q3d0ni;

iph39fny3b3n03 g i 0ipi03cnnb 0^36^336911 nj>n 3qjhm gi6i0fii - 0-6 °C; 530^6-
0 my3i 3(i03 0u36ih3369n in 3qjhn 0ipi0n - 10- 15°C. x ' 3n^ t>3, ihyn g i bmftn-
3honn 6n63R3i6n i 0 ib ^(*11115(1303636 0°C-b3 R g6i3 0ipi0 (j3033h^3hi'b3 [ 2 1 3 ].
yQmhj(36nb iQmyhi g i y3i3(i0m6i 93ndp3&i ginBymb b3Qbiyh3Q <i)3033hi^)3hib3
0bm0mg 0 i 9 n6 , hmg3bi(5 0mbi0big3630(i q jib i j>i30n0ni, 3.0. ©1536^3 3 33 3 £>i-
0m(p3(id3630ni (&i0mb30ni "bi0onhnb 0mb33636nb 0gj,m0ih3m6ngi6).
6pnb d jm h j 6s b j 3*hdn ih b 3 & x n qjoemgm^n^hn 3i g 36n, hmfcmhnisii 6 iy -
mqj36nb 0m06nqj36i, qjmcn036nb 5iy3ncnQ36i, qjmcnm0(3336i gi 0mbi30nb ip360b
ghm, 33nh3303b ym30dbi, gi 3ming3&30n ihr»i6 n0 5ih30m cgij^ml^&b], hm6-
0360(3 i6xD^36, i 6 b(p3gi336 0m06no]36nbi gi gi&3h36nb 3hm|53b3&b. i 0 936-
- 310 -
- 311 -
». 5.27a,b 33hm3nb 91 fihp. i3ohnjnb 6i 6ncnb <336mcm6n]hn hQijdn (Ihne, 1905; Caprio et al., 1974; Schnelle, 1986)
{W tfo a n u & i3^066 9036 ,!>rt0b 8(n«?6- a 66300 ) 6 6 a 668 a
Days of the year
6*b. 5.28 s3b^hnnb iC^D^Sn bn9ic>on<n {)i 9m633x n ien3i^nb 5130361 qjo&Ticmarrjh 90161(33936^3
(nb. 151130133 61b. 1.37 ) (lyiqjhj&ncn ic 6 n96x n * bb3igibb3i 9u36ih33&n). <3360100150311 0m6i-
06083630^13160 bb3im 6i bn9ic>cnb lbhgibaii6 ohoug 3njonnoi36b 3itfb3g, hm3 ,5333^ ^ 3-
hi i3 933oib333i3n oii9i3ri6b 5i69bi'bc'3h30 hmob. 0i9n6, hmiji 306n3363cm 0336010^411^(1
bb3im636nbib £>0130616(10 3^0100^3110 qji^ynhm qjoiôOohnoign (Roller, 1963)
016333*8015 ^ S ^ h i ^ h i b gngn 86083630016* i^ b , 330(133036*9, (101501(115 603010-

3(136*01* 3hRg3Junnb b303386ym6b. >0 ghmb 5*9*86y33,5n 06083630016* *j3b *h*
‘b(T3h30 iSDSôhi^h^bb, *11*839 (53333(1*^3(1*01* 3*0b, 3.0. ^^Si^h^^hnb 06(535-
h*ob ghmob 8016*5330180, hm9 pnb ghoib*(5 *po6o3636oig* b30b*yh3QO ^3833(1*^3-
(1360. 5*g*96y33L50 86083630016* ifob nb3cn cg*J^mh36b*u, (101501(1015**: 6*53360
6030103(136360^ g* Bypob 8*11*50, qjm^r^hnmgnb 5*3036* y3*3nom6*b* g* qjoi-
01^10(5336^3 g* lb*0cnhnb 3mb33636nb 33homgob g*6y36*. bb3*g*bb3* b*b3oi6*3o
cgmoiQob 5^3001036* g* 1533®* 3b*gg3&* ghmnb b*65hd0O3m6nb 338150(136001
(0*5., *hyn, *03nb b3, (50(10030, 6nqj30n, 836*, 6353hfib*0n). gj36morTi5n3hn 5*036-
ghnb 3b 601001 <3*^360 oi*3b 063636 3 *9 n6 , (101501(115 ^n ^>(313(1300 ^3833(1*^3(13-
6n 0*31538* 5-10°C-3g3.
q j3 6 r i( Y r'd r,3 h n 3 h r n 5 3 b 3 6 n ^ h n t in ^ & t a g* q ^ m o m j^ h n 3h oi-
I53b36n *3 (13500163680 330603636* n3 *95003680, bsg*u *hb36ri6b 3jhnm gjcn
6^cjjjdn g* 5 j* c j* . 63o3nb 33hnmgn *hnb 3363i5*l5nob ghmnb dohooi*go 9 m6*y
33010. 53*03nb*b 3*hi530i7i3636o g* bb3* 6*c*bm3*6o 91536*113360 ^3636*; <301-
oimo3l5336 6336b 15303* 0301010360; 3*h*g363*63 63360 53*0306 ^homgob 30(1330
6*b33*h3o 5*115*336 6363(10 qjmai036nb 3 33 l53brn6*b.
63080*6 (jhm3n53o (13500163630 <33601001503!™ ^ 3 3 ^ 3 6 0 830603636* 801300
Bpnb 5*63 * 3oni6 *3 n, 013815* 3b * ^ 3 ^ 3 6 0 *h *hob nb3 oi3*o9ob*u33n, (1015011115
853301(10 b3<bm63ho 5008*1506 8jm63 h35nm6363n. 50o9 *l53ho 03*^01(1360 0153036*
oi3noi 353*1501(130 b 016*80(5 (6*b. 5 .29 ). ,5(101305360b '601503(101 h35 nm63 n 033-
6mom503hn qj*'b36nb 15300036* 6*5036*g 33b*3R633o*, boi0m ^01503(10130 -
0533oih*g 5 *8 mb*l5X n- 6*03^36001 8gog*h 353*^01(130 lbm6 *3 n 63*03ob 33hnm-
g3&b 33ndo36* *g6O0O 3jm6g3b b*53*mg (13530*1130*9. 3o3b3g*3*g o3 nbs, hoi9
*b30i h35om6368o oi3nb b*83*001 (53033h*(53h* b*53*mg b^*6ooDhn*, 3 obo 9^3-
("*83(10 83hy3m6* ‘601533(1 86083630013*60*. 8u36*h30i* 5*63001*1136*^3 8oij83-
- 312 -
8( ^ 0b5 & i 6300)6rt050b b08C%D<f>0 809^06560066
9 36 b Bpnb 5b 69b 30m 6b8 o gpnb q jh o b g 3 8 6 0 8 3 6 3 0 0 1 1530^ 00^15 i o ,

3n 6b n gb 6 b9 h 3 5 0 0 1 6 3 6 0 b 0 15 3 6 ^ 3 3 6 0 c g o i^ o i3 3 h o m g o b :> g 9 o b b ^ 8 b o ig3j>hd 6m 6nb-
h36n bhob6.
^3&ni6n ^hoi3oi 36ob 8bhbg863b63 ^ 33 6 8 0 cg36oi0oi5o3ho 3hoi(53b36 ob 9^953-
65 03bbdo3&3Cn^> 8boi0mg b^^nb^nijhsp, 3.0. hoig3bb(5 nbn6n 93086633^ bbb3-
m&3&nb:> g i o6go3og3&ob gng hboig36m6b8o. 5b63oaibh36ob 3hm(53b3&nb 3308061-
I5dbb ,j)hm3n^3o t>3 8(536bh3368o, hm^mh^ 63bn, bg&O0O yj3b jj,jn 6nj(jnjdnb
ghmb, 3-fi. 630061980 mh^h (6bb. 5.30). <3010101015336* ^hmSoô ^ 3 3 6 8 0 0]n-
6119636b 3oi3pn 6onb 5b68b30m6b8o gb d(yi3hp36b 6nbgb5nb 5b9m9hm6nbbb p i
gpnb bb6j>hdon3m6nb 338(5oh36obbb. y3i3o0oibob 33hnmpn'b9nb 9nb3g3noi 91536b-
h33&n 03fido36b gbnymb dih^pdmy^^jnpj (9b5„ Hibiscus, Heliconia, Cocos, Carica
papaya), Ggnb bbj^g^bbji ghmb dmyj^jnpj (3b6 > Cassia fistula, Spathodea,
Lagerstroemia), jJnbrvpjhip dmyj^jnpj (9bg,., Dendrobium-b 6163016360) 9b
dhs3sp6pn±6 nb^jhjspjbdn dmyjijnpj (0b5„ 6b863ijo) bbb30i636bg. ^hm3n53Qn
(jy336nboi3nb gb8bbbboboi3630ob y3i3o0nn6ob dohocnbgo 33(1001906 bhbhb36m6b.
bhb36m6b 636nb9n3h ghmb 9my3b3no3 b336n, bmom 3hoin bbb3oi6ob 013 n6gn-
3ngnb (gb ‘601533(1 3hcnn n6gn3fignb bb3bpbbb3b ^ m ^ o b ) <^(1503680 y3b30poi-
6b bh 9n9gn6bh3m6b oib6b6hbg. b3'bm63ho y3b3nQm6b oib3gb3nh330bg b(3'n6n06Q6b
n9 (13500163680, bbgbtf 63*C3bb ^363Cbh3Cn 6b6oboio y)3b. bb30i (13500163690
"601503(101 b3 8(5361(136 5bbfi6ob y3i30Q0i6n6 3nghm33hnmg3on n6g3j(5oi [4 ],
vjhrn3nj)X iSyOD^n 860033 6bymcg36b 8308036* 833b3g30i 801300 600b 5 b69b3-
Of'A^n (6bb. 5.31).
Climatic factors
Day length 12 ± 0.5 h

Radiation
Air temperature
Daily temperature fluctuation
Precipitation
Phenophases
Bud break
Flowering
Fruiting
Leaf abscission (deciduous)
Leal change (evergreen)
J F M A M J J A S O N D
64b. 559 516180 (6®N) 81(119963163 ljhm3nixn (W** *0 815361(13011 b]<brn6]hn 51630011113606 3hm-
(536360 91 >,cndi(53hn (giijmhsbnb 6003(10 (5300006360 (Longman & Jenik, 1987). 33(5ihn 630836061
pi 8mh6y3ob 513036b 89hico ljhpi3ni(3cn iSynb ^3 8(5361(13011 isohao&'bj nb. Borchert (1994)
- 313 -
&6 6 3 9 (% &63=?366 90 3 6 6 6 0 b 8 6 ‘ 5A-a66300)6605683
fob. 5.30 ynb^p-hn>>;>3 n (lO ^ N ) jiis m b bjSn >>>86fu>0Dhn 'bhpnbs e» y jig n jy n b n b gi0m yip3& xoi»
Sijhnb gs 6iODjOi ‘t'3 (Alvin, 1964). &)6ffe6n ^hm3njxn i4a°b bo 9u36ih 30u j,s3-
bnseohn 'bhgnb goSehmSo^hncn 3 m6ri|jnnhn65nb 3rn63(Jo9o&n nb. Bethel (1990)
(hb. 5 3 1 3i6i9nb bjdftihngx S.

<3mcnom3>S ^y]3n bb3ipbbb3i
bibn^mybcf1' qjmhOnb
duofchocre 6iymqj3bb3mnshm-
6nb bobmb^hn
Oii3nbo&oho&i6n (Howe &
Westley, 1986). bb3i 3m5iu3-
8360 nb. Jordano (1992)
- 314 -
86m a ^ &d6300)660&0L 1)08067)60 808^ 060630^
5 .3 .4 .2 e o 6 e h n d h n 6 n c n 6 n i: 6cnohn h b n co b n u qj 3 6 n c n 6 n^ e:> i>en-

a ^ flb l53CflCD6 i
6903(10 h&mojbnb i6ioob3ho 9369(1^133^(10* (6cnjhn h^mQjbnb cgj&mdjtfw»)

6^8111^95360 qj36moni5nnb (1*111936016(103 a^cnrripb, hm3 x nl5 ilXinCD&DO0* 8h*3*00
b*30cnbnb 5ig*bi^hx*e- 3*060*03(10 i j (jri3rn6nb bs65hd0O3n6* g* 90033(136150*150-
nb 1133000, 3.0. 03hj6ob 836360b ighoxn s6 530*60 Shm^bo 9^3^130936300^
5ih33n qjyj^mho&bo. bm5*g*g, *0(13390 33hj6nb 836361b b x b 36ym6b ob 51(13-
9m q3^ m h 36n , hm0p36o(5 b30biyh30n*, *5(130133, bofthd^Sn b h g o b io ^ n b g* *0m-
6*y*hob 5*63oai*h3&nb*cn3ob. *0 O(jm0 , *0(13390 03 hj*6 o *b*b*3b 606* 690b 333-
6nb (133000b oi*3nb3d3h3636b g* sg(i3 5*b*qjb30b3 *hb3& xn bhgob 3ohm636b.
y330iq j3ho, h iu 16 3 0 3 6 b 8u36*hob bb 330ob b h g ib g i b 3gb 36ym 6b qgmcnonb g >
i^h^nb 3hm|53bnb 5*3+10*336*^ *ob*b3&* 530160 03hj6ob 30333601360b 90033-
^6(50(136i 3 o (61b. 5 .32 ). 6903(10 h^mgnb ^ 5 1 6 3 , 530160 33hj6nb 33(139J ^ b 339-
0360b bobfo g i b o3ôghm 33 *b sb *3b ibn0O0i^36001 3rri0ihi536ob gm 63b. 5*6*-
013639016*, ,$3033(11^3(11, bi333&o 6n3om3h3636ocn 3<bh363x y fT1<3 i. Bygnoi 3 m0>
(115361 9 1 y330s 83b *d0m *h*b30b*yh30n 3ohm6 * (3*h *b n li)36 ob 8 3 8 0 ^ 3 3 * , I5bm33-
0360b 803(1 01536*11301* 6i6n03&nb 83^8*, 53*03*, 6o*g*5ob 91,533361, bo^b3 9s
yo63*) 1693636 3nhgi3oh *6 ih*3oh0*3nh 5139361b 60o3ho h5m0ob ^ 5 * 6 3 ^ 3 9*
3ob R*3my*0ob36*'b3 (61b. 5 .33 ). hmgobitf 5 1 8 ^ 13 3 0 3 3 0 3j536*hnb bhgob 8 o3-
pn6*h3n6i g* 3*36 oi03 hn ijj,03m&0b oii3ob36;)h 36*60 V jb îg *hob I56m6o0o,
*03010 06903090b 60o3ho h5m03&n 6sh8m*g53636 606* 693630 bhgob 8*8md-
(11336300 d*036ob 153009360b 3(10153*11 ihjogb: nbn6o 5*3mg536o*6 *5(130133 i 3
693680 *hb3&xh]9ri ojbiynha-
bnb is^bso^hn
3ho«i3hoim6s6n A narrow ring
(Fritts, 1976) is formed
- 315 -
aoosafnob 8 rf>g;a-ga630 Qi;>fo3 ii>a83
Gradual decrease in annual rings with ageing
- 316 -
8 6 ^ 6 C!6 &66300)66360b b08n6Dff>0 808^06660066
6sb. 534 tpbtpgnb dorj&an jynb b jp i bi'bpgsh'bo 3m'bihg Tsuga mertensianaA) 6cnohn h&meo&nb
9ob6i3(xib biojodsx'bO biSîOfn 6pn;)hn dîjhs^^ho&nb h;}yn6b^h;j)uni (Graumlich & Brubaker.
198 6 )
- 317 -
U&foObO: ^ 6 b O S ^ m t)
6. ai536ihDD6f1 biihObX
aQ an a^h on bM n
93g*0n6nb y3X^> 33016380 8(536*113360 5*60(590*6 8(1*3*90 b*b3*g*bb3* *h*b-

30b*yh30n qjij^mhob 5*39 36 ^, h*y 3 *oin *h3*pnb 8 3 ^ 3 9 3 * 6 06333b. *hb3&m6b
nb3cnn 50(06*03(10 h35nm636n ((110510(10(5** 53*C3ob 13(06360, 9*80*83630 6o*g*-
50*60 *9500360, 3mQ*h3QO g* 9*(ps03cnob <bm636o), b*g*(5 I5bm3h36ob*oi3nb b x
b*yh30n 30(1016360 3 bmpfog 6unh3 ghmnb 5*66*39(06*80*. b3303 oiob 3hcn 83*-
0139^3 *g*3 n*6 nb ^833(163(0 *jij03m6* nb3 6 *ho3 *hcn*, hm3 3393(10 8(536^3360
5*0)333630 nj636, *6 b*3hoing 5*6 *9 5 3)^ 36 . o3 *95(103680(5 30, b*g*u 5*h33m
qj*j^mh36n 33g3o3*g b3pb*yh3po*, 8(536*(i3Xm6ob dpo3ho 5*63oai*h36ob 5*3m,
8(536^301* 3hoio 6*6non 5*6o(5gnb *h*b3ob*yh3po 3ohm636 ob 5*3C36*b: b33&nb
3 ôghm 3*h3ob ^338 5*6*013&3on6nb dgo3ho 93050(50^(1 *h*b30b*yti30O* j33^yn-
b*cn3nb; 310633(136(5(1* 03b33&nb 60*9*580 5*3h(53036nb*oi3ob b t p x ^ b *3 3bho3
b3bl|) b*b3fn636bi g* 069030936b; 6om8*bnb 8*p*0o 3016(536^*150* b3pb 30ym6b
3*h*'bnlj36ob* g* qjol3(oq)*536ob 5*311*3936*6 g* bb3-
y330* ob qpj^mho, h(0830(1(5 3y36*hob*ai3ob *h*b30b*yh3po* g* 5 * 6 ( 0 6 3 3 x 0 *
îd n ô b cn*3ob363h3636ob *6 ghmob 6 * h i 3 3 X 33(1001980 * (^ 3 6 3 0 0 30(110636001
(*3*3 3 phmb, *h *hob *3(50936300, hm3 050 3(536sh3b bibnyfn^bom b*3o3hro36*b
3J36og3b), bm5*9 *9 , 06(0936* "6(3(13639 q3*j^mh*g".
6 .1 UbhDbfl: Q^hGP333} bn69hnafl
6 .1 .1 h i b 6 f l0 6 ^ 3 b U b h O b fl
b ^ b o b 93qjo6oi5n36nb 333^36(06* *3 3m3036*b *(763(16 (1015(0(1(5 b*b0|5(0(5b-

On rnSô33309*6 5*g*bh*b, h(Ti330O(5, 330(13x36 yri30ob*, 06333b 33 :1(533*0 1530°"
03636b n69030gnb mh5*6o<b*unob y330* 9(^163^3, 3*5h*3 833390* 0(53010 1539093-
63611b 5*8016333*15, hm3036o(5 01(15*60^880 33*06 ^ 0133636. o3 838016333*80(5 30,
hm93b*i5 6^(136390 8013036* ghm36noio*, 8obn 3*hymgocno 5*3p36ob b*65hdpo-
30160b 3s,336*boi*6 3(101*9 3(536*hob b*bni50i(5bcfo *^03016* 3036x016b (6*b.
6 1 )• 3(536*hob *9*3,3*15006 5*11333300 b*t>(?3h36ob 5*g*0*b3ob 338935, 5*113563-
0*9 83386633300 (0*636^3(10) 63083880x0 qjiJ^mho 06333b jhm6 o3X 9 ** 3*93~
6*b, *6 833^533*9 l5X n0D^D^b.
60,^306 3ohg*3oho 860836x016001 “b^h^bo" 6o86*3b 83'b p x X 30(1106366
(0*cno63ho: Stringere), *6 5*b*ôhb, 36x3^13&sb (o650ob3hn: Distress). <30^03*80
3b bn^yp o65pnb3h 36*^3 6o86*3b bb3xob o3 806*5*6 g*d*6xoi6*b, hro3x
o(5 6*h3moj3636* 8*6^3 bb3* bb33(Yib 8bho9*6 d*pob 3ny3636ob 838016333*30.
*3 3nj33936ob 8n3h 5*801633x0 bb3X<ib 5*6303*9016* (bo5hd336ob cg*h90160010
bb3*m6s) *(510608636* 11015/0(1(5 Strin. g*d*630m6*b* g* 5*6393*9016*6 8mhnb
- 319 -
9 006 6630 60 ISfoObTJC? 9 CJ& 0 9 6 6 0 0 6 6 3 0
Normal Stress Recovwy Complete Normal Stress Reoovery Incomplete

slate stale
fob. 6.1 a,b jjihjQmb b^hobx 11 g siyn hjinb 6^303^ d(jo6shnb 3hm{riJly>;X'<T>&:>'b3. a foKJmjca
bn; b 3]h 9i 6o6 lj x n b^hQbn.
93qjihg3&b (].n. Qpib^n^hmbnb 9 ng 30n) g i 5iQib30i jgib^n^hngiB ( 9 ojU3-

3igngi6) ôsbijjn^h (033jli33ig) goqjmhSnhj&igm&iSn ihnb Oi^hnnb 0130636^.
bij)9 img b9 nhig j)i9mcnj83&nb 6y3ngn - Stress-Strain i 9 5^536001 j)i9mny3636i
6 nrri0 n 5 ni9 ni3 [2 7 2 ].
0ib 030g3&. hi(j ^>3h9n6n "b ^ h n " nb9ih3&i y330i 9m3p36nb 9 n9 ihoi, hm930 -
nu mh5i6n'b3nb gi^3nhai3ib 06333b, gn‘bn])ngi5 iCO&DCn a^ jn n6D_
Qnbb036i, 3.0. 9^3036;. g i 9 nb 603(1 9u36ih39n j>i9 mB33x n 9gj,m0ih3m6i "b^h^-
bnb" bibncn nj6i 9 np*3&30 n bigiSihiyn 36i9n. &i363&hm6nb oii3ngi6 ibi^nQ^-
JOig bi^nhmi li)3h9n6nb ihbnb 'bjbîg &i69 ih,j3&i: b ^ l^ b }^ qgij^mhn (i6 bjjh^-
jpihn) ihnb b^hjbnb &i9m06333n 9fTi3C36i, bmgm b(jh3b x n h3ijijni, g i 33b i-
bidnbig, b ^Q b xn 9gi>m9ih3m6i, ihnb h3ijyni g i 936336igfn6i (ig i^ itfn i), hm-
930015 5i 9 mnB333i bi^bmhnb 9 mj93 g 3&nb 03 g 35 ig.
&m0n icn6c330369n ftnrnQmyiiSn b '^ b n b 0 3b 6 i30 i b x Dgjhn gng 96 0 836 3-
0m6ib nd36b. b^h3b30 n 9 m303636nb 03b6i30nbib 5 i 9 mny363bi 53033nb bi^Qimg
9hi3i0O33hiT»3i6n frbjbn g i bi93i03&36n. j339mcn ip>363hon hi9g36n93 9ioi2>i6b.
5 i 9(pn<bni6360 nb 03b6i30nbij,36 0n0ihoix*i 53033nb 93cnmgn [ 140 ] rnhn36,yv-
h3&30ni 6ipn‘bni636nb {>i0m06333bi g i 9 nbn 9mj03g3&nb m &n^b 9 mhr»b ih b >
630 ])i30nh3&'b3. b^h3bn j,i6nbno3&i hm&mluj 0n0ihoi30n, ^ 6 ^ 3 ^ 3 0 0 qjiJ^mh3-
6nb 0n3h &i3m63330n 9m3036i. 6n6i 30i6'b3 6i0nB330ni bljh3b30n qjij^mhnb
9mj93936nb 03ji6n<b9n. bi 6 i 9 bjjh3b30n cgij^mhn 03bi&i0nbn lîjynnb ig5«0b
- 3hri|jm30i'b9ib 3ni(j'633b, 0nbn i330nli)3Qi bij>hd6m60ig 0133036;., 3n6ingi6
0U36ih336b j>iifi6nicn 0hi3i0n b3q}3h x n 03Ji6nb0n, hm903&n(j 5ih30mbi gi
D3^3gb 9 mhnb Oi3h0 mgn6 i 9 nj)30n i6 jn0n3hn 6m6ib6mhm6nb giOyshQbib i63-
03636, i6 i0unh3&36 i0 i 013 n9 qjij^mhnb &hign36l>>b. ai3ncn 3hm^m30i'b9i9ni3
2>n ihb36m636 93ji6nb03&n, hm903&nb 5i0i[pn'bni63630 qjyj^mhb i93bh3^3&36,
i6 b30iu oii3ngi6 0^03636. cnb3giU3nb &3qj3h30n bi03i03&i6n ("Avoidance",
n e ^ c n b jh ig : o u > 3 n g i6 i( jn o 3 & i) 9 3n d0 Q 6i 033 ig ih m c n 3 h r r i^ F i3 0 ib 0 n b B n 6 i-
ip0g35m&nb 36ihb ("Tolerance", n6&0nb3hig: ^0311*6^30016* 9 ij,., nb. 6ib.
- 320 -
b&fssouo: ^ 0 6 ^ 3 0 3 0 do b06 Â nan
6.73). b^jbob 3mj33g36nb* g* h3bn6lJ36li X rT1kn6 3m8*i!)36n6 336*6*8060 3hm-

^ 3 6 0 6 îpn^njjjhn 5*66003* 6*93*036*6 od033* 5*0m3*ht,3omcn qjsj^mho-
601306 b^nqjn^Qhn 0mj0x3&nlJ 3*^3360, (10100366*15 gngn 86033630016* *^36
3mc3i 303hn 6nmpm{,nnb, jimqjn'bnmcmftnnba g* 5*3my3636nain 3hm6038*vyiin-
b*oi3fll>.
dpdmdihjmbnb d jb 6^ c n b ^ j)j 6 0n0*hcnxn i30 33o b 03oimgn [219 ] b ^ h o b x
bn^^nib 5*60600*36 ho^mh^ 906*80311 Shm^bb, hm3 x n i 5 80130 mh5 *6 o'b36
0mn|ji3b - *3 ghmb, 6^(13601(106 3n3h 5*801633300 h3ijynnb 5*hg* 03n66*3036*
bb3i, bvi)h3bnbcn3nb *(1*633130030^0 6*yi 01636013. 6(5(13606 0n3h 5*901633300
h^yfonnb 01*306363(136* 5*80106*^36* 9u36ih39n *(1636x^1 9gj>m9ah3m6nb 83b*-
6*3nb*g 93U30b3 n ( b ^ h j b j p n b n B g h m d n ), 9*5h*9 3b 9g5m9*h3m6* 36361 *5-
(130133, 31101(536366, hn0036nu □'bh^Bg^Qyrnoj^B mhîBn'bSnb 6*3(10101 9g5h*gm6*6.
3b 3hrrii33b36n 9mj33g32)36 50183366*^01(130*0 - nbn6n 0633336 6*601301(36001 <336-
fonôb 6mh3*on'b*i5n*6 (3m03fnblji'bn). b ^ b x 0 ^ J i y i * ihnb mh5*6n'b3n6
3(5930Oi6* oi*3ng*6 inijngoib QDb^hjJ^njon 5*3036360, (10190360(5 n633336 3hm-
^miOi^nb 60539006. *ftg*6 5*9m0gn6*h3, b ^ h jb n b g n S s d n js 8010(3*36 g36lj)*6n-
Cnti^nnb 5*90196333, gjb^hjj^njg t)md^m6j6^b ("Distress") g* 0 nb b*6n-
6bip9g35n, d i b d i d n c n b n h j b j c ga h j b n b ^ j S i j j c m d n b 5*9*0+19*33630 353^0303 ^
("Eustress"). *9 nôi9 , b ^ h jb n , d j ^ j j b s g* h j t n b f j j S ^ j g n d i 36g* 5*6306000101
3(1013*630101*6 5*39oh9n. 5*801063336, 013 ah* 6 ^ 3 6 0 6mh8ng*6 9(10136001 5*-
9*bhib, 013 833^533*9 1530003636b, g*0mj)ng36xni 635*^n3hn g* im'bnôoho
(13*^503606 c gs h g m d n c n j q j j f o j h m d s ' b j (nb. 6*b. 6 .3 4 ).
6.1.2 hi beDbi bibhObflb 3nhm&069n?
b^h^bnb go6*9n53hn 5016(3303(5009*6 5*9m0gn6ih3, g*(j3nhoi3nb ghmb mh5*-

6n‘b8ob*oi3nb g*0 *b*bn*oi36xna 6;>hi3330n 6,5*903606 Oi*60 n9 g 33hm6 * (6*b.
6 .2 ):
5*65*006 qji'bs: b^h3bob 6*h9 m9 m6 nb 3nhm6369n bg3&* b*bnum|5b0m sj-
1503111606*01306 6*^nhm b^h^j^hxri (9*5-. 3(101^306360, 60019396(1*6360) g* 0336-
jljn]hn (6omjn9n3hn 3(101(536360, 6n3cnn3h36*cn* 1530*) 3hcn33036nb 93b^*6n0n'b*-
13ns. 9*9^30(101*30 03*^01(106 d0o3ho g* b6h*qjn 9mj03936* 06333b 33h3gnb
3cri0n*6m&n6 96n03630m3*6 g*h(j>333*b (g*9y*30 93j*6o'b036o6 *9mj93g36*0g3).
5*65*006 cg*‘bn6 03*11503690 6936* 6^ 36 30 0 (13*;)(3006 g*6*6yn6n6 5*00130063-
6*. >9 ghmb 6030^13(136*01* 13300b 5*^*601031™ 3(101(536360 6^*11601636 *6*6m-
03(1366. 013 g*0li)3nhoi*3n 03*^01(106 06(536603016* n5n33 (1636*, n6y3&* hjb^n-
tijynnb 3(101(53636^ (10103001*01306*15 g*0*6*6n*oi3630o* h33*h*lt(mh30 n *Jii»n3m6*
(^1*1^3063606, *6 g*0(3*3n 6*3(1013606 denovo 606013^). 36 3(101(536360 5*g*gn6
3 fn 9 g 336 cn - 6n6^(pdgj^mdnb g j * b * 3 n .
6n6**tp9g35m6n6 <3*'b*0n, 836*6*8n6n 9mcn6m36n036o6 93936*g, 6936* mh5*-
Bntignb 6o6**p9g35m6n6 5*63306 36*hob *9*^ 03^ (5*8016(101016*). *9nb 9x 3"
b*g, 3o3b3g*3*g b^h^bx0 qjij^mhnb 9mJ93g3&nb 5*5hd3036nb*, 6(i)*6n0n'b*ynn6
3hmi33b36ob 5s*j^n3h3&nb 93033m6ncn, 000^633* 6n3ain3h36*oi* 13300b 6mh0*0-
n'b^n* (835336*). bv|)h3bmhn6 9mj03go6n6 936y33o»n6 939g35 0U36*hn6 5*d-
COQh3&30n 6n6**(p8g35m6n6 36*hn 33nd03&* 936*hfi36g36 6*h>)333on ghmn6 5*6-
3*30(^6*00 .
- 321 -
9 (3 3 6 ^ 3 0 6 0 LGrtObTF? 9 ^ a n 96fn3 n 5630
Alarm Phase of resistance Phase of

phase exhaustion
_____________ Duration of __________________

stress factor
fob. 6JB b^hjbnbi gs 3*6^3 3(j36ihnlj Uibjbnb 0mg3(yi (Selye, 1936; Stocker, 1947). b^^b^co 031J-
ijmhjbn 0633336 bibnuPHjbom 3hmu363606 g3bî6n(Yi'bi(}nsb (*5i65i3n6 qjitii") gi 9|J36ihnb 0336-
JlSrwn6iC3hf' iiyismbi 3U3fo (6^36300 338905 3mgnb “h3'bnbj361l30(ri6n6 qj^’bi": 8(336^3
35336* sbjc bn^3^jrub gs *93036366 b^lrjbnbigSn 5i9do3mbib: 013 6,3(136300 'b33mj9og:j&i b*6-
Smipji, 6)536^3 b6hi<gig 36(136936;) ubpn33p3mj93g36nb 6mh0ip3h gm63b, bmpm 013 ^3601^9393-
6b 5s6bii)30ih36ncn 6165(1d(yi3ni, 33bsdca6oCni 5^63001^36 933^333^ gi'bni63&36n (Larcher.
1987; Amdt et al., 1987; Tesche, 1989)
n3 330oib333b9n, 013 bjjhpbx11 3g2>rn3ih3ni&i gngbi6b 5hdxg3&i, i 6 3 nbn

n6^36bn3m6i 3 ^ x fT'^ )> 3b35ih3 &igignb bsdmqjnwnb cgsbsdn, 06333b bb3i
btjh3bmh36nbig3n (3 s&., S ih ib n ^ n ) 6n6iiC>3g35Pi&nb ] 5 ihnb gsj33noi36:>b g i
3 nb 6iigh33iQ gi(p333ib. 013 b^h^bmhnb 0frij03g3&i ghm^bncnni, r)h5 >6 n 3nh3i 6g 3C 3g 2>m9 ih 3m6 ib gs, bi^nhm3&nb 03801633-
Time [ d ]
fob. 43 <3n^™i63n6n(jmb c^'bpdn fShgncnb sd&>& Oxyria digina-dn 9nbn d(xi3h bnEôico^
3jtoro6flh36rib 338935. ♦ dc°Dhn 5i 6ioi3&:>; o b^b^yi 5^6^0136^. dpnsh 5i 6iOi36i'b3 C0 2-nb :>bn3n-
C ibn i 5nh33pn33 geob (Ivpjunnb g A id n ) 85330*1^9 3038^ og d u i d y S p> 9y i e c o b n y i
6»hi33X 6nCie (iCÔ S1* ®>fo)- bahslxgiiynhnb. *63 dpohn 516*713606 Vjamja3e]6nb
33apJ,m9n 5>&Moe36ril»b gm^T)bfi6oi3(bn n6y3&6 &&)ohx! iCD^- <b"5 QC3>> 6n6ionc3'b3
9mo^3*>D&X <1 q}ran(v)6n <430010936016, 3i 9n6 hr^js bjb^ (63636(103) 5^6^0136^0 dymqjn <grocnc3dn
hfi3&nb6 863*63 (Engel et al., 1986)
- 322 -
UftfoOM): ^ ( 0^3036 b06 ^ 6 D 80
3*9n, bg3&* h ^ ^ jh iyn n b qjibidn 6 *h3 oij36 o0 n 9*^ 0*636 36^ 5*gi0ib3i.
013 5*63*‘boij>*g36oi y3X ^ *9 9*530(1336*^ ogn'bnmpm^n^hn o>3ipbi'bhrvbncn
b^hjbn 3 m{,3330n636i hmj,mhu mh5 i 6 n'b9 nb gidi& 30o 9 gaf»i9 ih 3m6 i I hm030nu
gib*6ynb3 n 06333b Qjb^^dnpnipfjmb. 3 38932) - 6mhdscnbmn$b g i hjbnb^j6^j-
pmbnh ]6ihnb fcibhgib. brigm 33j>336*gm6nb 6;>h i3 3 X n i 93 pnli,x nk &i(>i0ib3nb
330cnb333i 3 n - q)j6junjh pih fp jjjjd b g i 5 m(ynb - bn^gnpb [ 135 ].
bvjhQbnb bn6ghm3nb cgi^b o b i j 3my3i6nQn Smg^pn ihnb 03iy»g ‘bmîgn,
hm030n(j *b*b*3b 33bidp3& x 8n8gn6ih30i6*b* g i (i3^g0^l5nD^b- 333d03&30n*
3m330mgfncn n3ib, hm3 y 3 X i b(j)h3b x 3hr>n53bb 3jm6g3b *0g*{,3*hn 3n8go6*h3*
mb*. 0138(5* ihb3&rn6b b*58*hobn 8*5*0001360, (1018036015 ib ib i3 3 6 q)i<b3&nb
5ihj)33x oi*6*8o8g33hmb*b (6ib. 6 .3 ). *06*01, i 3 g*53oh33b3bog*6 5*8m9go6*h3,
y S X ^ D 8 6 0 8 36 30 ^ *6 0 9 * ^ 3 6 * nbii, hm3 bjijhQbn ghmb3 gsdm ^npjbjpn dmjpj-
6ii. *80^018, b^hpbnb Sn^h 5i 3m633x n Shmu^baftn 3693 5*630^00101 3ionn 3o3-
gn6ih3m6nb ib S jJ ^ n , hitf 6n36*3b n3ib , hm3 6m6*b6oihoibnb gn6i 3 nyib
>>i6m536nb fciSm y^^&i yr*i330oi3nb ih ihnb 33bidc3b3pn.
6 . 1 .3 hFldPlh ^ a n 3 fll5 6 n 01 biihObfl
bb3gibb3i 8(536^3, 6363i!>oi3hi0 gDib3h3n6nh363cn hoijijnnb 36ihngi6

(‘hjsfonnb 6mhds“) 5i0m3gn6ih3, 3hon &*hj)33X b^lvjbrnhfto hQi^nh^bb ih>
3haij,3ihni36ig. biCiD3Cn n6gn3ngn, o3 nb 0nbx3ncn, 0*3 h01501(10* 3 nbn ibiyi,
ij^nsmbnb gi 8353360b bîgni, 630n6ignb* g i 9P3-(?*3ob SQhnmgn, b ^ b x
q3iJi!i(Tihl33 (13*50(1366 5*6663*336x^9- 8o3b3g*3*g f>3 nbi, hm3 b3 nhig ogytamhob
0mJ33g36nb bndpr^l^bi g i 8u36ihob biiib^bm hQijynnb n6^36bo3m6ib 3 mhnb
*hb;)6m66 b id in g jphj,n ynf^gijjni, 3b n9 ib ih 6 n36 i 3b, hm3 ym33Q 50165(13-
(IX 933oib333i3n 3u36ihob 9*'bn*636ob bndoo3(i3 333b*6*03&* qjij^mhnb 3 mj-
03936116 n6vi)36bn3m6ib. *5(130133, ih ihnb i3l5nC3&3Cn- hm3 b^h3bnai 5*801633300
l33CnC3636nb 3 x D 6 n oymb u*08bhn3ig 5i3mj,33Oin0n g i3i<bni63b3gn i6 g*8u*30
jP3Q36i. 9*9536* n3 nbi, 5*6o(5gob 013 ih i 81536^3 b ^ b b , 33 b*dQ3b30 ni 3bn-
Qmg 9ob 601(18*03(1 j(533i<b3 g*53nh336ob 838935.
ihi6mh8i03hn g i^ n h o ^ i 33nd03b* ftiSmnbi^mb 8h*3*0n bnd^^dnb 033-
30m6ncn, (1018036015 03bli)*6oco'b*i5onb Shm^bnb 9*g536nb bi33i036ib 63 ^ 0333^-
b^jhjbnb ^hn^jhnjdjdb 31,301360*6 ^ 3 0 1 3 3 15300036360, hm3o36nu gij>i3-
3nh36xoi h33ihiunnbi g i h3<bnbl536lj x m‘!,n^ 33ji6n'b336oii6. g 3b(5h3Jijnxf> 9^
imbbijhjliinxf* 3hrnj3b3&n 3hai3i630ib <3*11*336 g i diom 3ha»3i63Cnob5i6 5*0rn-
biCW33î hoi3Qni, 5i6bij)30ih36nai, hmg3b*u nbn6n 3hoighmx>e 801^0x3^36.
^fpnbm 6jdnb byjunqjnjjhn 3033^1,0, hmftmhu 63bn, 3Qo6g36i j)f,i65h3lj30n
9mJ03936nb ig&nc3&3n. *b3, 8*jppncn*g, 3 bnb dpn3hn higoiuni 06333b (joQ^
*,r,inQ36nb 3336 hi6 nb gihp333*b, bmgm ^mjbo^ho nm636ob yn6(J36(!)hi(5nnb dgn-
3hn 0*^36* - 36<bn03hn 3hm^3n63&nb g*‘bn*636ib. i j x ^ 6*3n3gn6ih3, 33ndQ36*
*g5f>Q0 3jm6g3b i 3 i 013 n3 gi^nha^nbicr^nb b33ijnqjnj,3hn 6n3636nb 6 ih 3 mJ3 -
6*b. b!J3UO<30j,3hn h3<bnbli,36(!,30fn6nb 83j*6n<b836n 80^(5*336 y3X i <336^03(1
90163b. b3 nh*g 3 *cnn g*6y3&* a*3m63330ni 53636nb goq)3h36tjni03hn jpij-
^0336001, hm&mhnu** bl|)h3bnb 3hmvi3n63bnbi g i b33yni03hn o<bm36<bn936ob bn6-
013^0 (6*b. 6 .4 ).
- 323 -
3006^0050 8 ^ a n 9^fn0 n w 30
Changes in die
70 kDa subunit of the
A 9
CAM
- ATP hydrolytic
subunit
0
tonoplast ATPase (‘ 70 kDa)
8
Increased activity of
the tonoplast ATPase
7
Accumulation of maJic 24 kDa peptide
acid at night
6
P E P carboxylase
Occurence ol the 5
24 kDa subunit of ’ 16 kDa subunit
the tonoplast ATPase forms proton
4 £ channel
m RN A for
P E P carboxylase
3 ©
Proline accumulation
Pb
2
NaCI accumulation
*
1
fob. 6.4 bvjhjbnoi &s3m633X n SmcnîSnpijdnbs Qi n’brnjBbnS^ftnb bn6oi3'bn (Heun et al., 1981;
Michalowski et al., 1989; Beck & Luttge, 1990; Vemon et al., 1993). bAlliumcepa-b <33633600 jbxjah-
i'bib 3C3j^hrtcg(Tih3'bxî 0ii3db3&3h3d:>6n. (C) "jm^hmen; (Pb) 200ppmPb-ncn 5-gcni6n ^>30^1^3393-
6ngi6 10 ecrnb 338^35 (Maier, 1979)
b^hQbxn 9g5mSih3n6nbo>3nb, *51130133, g*9*b:>bn:>ai36xn:> shsbyjijnqjnj/jhn

5^00130(163636(1. 33nh33Q3b yoi30fib>, nbn6n 363363636 g ^ jn h a tjn b b n d cn jh jb .
3hm|53bn 831^036^ ih*b33U(iqjm,3h:>g fi>noi3:>0oib, 013 b^H^bx11 qjijiioihnb ,yi 3 nb-
5*6 9*9013^9360*9, yoi330oi3db 3hoi6>nh>9 (b,j3h3oi,jn3x > p ) 9 n0 gn6 i>h3 m6 b.
b^h3b30n 9g 5 oi0 >h3oi6 nb *h*b33unq}n>,3hn 6(19636(1*: 3&bn93 &nb (5*6b*j,3cnh36noi
33hmjbnei'bnb, 5p3<s*cnno>6hxoiy>^,fib, 933(igtioi*b>,oih6 *li)h3g3jlj*'bnb) *j*i>n3m6nb
1530(1036*; 3oi0n*9n63&nb 6(1011*16013^11; *6li>noijbn0*6lj36(ib (ib^mh6n6nb 83*3*,
^myno53hm0(i), b^h^bnb 03^s6m0n^36nb, oib9 oib3h*g 6 ( 13 0 1 ( ^ 3 6 3 6 ^
( 3hoipn6 n, 63^(1636(1, 3oi0 noi03&fi) g i 9h*3*0n a^mhign 9 i336bh3Xf> 6*3hovib
(3m0n<336m036n, *6^,011511*636(1) bx^>b*0(i bn6oi3'bn g i 9*5(101336*; b^>h3b30(i 5mh-
9m636nb (*6bunt>n6nb 93*336(1, g*b9 n6 nb 99*3360, 3010036(1) 6 *h9 oij0 6 *. *h*-
b33unqjni3hn> *5(130133, 0306h*636nb oi3nb3636nb (0396h*6nb 301,53615(1*0(1, 6(13-
01113(136*01* ,ihb6 b3oihljn) (530(1036*; b36oij3db n6lj36bn3Pi6nb 9oi0*^)36*; agmôv-
bii6oi3‘bnb 9*01115363*; 6n3om3h36*oi* Shoig^nnb g*6*0n 90163; 'bhgnb g*he'-
333360; *h*bhxn 60196110336* g* 6**gh33n 9*631136*.
b(!(h3b3Cn 9 g5 Oi0 *h3oi6 nb *gh3X n 5 *9 oi30 (i636 nb 3(1933 3hom 0 *R3363 & x n:>
363h&nnb 9n6ng36nb 930(5(1(136* (93^*6oi0n'b9nb g*h(?333(ib 5*0oi), *6 33h3g9n
h33ih*yn30(i 3hmi53b36nbioi3nb 363h5nnb 9019*^,3630(1 b*hj3* (6*b. 6 .5 ). cgmb-
qjoihnpnho&nb h3ij^n36nb 3 3 ^ 3 9 3 ^ 5*001, 9019*^,3630(1 b36oij3db 3nhm6369ni5
yi, b*j>9*hnbn i^qj-nb bn6oi3lbn *h 6936*. 363(150^ bî^bn (AEC; [7 ]) (jmpn*:
- 324 -
b&fnObO: b06«?rtn30
6sb. 65a,b ddn03 33(iie3&nb 5^3036001 b^lvjbn. a Beta vulgaris-9n 3^6gta(xVb

Sjoi^bjbnbi q1 ATP-nb ihmgjjymb gb^jncnjbi HgC^-nb ’b^Smjaoejbob 83935^ (Luttge et al.,
1984); 6yic9(j36bh33n (Euglena gracilis) be^Snfo^nb bnb£)dnb o6jh&nnb b ^ ^ b n ,jmjbni,;jhn 8ih-
nC0*f'b b3&c3î03h im6(j]5^hii}nb'b3 fb jg i jhiojnyi: 50p.M ZnCfy 9p> fchiejnyi: 0.1 JiM CdCl2;
J33ei ahsqjnyi: 0.01 |iM HgCl2 (De Filippis et al., 1981)
AEC = ([^03] + 0.5 [igg]) / ([i^qj] + [agg] + [1803]) 6.1
^cg/ipo) a^qjihg^bnb 0.6-b] 91691 9115380b ghmb, 363h^nnb bî£)bn 803-

0100136b bni3mijbgnb36ihni6m6nb g i^ n o iQ & i^ . Qomh] 9bhri3, b ^h ^bx 5nhm-
&363n babnismijbcm qa^j^n^nb 6mh8i03h gm63<b3 83^611163636919 8u36ih3b
b^nhg36i 363h5nnb 9181,536 noin 6yihm3&o. ibQ, 8151900119, 3 x >rn<3 nll)3&3n 83-
n6n363&i 363hj,nnb îh6o gi6ibih336n 8386hi636ob ^03->'b36nb ijiy^mbob ib>-
SiCT136919, hm0Q36nu 3633016 8ihn93&nb nm636nb (jhi6 b3mh^b (5n(jm5Qi'b0ngi6
3 1 ^ 9 8 0 . i;)3gi6 5i8m8go6ih3, 9;)ndp3&i 9131^3610 1, hm0 3iQmqjn^36n 8iho-
C36nb Sipign 3^16(536^1150360^980 h3‘bnblj36(j3Qn ihm6 g i ihb3&m636 0396(130
gi^3nhoi3nb 30(10163680.
y330i nb (131^01, (101839015 1 ^ 1 ^ 3 9 0g5m0ih3m6ib 530633636b, 3Qbid0363cb
bgnb 5i0m3oy36mo» 9u36ih3cni 85118601601(13 bib3m636o, 51(13801 qjiJ^mh36ob
gi^3nhori3ob d n m n S p n bmgm 8(5361113, 16 8obn (5193339° 61609360
(815., 08016090^36390 D3h3gob ^,39^^360) - 6om<bm6g36ig. 810m 5180^36360-
bib 3691 53ibbm393b, hm8 b^h3bob ^ 3 3 9 0 goi56mb(yiyib 6086360 83089361
51801633390 oJ63b 633(10 bb3igibb3i &i8pn,bni6369nb 803b. i80(jm8, obo6o ihi-
b33(5ogo*(3ho 60863601, hnj ih 5318933^ 50163(13(3390 b^h3b30O gjiJ^mhob gig-
5360b bi83i936ib.
6.1.4 UbhDbfl Q) 9l5D6ihnb bflbnbbCD
6 .1.4.i biShDbn annum aonx nhb^frbab

l^f^bn 8mj83036b 8(536ihnb 80139 01(15160^8^3 818061(5 30, hmg3bn5 giôh-
0131b 8(536ihnb 0bm9mg 3hoin 616090 51601390b. 8i336ihob mh5i6olb88o blt)h3-
b390 h3iji3n36ob 3mmhgo6i(5nib ibmhi50393636 a}n,}m3mhdm6jdn. (101501(1(5 j)0 8(53-
- 325 -
9006660050 gânaaftan M ao
6*hob hm03Qn33 6*6n0O b^h^bx 6*6n(3gnb, 3mh3 m6 *03 h b n b ^îSn

930603636* 33b*b*3 nbo (5300036360 (hm^mh^ *K*b33(5ngoi3ho h3*J(5ru>), hm8p3-
60(5 3(5oh3 ghm3 n 3(bh36330ymqj36 33(5*60103(111 *j(i)03m6ob 1530003636b; bm0m
b(ih3bxn qjiJijnhob b*65hd0O3*g 8 rrvjS39360b ghmb - 3^1363309010)36 bnymu-
b 0 o b 33 6 *h fi3 6 36 n b *O T 3 n b * 3 (5 0 0 3 6 3 0 9m h<gm 0m j,O3h-j)3 6 3 ^ n i3 h *j^ 0 3 m 6 * b . 6ycnb
03030(50^0 qj3b336ob 1^ 6*30 (ob. 6*b. 6 .59 ), ^ 5336 0 6o30i03h3636nb g* 3*656*-
gob 6*5036016* (nb. 6*b. 6 .51 ) 0633336 *bo3o0*d)3&ob 5*6*6o036ob g*h(p3Q3*b,
3n6ob'b3g*/3o6obj333* 6*6o036ob 33g*hg36ob 33 (530*b, *gh3X y3*3O0m6*b g*
q3moim0(5336*b. o3 ob (5mg6*, hm3 3(536*113 b^h^bnb 30(10163680 3cn0o*6*g n p ^ o b
3 ob&*6 oi*3ob g*b*h633*g, 53*od3036b cghcnbooig 803395301 8*11,503 bnb^3936^)3
R*O)*h3630O 3jb33ho336li)36ob 33932,360b 2>*g*ti,*6*b 801300 3(536*hob ^(533*^3.
6 . 1 . 4 .2 bibhObfl fl6 3 3 3 b 90130(1 n h 6 ^ 6 fr b a n b 6 ^ h Q id a 6 ^ b
b^f^bob J338 3 ymqjn 8(536^3360 3hoi3*630iob 8018033601 g*l53ohoi336'b3 g*

5*113801 q3*j(jmh36nb 013001 63330 ^38(11^83936*^1315 50 (13*50(13636 6mh8*p3hn 3 n-
hm636ob*oi3ob g* 3 *b*bo*on3&30o j^33ob5*6 5*6^33*33630*0. 63636*80 8(536*-
h3<b3 3hoio 50165113^300 b^h3bmhob 3oij83g3&* 0830*010*. (101501(1(5 &3bo, 8(536*-
(13^33 3mj33g36b b(jh3b30o cg*j(jmh36ob 801300 5m8303jbo. *b3 8*5*0001*9, 8bnb
8*r*0O h*gn*uo*, 5*0^3(136*, 53*03* g* 8*ho036nb j)m6(536(jh*(500b 8018*^36*
6*h3m*g536b 3hcno 33mh3boi*6 33 ôg30 O b(i)h3b30O qjij^mfôb 50i86n6*(50*b.
>•' d,1^ '-j 1 > 1 <j 1 c r” ' a1 0 20 40 60 80 10 0

3 f I S 3 2 i l l !
Water saturation deficit [% ]
fob. 6.6 a liynb ^39* bi'bf'^h'b] QpVbihpn Pinus cembra-b 60633600 yo&3obig0n gi
5s8m8hm6obs93o h3<boOl|>36£}ef,i6oO 6003(10 906*801^. FR 1406300*980 h3'bo0,j36^3ero6:>:
hi, hmSxtOO 6063360b 5-10%-8o j>i6fnobi^36i 5*^063001 5*8^633300 9^0*636360; DR j>*9m3-
hm6nbig9o h3'bob,j36£)0(Ti6i: d^'bmgocnb 1-2%-oon 9*130*63600*0 5ypob 333u330fndnO 3*833636300.
5h*qjoiob 9s8^hob3Cn 36*60 3mo(js3b 6pob o3 SohnmgO, hnnj* 930^^036300^ 003380 15-gcn*6n ^16-
3360O ihb36m6* (Pisek & Larcher, 1954). b 01536^33680 5*1406300*980 9s 5*3m9-
hm6fibigdn h3'bob^36lj3C(Ti6i; R Raymonda myconi p* PPolypodium vulgare ‘b*3oa*h3o g^nphi^nh}-
630 3nhm6368n. i53333h*»j3t1X 9*^0*636*9 80(^36300* oO ‘bc'3*hn, hr*nj* qjmoicoO 10% 9* 83^0
^30*6936* (Kappen, 1965)
- 326 -
b&tfObO: ^66^3036 bOBÔSt)
20 30
Temperature [°C]
6ib. 6.7 0ihnp3&nb bb3igibb3i imfjojB^hsun^&nb ’bj&njQ^pD&i qjm^ml)n6oi3'bnb iM93:jhitK)hx

es9myip36x3&i'b3 (Havaux, 1992)
^1130133, qjm6nb 836;!86390 3ohm636o, hmfjmhn^bb 5i6ioi3&30m6ob gpn^jhn yny

0n, 56 630n6ignb ghm^bnb 1330^ 5539365b 5693636 3^36ihnb j 13335^3 b^h^bx
bn^3i(3ri363n: b^hgbmhnb 8(1^839365 63636580 556bb353g365 3 nbn 3mj33936nb-
5556 0Mbmhi^mhn30 3ohm6368o (855., 3hm0o6nb gij)hm336i ^1303001 95^30(10139
3j336ah30n), bmgm 3353^>unnb b3<bm63hn hn^330 m6 i dnh330ig (3390b 3(336ihnb
jl533bb ihi 3 ih^m 5 9 0 8 5 ^ 0 , ihi 3 ]g 63&nb3 n3hn bb35 cgij^mhncn g5vi)3ohoi3ob
ghmb.
b(ih3b x r> qjijtjmhôb 311136060113630 g i cn>63 n3 g 33h x 3 mj6393 &b(b 3 8(336^3
h3i^nh36b 5^ 0 ^ )1156 36^ 3 3bb3bnb: frsdpnjhjdncn, djh&npjbncn, fr^gsqjshjncn gi
djbh j6jbnaix 5 in. 5^330^3 b8nhni 9hi350n qjij^mhnb 0mj0393&nb 8393559 6>
6(11633390 5id0n3h3&30n gi3i^36noin 95^311(1013360 0633336 9585^36(101
95h(p33336b g i id0n3h3636 dnhooiigo b^h^bmhnb 5539 36^ . 63mh3 3bhn3, bli)h3-
b x 3nhm6363n 83069365 6ih8moj86ib h33ihii3O30 n 013 Ii3'bnbli)36,j30rin6nb 5i3id-
Qn3h3&30n 3hmi33b36o, (1(11693611(3 0633336 8111394119630 bn63h5nbli)3h1 i6 56^55^-
6nb^3h h3ij(3n5b: 15030 1b i 3 oihnb 8jm63 h35nm63&ob b3 815365(13360 yn63nb 553-
0360b 33g 36ig 6i yn63nbig3n ^m03hi6(i)30m6iboii6 3hoiig 063636 fciSmShm-
6nbig3 n 6n6i5(p8g35m6nb 365(16 (65b. 6 .6 ). 3iho036ooi 95^30(101356, 060695 80-
6393001 013 hm5mho5 6^(13606 6565(^9030165 95 06,53660311165, 83060365 5i‘bih-
gmb 6015030^980, 5igib3h3&nbi08n g i 553m3hm6obig3o 8g5higm6i (6ib. 6 .7 ),
56 33ih6o0mb 8(336ihnb yo03nb 8n8ihoi 5i8m6hoim6nb Shm^bo.
dhsjQmbncnn hjbnb^jB^jQmbnb jdshnb J>i3m333i336i 3hoio b(jh3bxn ojiJib^-
hnbig8n 8353360b 3hm(33b3n bSohig g i ^ S o h ^ x n i 1509360^ g i 60(11838-
6(1563606 b^h^j^hob 831330560156, hitf 06333b 3hmli)m3gi'b 8 ob b 015590 bî6n0-
3hm6ob 36ihnb 5650193656. 35ho036nb, 55gib3h36ob g5 535930b 6,5(1363606 ghmb
5(163639^ 3hm(j3o636ob b,5(13^3(1390 8b5i3b36i, i06ion, giii38oh3630oi ^30 1
im5gi3l5ii303h <336(116360156. ob3oio 5115633(5003033(10 (135:1150360, hm5mhoi55i
qj053m6mo036ob gi5hm336i, i6 33hmjbngi‘b36ob 555:^503365, i5h3oi33, 0633336
3hi3509bho3 83533636b (39^(15006033(10 (15905(5006, bm3m36nbi g5 5idm3ob
6n6ii(p3 g 35).
- 327 -
9(306660050 LSrtOUT)^ 3^ a n 86 ff)0 n 6630
6 . 1 .4 . 3 biShObflb e ^ d C 3 3 (lb b ^ b D h f l
b^hgbn ih(?333b oD^Jbrohig 5>chhor)0 S m cQ ix ^ ’ 03 ^091)0 Qb

mh5 * 6 n'b9]c 9^6336^3. £,36361^ hgSihi^nxn ij^n3m6>, ibi0n 9gj,m3 ih 3 m6 nb
fii3myi0o63&i g i 3qjhm gngn 6o6ii(?0g36fn6ob ]6ihnb 336^63636* 0mooibm3b
<b3g03(4 053h6 :Wn.bDC g>5 ibih 336b.
9U36ihnb "biboumubgm b ^ i ^ o i " 1^ 1 ^ 3 0 0 3ohm63&ob 3jm63 3ib n ^j) 9 n 3 o-
3 bhoi30nb ihi bhgnb dsjbndjdnb 8op633obi2)36, ihi03g bhgnb bsdy>pm n6^j6bn-
jmbnbs g i bsbnijmijbQm jbicghcnbmjdib dmhnb i,md3hmdnbnbsj)j6. 01536^3360,
hm0Q36n(3 93353360*6 bi *13360 6030103)13636001 c'*ho6 gi bo33h*gobij)36 3 ngh-
32)OQ 6niQij,b, obhg36oi6 63c* gi b3 oh*g 3i^ ih i bb330O iJ30i, 3i5h i0 (0o}b3gi-
3ig bii33&o 60301031136360b 6ii036m6obi g i Sygob QQqjnuoôbi) 0 6 ^ 6 36 36 36
3o63hi03ho 603010311360b 3 i p i 0 in e^ ^ h itfO ib gi bii>0ihobig 3 ip i0 6ypob 3 m-
^ 36 uoi0b- ihoggco h35om63&nb 'bmj^haio 0 ^ 3 6 ^ 3 (i^>Jii3^36o, 6im6i6o) bb33-
gob bo3b3&3jobi g i (P3hm<b3 gi 3 l3330O 3038366360b 33)10053110300 5i60i53&ob
60-
cr.\
100 ■
80 -
S£“>
5
•2
§g
c
50-
rX : ' •
aT
w 00 ■
^<0
• • • . / V • •
40
NO
®
■a 40 -
• ro
2o n
0 *
30 • •
# 1
0 - 1 1 1 ' 1 1 1 1 1 1 *T
1 1 1 1 ■ ■ ■
10 20 30 40 50 60 20 40 60 80 100
Tannins inleaves [mgg’DM] Frost resistance (relative)
fob. 6.8a,b 'bhpi gi ho'bnb^jEijX'mdi. a ^mSojjxn b] 9u36ihnb Cecropiapeltata-b {,]6r\yi3n 9gn-
eihm ,3160636001 pi 3b 8(5361)13360 35301 ihni5 eXiX0 963h3&nbj>i6, 0139(31 030011113635 ^cghm
9^nh3 hing36n6noi qjmoiQQiib qjihcnmdnb 3)101330^3, 3<iph3 ,ji6n6nb 9(jnh3 hipip35m6nb 939(J33(V>
535m^n3nb 6ih3m9ie {)36c36n (Coley, 1986). b Pseudotsuga menziesii-ob yn63i{(i 3do3 i>36m,yi3ob
n p 3 m 6 n j3536 n 0 ^ 9 3 6 0 16 o q jh m 6 3 0 1. 3 1 ^ 3 0 5 0 3 3 b ib 3m 6 n b, Q ifrh iS yn6 3 n b ip 0 n b 3 6 b r y ,]h n
636m^n3nb 6ih9maie{>36p36n (Braun & Scheumann, 1989)
33833^6001 i3(3oh3636 bî^fi^ho bo3 yihob 036ihR3636obioi3ob biôhm 363)1 j,n-

ob gi6ibih336b. 3u36ihob 3î030o (56013303606* gi 3ihi‘boli>36obi5i6 oii3-
gibi(ii3ig gi 83^0300 6i6o036ob 03bi6i(53036ig ib333 biôhmi 36311500b 0633-
bôh36i (6ib. 6.8i). b ^ o q y i^ h o gi6ibih33&o 03od03&i 5i0mooi3i0mb hm&mhu
5 C 0 2-ob biôhm36i 15 g i 0U330n 6i3hoiob bo6oi3'bobcn3ob g i 03bi6i3obig, g i-
2,ih530O 0ibob i[ng536obiOi3nb. 03mhigo 6o30i03h3&36ob bo6oi3(boboi3ob biôhm
- 328 -
U663U0: ^66^3036 UOG^fnOaO
(jbhnpi 6.1 3u j6ih30tt Sîibmp^hn 9i 6jb ih j36n a^mhien b}6b^>6(sn36nb 6nmbn6on3'b3n (Merino
et al., 1984; Gulmon & Mooney, 1986; Williams et al., 1987; Diamantoglou et al., 1989; Kull at al.,
1992)
33,ji6nxj3hn gifbbshjri
6tj3&shnb 6i6n^n (6*1 bClnrfb* 99hipn 6ro6yl)
9i8U33Cyi I>j6b*s6un36n
1^1606360 1.55 - 1.6
[30^6^536jhn 5cnj)m'bng36n 1.9 - 2.1
sCi)i0nine36n 2.8 - 3.3
3m6m^3hi36fnne36n 2.8 - 3.5
C>03jbn 3.32
33hoe* M W * b36b^6(jn36n
Cn&6n6n (6n63m3i 6ons BjtvjiCSn) 2.44 - 2.49
|Vi56n6n (qjih3coi3b(yn3s6oii 03hjs63n) 2.48 - 2.52
£Q5nc9m6*u3C3 nh&»6ni3ta
hbnpn ajmcnp36n 1.3 - 1.8
biC3hmq}np3hn qjmaip36n 1.35 - 1.55
6116336(1 - 1.5
ihi0n66n6n'bfih363cii
C^hmoin 1.1 - 1.35
pnj6n6n'bnh363pn p3hm36n 1.4 - 1.55
0î6mQ3h(i gi6ibih336n b id i n g 9i(?i0ni5 nymb (ubh. 6.1). b9 i 013

n9 b^hob^cn qjijv!)mhnbig9 n 6n6iip9g3&m&nb i 9 i(p03&i ^(130133 9mj93036b 603-
cnnjh^biOU) ShriQjj^ns'b], tihgibi g i 5 i 9 hi303&ib 3 : Sphnmgxn 5 ig ib 3 0 i 9mb33-
6360b qji'bi9 n> hi(5 i X nCD^DCni yn63i5i9d03m6nbi g i 6ycnb goqjnisn^nbigQo
0Qj>higm6nb J>i9mbi939i336pig, 06333b n9 ghmnb 930ijnh3&ib, hi 33hnmg9o(5 63-
bid03630n oj636mgi ibn3npi^36ob Shmg^Juni g i i 9 n(jm9 , bhgnb 3hfnjj3bn rfb-
(T3Q3&i (6ib. 6.86).
6 .1 .4 .4 6 ^ h R 0 6 i b ib h O b X 3nhP1606an
9(j35ih3 &igihR36i b ^ f ^ x 3nhm&3&9n 9bm0mg 9i 9 o6 , hmg3biu: 1 ) ihb3-

6m6b bi9 n9 hm3bnb ou>3ngi6 i|yiQ3&ob (b^hjbnb shngjds) 03bid03b0mdi; 2)
b^h3b9n 9 ymqjn 9(j36ihnb bb3igibb3i 6iBn0nb 9 gj>higm&nb 36ihn (b&hjbnbigdn
hjbnbtij6$jQml>s) bi^ 9 ihnbig 9i(pi0ni g i 3 ) b^hQbocn 5i9m63330n gi“bni63636n
93d036nbgi53 ihig, bh30 ymcgn0 ig i,m9336 bnhg36i (h3b(jn^3unnb 35 ihn). 9 ih^03ig,
3b 5i9mnbi(!,36i ojmh9x noi:
ftig ih R 3 6 i = b ( jh 3 b n b i h n g 3 6 i , 9g j , h i g m 6i , j, m 9 3 3 6 b n h 3 6 i ( 6 .2 )
b^,h3bob ihng^bi 93ndg3&i 9mbg3b bnjhujQsg (9ij,., 9o6nbj330i j>igi'b i9ov

Ii3&nb mhj,i6m36n) g i ghmdn - 33&3»!>il5nnbcn3nb b30biyh30n 33hnmgnb i9mhfi3-
3nb bi93i0Q6no) (9ij>., B3fi9nb 3Qhnmgnb 305393^36^. 9g^higm6 nb (Bn6i-
i("3Q35m6ob) 30ihn 3qj3d636i b^ bob 930unh3&nbig9n 9 n9ihor»x ihm^bQftb gi
9nbig9 n ^m03 hi5^m6 ib. yn9336bnh3&i (gi‘brti63&nb iP9636i) bmhun30p3&i 9^3-
- 329 -
8006660360 bftrtoiyac? aânaoAonboao
6iho6 633150193(10 616093606 (5161603606 9901(10360 93 h35363hi(5nxn Jbm 3n-

0360) 6183*036001. bSnhn g*'bni636nb 930016333100 0(5361(133606 5*91(1^33* 0n-
0*hcnxni 3D6>Si l5nD^f' 6i0h*X3&nbiJ>36.
6 .1 .4 .5 b ib h D b fl Q) D 3 H C 3 l3 n i
Igi0ni6n6 93966*3000 03^603(1363690 (560160901, hm9 b^h^bn 0 6 3336 0 3 3 6 J-

(503(10 *joo30i6ob g* 3353360b 36*hnb 5*‘bhg*b. 3b 93&X3&* 6*0oib*g35O* 6nm -
001500b 93301 gih&nbo^nb, 0*01 3nhnb, 01536^3011 3501001500610130^ b ^ h ^ b ^ cn
bO03*(5O36nb 6*50360161 06333b a v»3 pn jjn b j 6ih n b g ijjjn cn jb ^ b , boi0m o jb ^h ;}-
9i 03hn bljh3b x n 9 10 3^ 0 1336 0 b30biyh3Qni bsdmdhoimbnbbcnjnb. 60(1360, hm-
501(1(5 9*030(1013* g i 6 o o 0 x n, 06333b 303(101 y>hftig 835336300 536010033606 6 n 6
6i0m 633ib, (1*15 ^>*(15*0 0213301361 91030(101306 5(1*0036006 51663(103. 3m >}0 ilSn"
360, (10109360(5 5*601590*6 6 o h 3 6 x n q}*;Jooih36o6 bi65hd0O3 3ovj03go6ib. m j^ n -
0*03hn 3ohm636nbioi3ob 6396*9(1300 "bhgnb 60(1*03500916 (K-6 ohi036n i) 5 1 9 1 -
9016 8 3 6 ^ 3 6 3 6 0 6 6oh*o36m<b3 (r-6ohio360i), hi|j 61(1301195366 9*030(101306
l530O036nbig3n 3301931503(1 9353361b: b30biyh30 3i 6nî^3n 195090 ij3 b ^m 6-
53(136(5016 0 $ b n fis$ n gs6 ^ d m d j jj b n b b ^ hjbn ), boi0oi 3i 6ooionb 6 i 6 i ‘b c'3h m
bni09n (3501001680) - i0O0*o3h-3g*qj3h 60(1366. *96101, ih 1I16360166 ^ i b o ^ ^ o
336*616060 6 0 (136 x0 32)01001503(10 q)i;Jooihn6 51(1303. 1000010, 6oh36o g>*0m6-
*50060 30 ih i, ih i0 39 bnuoiybonb 3 3 0 1 9 5 3 6 x 0 6i6n0ni. 9013390 6 0 (136 x 0 QPd-
Ooiho j96nb 6x 3Jl5n3h 6 i,bc'3h366 9 1 63hon6o6 601501156906 9 3 6 1 ^ 3 6 3 6 * 9 0 9 m6-
16003 93301 3(101(5366, h016036015 3^ 36 339 9 0 1(336 0(5361(106 60(13606930951*19
6ibni5fTH5b0oi ijon30i6ib.
6.2 W
63bhn3nbOhOLOCfl qDtonhObfl
51(13001 03*^001(13606 6^(1360 6*h0oioj0636* 633(10 *6 6*50360 363h500b 9 n6 m-
936nbib, 6030103(1361011 (5390b bBhicgn *6 6300 0n0gn6*h3m6nb*b g i 3(5bm 5*3-
0360b 3393519 (6*b. 6 .9 ).
sbnmfijh cgsfomhjdb 3 mhob 83060361 5*0oi3*hfiooioi 5000*03(10 cg*jooih3&n,
hm0036n(5 0mj0x3&36 *o0oibq}3hoi3o, 60*9*56* g* 6y*03o: 3toob (ngoiynob
boh3bn, 51001633390 d0O3h 916*00 16 0ipi0n 5*6 10 136 x ^ ^ 0 1; d0O3h 0*Cn*On 16
916*00 0 38 ^10 3(1360 33bi6i3obo 01*609330 001303636001 - 9063001, 6o*g*5nb
5*9063001, oim30nb 91 906300b b*qj*hooi; 6*03^3606 615036016* g* 53*03*; 01013-
0nb b103ihnb bo0i5nh3; jihn; 6n*gi5ob g*00*336*; 60191590 b*533&o 603010-
3h3636nb bo0(5oh3; 60191506 03»43< 16 89133 (13*^50*; 0oidhi3n g* *h*bo*&nQ3hn
63660(1*0*^ hi(5 03Ji6o53hig 03^(101*36 0(536*1136; 053(1030 *6 6903(1 9*03601636-
300 601915360, hoi0036o(5 (j>*ho6n* 3*65619001 g* 663.
bnm^jhn b^hjbjQn 30(1016360 5*6*001636300* 6(536^301* 0*p*0O yn6(536oh*-
I5006 *95093600 9 1 3ih i‘boo3&nb* 9* 015361(106 6^*0390 (56013x36001 g*o3ohoi3o6
3ohoi&363n. 63636ho3 9*0o3nhoi*3 qjijooih36cn*6 3hoi*g, oi*3*g *g*0oi66 3 y
- 330 -
6 DG0 ?>An30 b6 Aaii3 «s?0 ^dfinfnObO
Stress factors
abiotic biotic
R a d ia tio n P la n t s
A llelopathy
UV radiation P a ra s itic plants
T e m p e ra tu re M ic r o o r g a n is m s
Heal V ir u se s
Cold B a c te ria
Frost F u n gi
Water A n im a ls
Dry air Grazing

Dry soil ■ Tram pling
Flooding
M e c h a n ic a l
e ffe c ts
Wind
Solifluction
Burial 6* b . 6.9 b ^ ; ) b x rt c g ^ j ^ y n h ; ) ^ ©* 8x n n
Snow cover
Ice sheets 3hcnn3hcn9i0rTiing36xDî (Kreeb, 1974a; Levitt,
1980a,b)
Sm*j3b 0h*3*pn qjrfbnjflhn, 83ji6nj>3hn g i £,i6bij)3cnh36noi, jnQn^hn giS^nhou^n

qjij,jmh3&n. 8icn 3 mhnb &33hni b*3 n3 n gjbj^mhn, hiQj>i6 3 icn 5n6»c<0g35 8(53-
6*h]b ihi ifob J>i0 m839 i 33& x n e^9 l3i 3n 83 }i 6 fYb33&n.
LPCJ)DDCn qjsj&mhnb pibib3Q3&i3 g i {>i6bnc3i3 ih 36gi SgjaBib n0 nb 9oi>
&3^gnp36i, hm9 nbn6n 8(^^9393636 rfbmenhgdxie- b^h3b x 3ncriih3&i9 n 03 n6 n9 -
6 ]6 i 8hi3ipn bb3igibb3i b^h3bmhnb cni68n8g33h3pn, 013 3hcn9hmxn 8(^^8393-
6i, hitf i3fi6hm36b 8u36ih30ii ih3ipb (6ib. 6.10). b6mh3g i8nb j>i3 m 6ih8mnj8-
6Q&i 9 i536ih 30ii 5i3lnj3036nb jjQnSi^jhn 9 i ggiqj^hn ojij^m l^nb 8n3h 83j86n0n
5hign36^36n g i bi*bc>3h36n (3i&., Sn^hgcn, 3mpih30n g i bn8ipcncnn) g i ijj-
h30133, “8(j36ih3xiT'i>nb iD6dxD&n" ihijntp9 n g i 8 iCf>ip8 oii3 n.
- 331 -
9 0 0 6 ^ 0 0 6 0 bfirnObac? a ^ a n a ^ a m a a o
D e U y td elon g atio n u nder s n ow (Sm. Sd)

S o il w a ter p o te n tia l below “ 2 M Po IF , M )
L o w carb oh ydrate concentrations p ro du cin g
w iltin g a t h ig h le af w ater p o te n tia l* (Sm. S d)
H igh tra n sp ira tio n r a ta in exposed p ia n o IF )
E a rly fa ll d ie b a c h , p ossibly related t o lo w
? s o il w ater potentials (F |
t
Inadequate ca rbohydrate reserve (F , Ss. Sm. Sd)
S m u t in fe c tio n (F , M, $)
t
Inadequate carb oh ydrate reserve IF . Ss. S m . S d!
D elay in b ud developm ent u nder fa ll
Dispersal p rio r to m a tu rity (F , M ) and spring snow a ccum ula tio ns tS m . S d )
O th e r u n k n o w n causes IF?, M , S?)
fob. 6.10 3*c|ic 9o'nl> 9uo6ihnb (Kobresia myosuroides) &i3h(joe06nl> cnQfHjnhî b^h^bx^
ojiiynhjdncn (Bell & Bliss, 1980)
6.2.1 a'bflb hOQfDbnflb bibhObfl
crjih-nb bnîh&nbi g i geînnbqyhn higniynnb ftidpnghgb^xyi 9 oi>6 oiJ0 nbib

81536^33680 300iihg3&i higoiyngco b^gbn. mhn33 336016333*90 bijdg 6 3 >j3 b
qjm^ni363h53l|)nj)3c 311(^153636006.
6 .2 .1.1 denohn hênîsnncn 6âne333cn w>h3bn

qjm^mbnEcnQ'bnb iSihi^n 8m6ym6npoi nbg, hm0 033d0mb bnox0 bo6icnQnb
3 ijbn 0i 03hn gqjgj^hm&noi 5i8my3636i. idoty ) 3 , £>h6o &i6ioi363C(ndnb
3nhm&3&3n bggbi qjm^mjnan^hn 363hfconb 6 * 8 * ^ 6 bh9 mj06 b, h^j 0633313 ^3*6-
(!)3&nb 5>3 my3636nbb (6ib. 6.11) g i iboSoQiôob n6^36bo3m6ob 8381501136:^
(cgmli)mn63n6 ni50i). di(vii6 3 ipico higoi^nnb 3ohm6369 o 5o&336 ,j36 obi g i
^nQiijmnggbnb b ^ ^ ^ h i *boi6g36i (q3m^mg3b^h3jijOi). 33bhob3&3ho 3ih 36 jn 8 nb
y33Cs(b3 Vje^ ShQD&ob i3X?iDXn dcmhm3C>bj>nb q)rry)fng3bl!)h;nhjn:> y s x ^ D b9 n-
hoi gi 3b 36g i oymb Qi&3h3&30fl q}mcnp36ob cgm(ynbn6cn3‘b3hn ij^ 3m 6 o b
Qij33no»36ob 3hon-3hoMi 80^3^0. b id i n g bdohoi jcmhm3pibli)36ob 3 iîh 3 &30 o
332
{>'36066030 UftrtObÔ 3 i>jJ6 P)6 060
1750 100 pmol m‘2 s'’

1.0 \ 100
0.8 i <►-o- ~o~ ~°

K , - 5- - -
0.6
Sh > * -
0.4
0.2
a
0 I- 1 r ■----- r ---- 1----- *----- 1----
0 60 120 180 240 300 360 420
ia.
| 1.0 100 2000 100 Mmol m 2 s '1

o
3
= 0.8 00—
0-^ L. StL —O
^ -------------, fb-'
NW L Chlorophyll fluorescence [ Fv/Fm ]
b
120 240 360 480
Time [min.]
6)b. 6.11 a-c qjmlynbn6<Ti3'br>b qjm,ynn63n&n|jnnb} e> ghm3n J>i6i 6nc3dj. a Hedera
canariensis-rb bn6xn(<nb» (S) gs fihgncnb (Sh) ognxncoftnb 9oe>h36>: b did&nb dcnohn
Ui6ioi(inl>ie9n aeiSdnhj&xn (StL) e> bjb^ bn6ioiQ3bois6 33633*300 (WL) qjmcnpj&nb 839^ 36^;
c ii33nhn jhomb 3bhn3. hjicjhn p> 3y)bn3i03hn <ge3iTih3bu;)6unnb (Fy/F^) qphQm&ibh pi, 9]nv-
hjb 0bhn3, qjm^m63&nb Jjb9my3636ib (a) 9mhnb {Jujfjihjou, Vj9mtn9my3i6nc mh bib3m&}9n
(Demining & Bjorkman, 1987)
Ulva Gracilaria
a e
ii
CL
61b. 6.16 bpjnb 6ybe0u36^33680 cjm^mbnfkno'bfib qjnvymeSn&fnyinb epn]hn gn6*3n^ 63636^3

3b6n^nj3n. PhN 6;fcyn-o}pyynbn6cn3V> hi£>ni(jnnb 70 % -b 3 'bcgnb ‘baQiSnh'bj; I0 qph-nb n6£)6brt3ro-
61. 6ybcaU36^h3. hm0x<X5 6n6ighm6b phdi b^bcoanmhbcjh tm fean &i(yiC32>r>cn Mhm baGUvnhfii
i^ôrybrii 3n<bhoi (qjpyynpsdnp^hn), 3 0 ^ 3 ”b(p3nb *b3ps q)363&3n 9)T>6nfoeh3 bibjpibi (Ramus &
Rosenberg, 1980)
- 333 -
3006660060 u s t f O b ^ a«;an 0 6 6 o n 66 ao
b^h^jii^h^bnb g i(boi636i(5, hi(j $i3mnbi^36i bn6ioipnoi gi^ o h o ixn ip ^ r^ b n b

j>i3q)3hx36i3o. 3hi3ipo J,hn3^m5i3i (6y ic 31536^33611, 0^36360, Rhgnpnb 9 my-
3ih30n bs3b36n, ^(*1503110)0 2)3n0h:>), 6yi03o Ridnh3pn (^hm-qjmoipniBn 9^36>-
h336n g i cgi63hm5i836nb 6363^053!^ fihgopob i 3î 6n y3X^ bib3m6i dpr^h
3j)hd6m5nih36n ihni6 bo6icn0obig8n (q3m^mci6n03hn 3(536^3360) g i higni^nnb
b i 63mi03 <b36mJ03g36nb ghmbiij 50 tbni5g36ni6. (bm503hoin 6yip3u36ih3
n63n6nynib 5i6nugnb 3533 gopnb biioi 363o g i 3iOin 6ib8oh6igob ibnSnQb^ni
b30 3qjhm iC363pm6b (6ib. 6.12). 8mrh36p3po p(p33&ob b3hnnb 838g 36, 3ngh>-
lt)nbnh3630 3gj)m3ih3m6i3n 3ymqjo 0fij3636nb 8mt>33ghi 3nhgi3nhn higni^nnb
6i 3C3^nb ^338 06333b qjm^mbn6oi3'bnb o63o6oh3&ib. j33iiynb 8(536^3360 j>i6>-
013630111606 33Uihn 3m3i^36nbib (dp03hn jihnb 5i3m b6hiqjo <3moim0i5336nbib)
2,i6oi5goi6 d0O3hn 5i6ioi3630m6ob 3mj>b. hmfcmhu 63bn, 8(536^3 3ncn 3qjhrn
3j,hd6m6nih3i J>i6ioi363pm6ob 3m3i^36nbig3oI hi(5 303(1m 3<33j^3hig oy363&b 00-
bo 3nj)336lj 3hn b o b ^ S i bo6icn03b.
pni 3i 6 nl|)i^nb (Sift., 3 ip ip 6 aii, 3g.M!>6m, t»e>3ob 6 i 3oho, 6ib336n 806360 g i
6icn3b36n) 8(536^3360 d0O3h 5i6 io i36 x 'T'6ib 3530i3big 0 ^ 6 36 - nbo6n, dn-
hooiigig, cgm(5mbl!)i6op3h36o ihni6. i8 i3 3 ghmb, 8io»3o(5 33bid03630fii 6 ih 0 mnj9 -
6ib qjmlj)mo63o6oi50i g i qjml5mg3b(jh3j(50i, hmg3bi(5 bb3igibb3i Sn'b^bnb 5i3m
0 ^ 3 9 3 6 ^ i6 63pg36i 3P3jv|)hm636ob ftigiîSi 5ip3o6ob (505080. ib3cn b n ^ i -
(5036b ig&npo ij3cn j>ig ilji6o0o i6 8mj83go b^H^bob 3ohm6360o, J>i6bi53cnh36001:
bo(5bob, yo63ob, &3i03ob, gi8pi836ob, ^ 53 36 0 60301031136360b 6i5036m6ob (360-
6ih3big ibm^nbi g i 30 5^ 30 3836 ^ 36 0 ^ , n6^mjb05X500bi gi 06033^(5036^ 9 36-
0163333680. cgm(5mo63o6o(50i 6ih8moj8636i 8 i 3 o6 , hmyi 3 i(pipn j)i6 itn 3 6 x |Ti6nb
3 ohm6368o qjml5mbo6cn3bob 33mhigo 3hm(53b36 o 33b 8o0gn6ih3m636 bh^po
06*536^31*16001. i8ob ji8m i bSoho 83ig(j>ob gm^mo6!in6o(5ni, i6 qjmj)mbn6oi3'bnb
o6(!)36bo3m6ob gi(533i 63i03nbib g i lbi8oihob 9mb33636nb 33homg3n g i 3 nbj)i6
&i0mb30ob ghmb.
q}fT\yTin53o6 oi50i - go6 i 3 n3hn bljh3b3pn 3 m3p36 i

dpn3ho 5i6icn363pm6ncn ^ 6 0 16 33x0 b ^l^b xn 0m3P363&o 53 ^ 0 3 33 6 3 hoi3 i-
63010b 3m8g336m Q3b^h3j(50x n gi h33ihij503pn 3hm(53b3&ob, h3'boblji36^3pm6nbi
g i gi56o63&ob 3i5ipooib gn6i3n3hn b ^l^b xo bo6 ghm3 ob ib ^ ^ S o (ob. 6ib.
6 .3 ).
q3mli)mbob^38i ll-ob h3ij(jo36ob (536^0 d po3 ho bn6iO»pnb 3 m j3 3 P 3 & n b
bi6yobo 63h^0p0i, bigi(5 bg36i &ih533xn 3hm^3o6xo 3hcn3X36ob (3 i&., D l,
32 -kDa yopi) bhiqjn giSpi. i 3 nb j>i3 n, 3P3ji5hm63&ob ^hi6 b3mh^n o’b p x o b i g i
cgm^mbob^)33i ll-ob 3<33ji!)3hm6i 8(5ohg36i. i b 3 0 i 3ohm &363o, 3 8 o i i3h3 b g i 3 (533 P
(pm6obdn36ig 33odp3&i fiiooi3ipmb ojml5mbobl538ogi6 £>h6n bo6icnpob 3m(5np3-
6 i Q303mh3b(536(5oob 83333m6ooi gi, 5i6bi530ih36n0i, bnoi6mb bibooi. jpmhm-
3 p ib ^ 3 6 nb h3g 3 j(50i 8 y o h g 3 6 i jb^ ^ m q jn Q jdn b (jnj,pnb d jd ^ m d n cn (6ib . 6 . 1 3 ) .
i8 ghmb, ib5mh6iôbi g i NADFH2-ob gib3ih36ooi gn-33mjbog 3ompiJbi6^n6n
h3g3(5°hg36i 3m6m-33mjbog i6oi3hijbi6ô6ig g i cb3ijbi6ô6ig. dpn3ho bo6ioripob
3ohm6363o 3onpijbi6^n6ob 5ihgij 36 i lb 3ijbi6l5o6ig bmh(503pg36i 1^8936063
63010b 5i68i3pm6i3o (6 ib . 6 .14 ) . nb33 3ompijbi6ô6ig i(pg536i, hi(5 bg3(6i h3-
g3j(5ni8o 8m6i6op3 6i3hoi36ob 83833m6ooi, bi58img b6hiq}ig 8o8go6ih3m6b
gi6ipo 5i6icn363pm6obib g i bn663P33o. 5ng33 3hcn gi8(533p 83ji6n'b8b 6ih3mi9-
636b 50n5m0iO)Ob 33î6m0O(b8n.
dpn3hn 5i6io i36x(T'6ob b^h3bob ghmb 5hm3g3&i i&h3boxn gi656igm3i6o
6i3hcn36n, hm83poii(5 333dpniOi jpmhm30ib^36ob 3o5836l536obi gi 8336hi636ob
- 334 -
6sbi 6.13 (yipijjmnQob S3di)ha6j9n JbiEi^mojncnb ^yi^yib {>s9sfHy>336](yi b£)9s. JcmhmScib^nb
jîoij&ob ^^hnmgSn ^pjynngSn pH-nb Jihp6:>bcr\i6 ^hoog (Kao jia m e s a x fo XDW'i’iBjbob
l)hi6bimhlyih3&ncn) jBbnSn pj-jUf'tJbnpi'bi. lyiQ>^mngnb a^S6hi6nb jpb63hn3 3yihp;)&i
qjm^mSjbnb ^hignjfi^n, 3nmQijbi6ljn6r 5ihpinJ3636i 'bjijbi61y>6bg hjpmjb bnb(j38i0o
cnnn&i/mjbnpnhp&x0 &036*o,nrn6H' GSH/GSSG; ibynhj>n6nb %p>3b/g33nphmibjmh6n6nb &p3i,
■V^/DHA^). 3j)35>ihgij96b %Qijbi6|yi6nbi 3nmeijbi6lyi6ig bp;j&i bnb6x33n- *6 dpnjh 6(jnh3
pi ^pQribnhp^tti ^mjbnpi'bncn. PSI, PS 11 - tgm^mbnb^jdi I g i q)f>ynbnb^39} II
(Hager, 1975; 1980). h ^ ijyii dpojh ^sBioijbi'b] pi 3nb&i6 pn}3fib 9Qji5n'b936n nb. Powles (1984),
Demming-Adams & Adams (1992)
Zeaxanthin qh
HC !
Violaxanthin q /V OH
Antheraxanthin
Duration of irradiation [min]
Ebb. 6.14 3nmC4jbi6tjn6nb (VX) {>ihg>j96i i6crohiJbi6rtn6ip (AX) pi 1b3iJbi5ijn6ie (ZX) CUorella
pyrenoidosa-Sn Qiîcn &*5ioi36nb Snhm&^Mn (Hager, 1967)
Cn3np]6 nb 9*^0*636*. * 9 nb 01*305*6 *6*1500360*9 *hb36m6b mjbo936ob h^g^j-

(b;j33hmjbo9-9 ob93 l!>i'b*> 33hmjbo9 *'b36 n, ^^*9*^360).
dpoQhn &*6xn36xcibnb b^h^bntaeSn 3 ^s~n6 *

dQoghn 5*6*oi36x<T<&nb5>*6 cn*3ob g*b*(5*3*9 0(j36*h3 0o0*hoi*3b bb3*g*bb3*
- 335 -
8006660360 u s f n O i ^ a^sp m fn o m aao
b^hbb: qjmcnpnb nhn6ag mhnobvjnho&ib; 9 n6 nbbogi 61600360b 9163331b, jprnhm-

30 ib^ 3&nb 9 mdhim6 :>b biibo 9 o0ii5nr(i jbm3O03 &8 n. qgmoT^bnb gmhbi03 hn 3 bhnb
6ob]b3&nb 9 ^nghm 0336^, 33n93h9 nbobi 9 1 3n3mg3h930n cg3636nb (615., 6063(^13-
63680, 638380301010(^3^6 b^ 81536.^336811 95 ^ iiD b j^ n ) i 390 ob 5 ^ 3 0 3 6 :> -
y330bqj3hn 3b ibhxD&b 51696633(1 cgnc^hnb qj36ji5nib. sb0ig 5180110 <30101036-
8n, 5i6bii3aih36noi ^hm idix 9u36ih3368n, i6onrnjm6n ”fihgn0i3b" 93^1^1gjriQb
91 ibQ33, 5333006361 3hai53ihn cgno^hnb hm08n. 9nj'i0fi 5^61013630^161^ ghmb
j 0 mhm30ib(!)368 n 81^39™ &b nb3om 918153x0 3n5936 ti3&nb iiTi6(j36^hn5ni, hmg>mh-
111511 2)ihm^n6n g:> 03^306(1.
6 . 2 . 1 . 2 DCibh^nnb^Dhfl hiQfW nnOl 6 ^ a n 6 3 D3 Cfl biihObfl
i^9mb€33hmb 51300b 838935 8bnb higni^ni 83ni5i3b 290 68-nb bn5hdnb ^ 10*
("360b ^33001 ahb3& x 30vbhioobqj3h 5 i 9 mbbn336ib, hsy 9 mo(5i 3b 5 hd30 îQ cvii 6
UV-A (315-400 68) 9 1 6i6nom6hn3 - UV-B (280-315 69 ) 5^8mbbn33636b. 6363-
6hn3 3nhm6368n UV-B-b ifob 916190 n6^36bfi3m6i. 9nbn 9m9i^36i 83b^dQ3d3-
0ni 9bm0 Pi9 bli)hivi)rnbqj3hmb m’bm6(ib 8hnb gihp333(ib (bb3i9ibb3i mjbngiSô-
6nbi 91 3191*15363110 6ib8nh6yi06i936nb 38nbd36nb 8 39 35^ ) 8380^333180 (61b.
6 . 15 ). b(p3nb 9m6ngi6 bn9i(p0nb 8i,j36ibi 9 1 53(115^030390 5 1 6 3 9 ^ j>036ibcru>6
3horii9, 30^hinnbq33hn hignitfOi 3hm3mh(5fi30 S9 8 i lj 30 m6 b.
Wavelength [nm]
6^b. 6.15 3Q|>h;>nnbo)3h(i hign>isnnb 96n3363C<n&i 3>33:)ii3hm&i. UVoz 9fn>,c>iiCCrn 3;>^n hipn>
Uni; UVi/2oz higni^nb bc^nb 91^163^ 3, 013 bî^mboflhman m'bm6nb br>0>>311033
J>i6ib33h3bx<ii. 1 3gm53(ib 03663600 b5^m(jnb6n6nb bn5cn3'bnb 91^3300136^; 2 (^jndQnb bijbnb
bSml^&nb 63^ 1536^(1 3033^ 11; 3 bib3nb 6m(x)336nb 33ng3h93Q 3}h3g3&9n 5hmj)m30i'b0nb 6it,bgn;
4 bn0n6g0n O)0i 3m6mne3&nb 3hpig]juni (Cappelletti, 1961; Caldwell, 1977; Robberecht & Caldwell.
1978; Wellmann, 1983)
- 336 -
6‘3606c«S030 ^dSnAO & O
3jh 3g9 o Xil*1innl*3D*in higoijjoob ] 9 > j 3bn 6i6o0o i6bmh6ohg36i

gi jp6,i36nb 9 i(pipo 363h&oxf,'£>nb &i9 (n 06333b 9dn9 ] 9 ^ 0 16 3 6 3 6 ^ &hd30-
^>CCVTi3>6n UV-A 9 mJ3 3936b qjm^mfTijbnQiS^^hip; UV-B, j^hgi qjm^mmjbngiun-
nbi, 06333b ij)h3cri33 q}m(i)mgi'boi63636bI 2,16^5301(136001 - 6nm9396hi6369n.
3(\5hinnb<33hn h1901^0001 5 i 9 m633x n Qi'bni63&3&nb QncDiXDhn 93 ji 6n'b9 n
9gj,rri9ih3ri6b o9 i 9 o, hm9 ig&oen ij3b 3^,53063690 gnbxojng-jhn bng36nb îB-
y33vjib gi p69 -nb oio9 o6 ob 353qjob gf^hobiynib, hiy 06333b 93<33j»J36b ,jhi6b-
jjhmîinnb 3hpi53b9n. gihgi i 9 nbi, 3Q5hioobq)3tin higni^ni o63o6oh36ib 3530136b
3nmpi|ji6 |io6 -9333 mJbngi'bib, hob 93 p36iQib, îh6n 5i6ioi363prri6ob 3ohm6369 n,
jbi5li)mq}n036nb 130500 3<33ijDhi9 33pih ibhxo6b gi 9 (533C <336;)(joib. jQ^hioob-
<33(10 higoiuonb 93932>ip 5i9m6333Q0 pi‘bni636i 930966331 12,(130133 36bn936nb
2.0
1.8
1.6
1.4
1.2
1.0
Duration of U V -B irradiation [h]

6sb. 6.16 Heamatococcus lacustris-ch 'bmmbSmho&Sn jfvrihmqjnpnb qjcomhjbyo^nnb (J3cnc0&0&n'
{>i3m633xn XibhMinbqjohn hspniunnb bifSfrhdpf^n 3mJ93£)&noi (Hagen et al., 1992). Xvjhinnbqahn
hipniijnnor) qjm,ynbn5on3'bnb Snhox^pn n63n&nh]&:> bgods ojm^mbnbijoQi ll-9n (Bomman, 1989)
ij^m & ob u30nQ36i9n O^hmjbogibob ij^03m6nb 9m3i^36i, jjnvj)mjhm0mJbngi(bob

ij^n3m6ob 9^33001361), 33h3gnb ihib3Qbiyh3Q 363112,3(50530 bî^3b9o
ip i^ n b g36igm6nb 153000361), cgmljmbo5ai3‘bnb 36ihnb 9^3300136190 (6ib.
6 . 16) gi "bhgob 3hm|j3b36nb (bo3hisxn bhpob, b16^3(13 3ih5ob 'bhgob) 9 1^ 3 3 -
3i9o.
39i("C3bo 9u36ih336n 330 53^ 0 5x ^ 0 9 hob 93933016001, 30^(11001x33110 hi-
9011500b dpo3ho 9 oii5 oij9 obi g i 35h3gob 633690 j,ibb6o0o <3oi3Pi6(Tiog36nb (6ib.
6 .17) 93933(116001 5ihj>i9 ihoi6 gib3Cn gi'boi63&36obi2,i6. 9 i 9 l533Cn 302,936,53-
60b bo6oi3'bnb 0693(50(1361 69361 3Q^hioobg)3ho higoiuoob 93833(116001 - gi^-
3ohoi3ob 9m9i^36obib, 9 iorio 5(1161536^1(501 obhg36i.
- 337 -
0036660360 b6 6 0 L 3 ^ 3 s?&n8 6 6 3 n 6630
6ib. 6 .17 9(3361(13011 3jh3g3&nb 0n]h

3tXjhinnbq33hn higniunnb i&bmhiujni.
9mbxn X>i^linn'x33hn hignujnnb
9bm0mg 5-10% 1(^6336 33'bmq)nonb
3jh3g36nb ^391 3(13693. 3nbn dnhncoign
6i6npn 3onn6oij836i 33np3h6nbnb 33(13-
93&nb 3ii3mc363n (Caldwell et al., 1983).
3^361(13011 bb3i9ibb3i bibrujm^bcf’i
gi<Tih336nbioi3nb nb. Day et al. (1992).
Day (1993)
6 .2.2 DjbibhDa^CDhfl iWaSOhiiMhnCn 6âm63030n bibhObH

bn^b] gb b n u n so ih n b a i3 h 0 m g n 6 b 3 n i3 h n 0 g j,m 0 ih 3fT t63& n , h m 0 3 0 c n ic n 3 n b iu
g i 9 i b i b n i o i 3& 30 n i 9 m Q 3 i3C 3& n b 9 ie > i0 n 6i< 9 3 b i& i9 n b ig , g i6 i0 n ^nSo^n^hn
]6 3h 5m . bn u b n b 3nhm&;j63 n 9 m d h iP i 6 i d g n ^ h g ^ b i, 0 i$ h m 9 ni032)3-
0 Q69 n 9 ] 0 i 3iQ n 6 i3 h c n 3 6 n oqjhm 9 m d h i 3n bp3&ru>6 g i & n m 0 3 0 6 h i6 3 6 n b Q n S n g jh n
0 h 33i»n h 6 n p g 3 & i. b n (jn 3 39 n & n m 0 3 0 & h i6 3 6 n 3q jh m bn b^n bgo&n:>6 g i 6 n m jn 9 n-
3hn 3hmu3b36 nb ^nj^ng^bib ^bi^nhm^bi ^qjhm 03^n ^^hfcni. yn63nbib jbm3n-
0 3& n 3 y ih , i^ h Q ^ i^ ^ Q 0 g g ,m 0 ih 3m & i9 n 5 i g i g m 6 . 9 y 3 6 ih n b i o i 3 n b 5 ^ 6 3 1 y 3 3 -
0 1^ 3 hignj)i03hn 3 ^ 0 3 6 1 1 bibnijfrnjbgrn Shm^bSn.
bn|ib3 g i bn(jn33 0 mj93 g 3&36 6n3ain3h3&icni (5301^3, 0U36ihnb 'bhgibi g i
bni5fTH5bcnb36ihni6m6ilb3 gi, 33bi&i0nbig, <bp3gi336 101 013 n9 bib3m6nb ih3-
iQb. 0mb33636nb qjibnb bi‘bp3ih'b3 y330i ij^n^hn bibniinijbpn Shmuobnb n5-
li)36bn3m6i 9 n6 n030 i 9 g 3 g ib x n g i 30Jl503iPni- b 3mh36n, i6 3mnj)n0 m3nghxn
0U36ih33&n 03 hi0 3i6nlj i lj363n ih ihni6 bn(jbnbig0 n 0j,hd6m6nih3 g i nly>636
63&nb3n3h ^ S ^ i ^ h i b . CDd^Cjh b(?3±hcns6, hm3 x<H5 gi0ibibnicn3&30ni 8(3361-
hnbi 013 3 nbn l5i 0 i 33Crl iTihj)i6mbicn3nb (ubh. 6.2 gi 6 .3 ), 6 ih0 mnj06 o£>i g >
<bni63636n, hm0036nu i3i0b ($0133636 0 U36ih 39 n. 6 ih i 33X fl bibi’b(p3hm ^3333(1-
i(bDh30n q3ih&p36ngi6 Cb3gi gi j 33gi) &i0 mb30 nb 030oib333ian, 3jh3gnb b^h^j-
(53h36n gi qj36jun3bn nb3 bBhiqjig nhp333i, hm3 3hrTi^m50i'b3i 301033 np333&i.
gi(bni63&i 03nd03&i j,i63ncnihg3b oii6gioii6rn6ncni(3, hnj ij,h3cn33 n6333b bibnu-
rn3b0m q)36jun3&nb 6m6ib6mhm6ngi6 5i3m y3i6ib g i 3jh3gnb gi(P333ib.
- 338 -
Ubhncn 6.2 :jfnnyicm^°Qh3Cn 9o:)6:>h336nb Q
* 0nyiromhjp6rfb83&nb aijbn3^C3hr» iW^DîWhZJC0 h^'bnb^S^cm&i SyQncn j^hfcn 6m9ihi535nbib (^nghnhQ-
6jq) qs QyZnghnhjbx? 9p&™9>h3nd:>9n (Larcher, 1973a; Brock, 1978; Kappen, 1981; Luning, 1984, 1990; Fujiwara & Miura, 1986; Sakai & Larcher, 1987,
Stetter et al., 1990)
- 339 -
gi 0@i66mb
636 3 &fn030 bSÔbTIÔ 9 6 d«S»nfn360
3 0 0 6 ^ 0 0 6 0 ustfObT)^ a ^ a n a ^ o T O ^ a o
in
S 3
n b Qi a
2 ( 8 ^ ®? 9
l£> 10 S *8 *8
»— lo »— r— un s
s>
&o
J -2
c O
: &
■A
T«
JZ>
tO ■?10 ° 3 1
■p, -O S> ^ “?>
a
-8 /n 2 2 c
4?>*_D?J) &r £^
^
2
- 340
5‘3G06<f)030 SAdSnrtObO
Ubhncn 63 b b ^ ib b ji iCn 9i>J0h 'bm6i 9n £jh^[>m3:>6 81536:^3011 qjmoioj&nb oi3h8mh3'bnb^36-

(C > W 3hn iW M ah i^hn b, hmaxyHS 06333b cjmcnQ^bnb 50%-noi 91^01636:^ - LT5Q,
ynBjnbo^nb ihnb 2 baw ii6n 16 9> y i b6nb 3jb3m 'bfxjm , bmpm bn^bnbo^nb - 30 6own6n; a -
33d » i \bn3tln i3nh(>36n; b - LN 2 = 01633190 it>m ,yib = - 196°C ) (Larcher, 1973a;
Kappen, 1981; Bannister & Sm ith, 1983; Losch & Kappen, 1983; Sakai & Larcher, 1987; N obel, 1988;
Larcher et al., 1989; Yoshie, 1989)
attf6s!v)att jecxgn yn63non gi'bnsSo&flb b n ^ n o i es'tensSjinb

I»'bp3h36n bibc^ho&n
^hntin^bn
+ 5-g i6 -2-893 45 - 55
j 3Diiynb 36^3360 + 5-g i6 -3-9^3 45 - 46
9ip ic 9oinb 8(5361113360 -5-916 -15-893 (-20) - 45
|]3&i)h(n3ni|3&n
3bhip063i 63 93hJ6n i636n -8 -g i6 -12-893 50 - 60
qjmcnmc9|i 336n 9 ;)hj6n i63&n -10^ 16 - 15-8g3 a
5iC9o?>n -5-g i6 -14-893 55 - 60
-5-g i6 -ICW 93 (- 15) 58 - 67
C4 6ipib(n3636n -I-9 1 6 -5-893 (-8) 60 - 64
3gi65mb 3hcn6cm 3i 5n 81536^3360 -6-916 -10-893 50 - 55
txnflnohn bihjjy3cn
biBiinhn 'bmpnb 8 ih ip 3 6 3 i6 3 d$\jbrv£rjbo -7-916 -15-3g3 (-25) 46 - 50 ( 55)
3^13330" h3pnj,j3hn 63360 -8-916 -20-9g 3 (-15-916 -30-093) i
33j> &3fij3&n -2 0-g i6 -25-893 45 - 50
q)mcnmp9|j336n 63360 91 63^360 -2S -pi6 -35-8g3 a ~ 50
3bns6n 3i6nip^3&nb 61^16(^36360 -l& £ i6 -20-893 (-30) 47 - 52
fihpn(yii6n 3i 6ol!)il£)6nb 6ieibm 3636n - 10-p i6 -20-8p 3 (-30) 40 - 45
b ^ n b 9ih ij3cm36360 -30-916 L N 2-8e3 a b 60 - 65
3bpmojn(536n -10-g i6 -20-8g3
- 10-g i6 -25-8p3 ( 42 ) 55 - 62
6ijCnb 8(5361113360 -5-p ^6 -12-893 38 - 44
3m0mnm)n9h3cn 2,308(1360 - 10-916 -40-893 46 - 48
Un^taScnhrefin h3f>nri636n
9ihig963i63 6n&3m363611 -40-e i6 -90-893 44 - 50
imhpp^hn ajmcnmcS^jGn b33&n -3O-916 LN2-893 b 42 - 45
ihj^nyibi 91 8013606 333s 63^^3611 -3O-916 -70-9^3 48 - 54
ihj^nyibi 91 3i(rii(^ 8oinb bipibrn363611 -30-916 LNg-893 b 44 - 54
- 341 -
9(306663360 L663U71C? 9âr)8663n6630
6 .2 .2 .1 3 jW)hD9 ^C 3 hn i*»D0 3 Dh>&3 h3 e>n QDe^flBIftO Q) bfllim iibCnb
iW aSO toiM hXfl b i <bC,3 h 0 6 fl
9(536ihobioi3ob 36103(1016^ gi 63961^300 ,1,3033(11^3(1360 ( 5 °C -

916 2O-25°C-093) ihbj&m&b 9bm0m0 6y0ob bnph0 ] 3 n gi 6010150b (?h0 i 033-
6369n. 66303010b ‘bQQiSnh'b^ 91 01639 6yo3&0o S^hy^mbb QCP^-pb-
93(119 gi (3i3^CifTih x ri lbm6nb 519(112,036001) b3<bfii63hig 9o6o9io3h g i 0ijbn8iQ3h
0 6 0 9 3 6 3 0 ^ 3 ^ 9 mhob, (n9i(5 93 od03&i 9 ni(p6 nmb ^(5(11(5600601306 b i 9 n3
bi<bp3h36b.
9(5361(13011 lb6n£SvJ3&nb 519163(1361 69361 gngn (11111936(116001 0oii6avj030o
363(15006 gi, 16 133 phmb, 6ooi6m6 ihibij)0ihnbn 51666330b 930016333180. 6363-
6hn3 3ohm&369o 363(15006 396n936x<ii3i636 Byihmb 6ih9mgj,36b 0 bnb higni-
(501. 5^(191 i 9 nbi, bncn6mb 9 mî6 i 933990101 6 2>^36y33C»Cn3 9 tiidhi3 3i3hnb 3 i-
6366. 3$jhnb ysjc& D d±(p$On tjjd S jh ^ jh jd n giq}oj6oh36xo:> 3.6. 51(191^3606
5 i 6 x 3 ^ ct)3 - 6(1909(11301 iqjhn ^n, o6gm3ai9o, 93jbnî9n g i 2Hpnq;jmh6ni9n. i j
5 i (6m0o0o i6bmc3(j3hn 0ifjo030o 39(106 57-58 ®C. 603930106 ^giSo h o b 2 3 %-ob
3i3hnb (i^ ^ h i^ h n b bi03i0m 9ij6o9393&o 40 °C-nb ‘63300111 [9 5 ], hi(5 60 3613b
n0 ib, hm9 dgo3ho (11901(5006 3ohm&369o ib30i 1950936311 9(5361(106 63833(11,2)3-
hi 33hnmgxie 3&e^ 1(^633936 50 °C gi 83^6. 6019151161 g i 6yoob 9i(ni0n ,538-
33(11,13(1360 912,1390(1363901 3Xi^6n'b9an6. (5(11(56190 mh5i6o'b936ooi 9 1616 9363-
Cn y33Ci<t>3 13^300 195003601 53o(b3h36n, hm00369o(5 6y0ob j>3933hi03 hi 9 2 -9 5 °C
HP6336. 3X i^ 63 ^n^ bmb0fn33b, 9n6nbj330i 9i50obi 91 (36300 6y03&nb 5 13-
036001, 6oigi5ob ,1,3933111^3(11 70 °C ip633b.
9U36ihob 5igib3h36ib b9 ohig ig5ogo y)36 poi, 9 bob hig0H5006ig9n 3jb3m-
6oh3&x 9* 3jihm 3i6oî^369n. 6i9i036‘b3, j3o96ih369o 9 1 160319^,06 ib0mb
6oigi5ob b3gi3oho 516612,301(136001 d0O3hig b3hg36i. l6m9o3h bih^y309n 60191-
50b <b3gi3ohob (2,3933(11,1,3(1360 60 -70 °C 1^63336, bm0m 39166^90 - 80 °C .
0bob (11901(50001 51001633300 510163(1361 5hd30g36i 9bm9mg hi0g36o03
buoinb 5 i 60 i 30 rri6 i 9 o. 16301 3ohm6369o 95361(106 (11(1516(113606 519163(1361 6ih-
9(11195366 bij)0img 9d0i3h 91^30(10)316. 2,1(150 (2,3033(11,1)3(1300 9 o9 (ii|330nb 0jm63
03(11010360 g i 931300360 ig3O0ig 1011616(13636 0113060 gi 51(133^13(53390 ^13(106
,53933(11^3(1366 9 1 i 9 o^,m9 , 9 ioio 5igib3h36i 9bm0mg (119936093 6301b 5hd30-
9361. 0139(31, 632,3036,53606, "6(11503(1010 91(11530(113636061 9 1 0603606 6019150b
“630130110116 i60(iib 90361(13 6160036b, 9bmbi3 (11(1516(11366 9 1 0336306 y30b
033d0oioi ioî6m6 40 ® C ^3933(11^,3(11, 6m0m 9(50(13 66001 - 50 "C . i 9 ghmb
obo6o o9ymg336oi6 j,3933hi^,3ho6 o9 031(1503690, hm90360o|3 6i6n(5m(5b0m 3 hm-
(536360 3jb^h39i 03 hig gi^3ohcn30oi g i 03î03h <b(j'3ihoii6 i60m 6n (616. 6.18
g i 6 .19). 9o363gi3ig i 9 obi, 6(1161(539360 9(5361(13011 600163110 63j)hm'bob 9361636
9 ioio &3636h030 5i3h|3303&ob 195091b oiootJ9 ob ih ihb36m6b. 3b, 196101, o9 ooi
iobb636i, hm9 iji0 g 3 i 9 9m3036ib ih 3ji33mgi bicni6igm y3hig(i>36i - g i 9 >
33h3&30O 0 i 9 i 9 ^,ioi536x n bi63oi3&ob 9m3m33&i 93b id 036xo i 9bm0mg 015361(106
(J3033hi^>3hnb 33h0 i 636 ^ ,x n yi>6^hm0o6 3nhm6360o.
60(5606 3(101-3(101 516612,301(136001 bi 9 o9 5i0m30n636ib 6ih9mig536b b*6d-
hjbn. j,y336nbi g i 63fij6ih3&nb (915., Ri3ihi036ob) bi6dho6 ghm6 30oiihg36i
5i66ii,30ih36ooi 9i(pi0O ^,3933hi^,3h36o gi, hm5mh(5 6360, 6i053hg36i aiooij9 nb
001300 0)5361(13x0 biqjiho. 6i6diho o6y3&i j,yob 16 63fij6ihob 61033600 (53(5bpnb
51636001, hm 9xnl3 339936 519190b J33^y3'b3 g i 6m0mb, 69 x o ^ ^ ,36 0 6 1 91
00^36360b 93933(116001 - 3ih336<b3. ^yob bi6diho 3(1(5309361 1-6 2A60V1 6ofi-
- 342 -
6 3 6 0 6 6 0 3 0 bG6 0 b3*=?0 ^ d 6 D 60 6 0
HLPh
20 30 40 50 60 °C 70
6sb. 6.18 b^Snb 6iQibm3i6 8(3361(133630 ji3933hi^3hnb bljh3bo gi J>igib3h3&nbig8n h3‘bnb£)6(E}-

pmbi. 153833(11^3(1300 8(13936(1 ibsbi3b g(nnb ysxÔ U^X 33hori@3o 6019150b ^39130(10916 ^38-
3jhi^3hnb 5hipn36^b. 63(1^0(^316(1 bilxi: 3i3hnbi 91 6nigi5nb bi93iom 8ijbn3io3ho 153^3111,53-
hi; 6y33^n(yi biVi: 3i6nî^9n uphn^bxn 381^00^ 143833(1103(11; bujhoboQhn bi'bn: 3|j36ihnb
mh5i6m3&ob 8ijbn8i03hn ^383^1^3(11; 813(1 bi'bn: 3i536ihob bb3ipibb3i 6i6nonb boybobigSo
h]'bnbi36(j3pm6i; HLPh: C3 91 C4 8u36ih336ob qjm^mbn6cn3'bnbicn3nb bn^bnb <be'3ihn (Larcher et
al., 1989)
Heat resistance Heat stress

T1VTL100
A: 46/52
G: 50/54
C: 48/52
48/52
48/52
56/58
52/54
52/56 61b. 6.19 Sempervivummontanum-dn boybob
b^h3bn (815361(13 n‘bh936i 51(1519 5161013630 91
jihnb5i6 91UX 3i(riCaCT'nlj Ii6nî,j8n) 91
biq)b3o8n bn(jbnbip8o 8nbn h3<bnb^36^3Q(n6i.
LT„ 8ijbn8ip3hn ^3833(11^3(1(^1980 1913,50(1361;
LT100 >J3933hii53hi, hmSx’bXÔ 9b<^»e
UiCi33Cn 0^1x0(136300 35(139360(^1 013
5191(16361 (Larcher & Wagner, 1983)
- 343 -
8006AAD360 U&rnObT)^ 8^50966306630
jihocn gi ^305jhi^jhi 3ih3ob q)ih&0360n i(j<633b 500 -700°C [251 ]. bi3i63&ob,

gi 6]fiJ6ih36nboi3nb pxfebibniOTjbQcn bSohn biBdh^nbib
hi ^qjhm gi6ipnb (200 -400 °C). 99 hip U3(jb0n jihob bnRjihnoi
3^13X 60 ^ (B^b- 6.20).
eatacn A303Dhi|S3h36n ihnb g3pi9n6nb bncnb^hn h^gnQnb îhymqjnoin 6i0i6-
bnb 93Q3&0. biBîpm g i 9 i(pip jom&hicgox 2 ^ 30 36 ^ 3, 9 bnb gibipn ib n d ^ n b
gi p i 9 nb 5 i 9 mbbn33&nb qjibob j>ibi6j>hd0o336ob 5 i 9 m, hsgnsunnb dsQsdbn 'bi 9 -
aiih9 n îhymojncnni. i 9 ghmb ig&o0o ij3b Si^hnb 9 ib ] 6 nb dpnQh £^150336^ g i
gij,hm336ib. Svjhnb (5030 0ib3&n (jnhj)30 ibnrib 9o933n6ncn ^ ig in ^ S^ b i ^tghm gi
6iC J)s63g36<b3(3. i 9 o(jm9 bn^n33b hmj,mh|5 Qnjjic^hn, ^ 3 3 3 êm&ifvjhn bibnicnn
,•16 660
6bb. 6.20 (jaQSjh^hnb 'bhpi pi ^jQSjhi^^hnb fchipn^B^ bi3i6i9ri bi6dhnb ghmb (Vareschi,
1962)
ifob gi 6ih9mnJ9636i 3i]hnb 15030 9 ibo6 nb 9 mî6 ob (ig33j(50i) biSîO^ftnoni^.

i 9 m633x r‘i 3m0 ih x n Siohnb 9 ib ] 6 ob 3 ;j0 m^hocn.
839^36001 9(5nh3 qjihoimbb] 15030 3i]hob 9ib3&o 6ih 9 rn9 m6 ob ig&op36ngi6
(9 ip i 09cni, 3 o93 x n b h ^ o ) b3m636ob 939331*16001 îgiig^npgQiii gi60m6obj)36,
bigi\5 obo6o 5hm3g36i g i 3qjhm 016000 3i3hob 9 ib^ 6 ob ^338 3;)l533i (o633hboi).
SiQhob 9h336nb 63333hii!»DhX Qiymqjib igj>opo ij3b 8 oihnico 9 i(J)Dh gm63‘b3u,
hiy 5 i 9 m633x ni 6oigig,obi gi 9(536^3300 biogihob <b3gi3nh36ob 9o3h bocndnb
5i9mbb0336noi g i 3i3hnb ^15036300 0336360b gi6pi g i^ n o t .
- 344 -
57)6066030 16601^0 ^6d6D6060
A ir minimum
6jb. 6.21 5 hipo36lj 3{>n bb3igibb 3i tabnumubcm cgnhBnb 3 |j36ih 33d3 n (Shmueli,
1960; Hofmann (jn$. Burckhardt (1963) 6063931101)
6ib. 6£2 gsftiQn gi yn633&nb &*6i6ne3&i 9391606^ 3. A 3yo63m bm6i; B yn5-

33&n jJn'bmg^hig 1(^6336 6n63b 10°C; C (5030 'bidonhob 6jm63 h3Mim63&n biS^^^m biScnhrtb
15365^ ^ 31136001: -10-gi6 -40^C-993; D biSîcm 6(Yi3hn 60606360 •40®C-'b3 j333mcr>; E 3mpih3(yi
yn63pn gi S3h6i635(j3Cn yn63i; 83)1 3i 3n bibn: -30®C 6o6o8i(T)hn o'bmoi3h8i; 6i(jo 536iyihn bi'bn:
+5QC 3n6o9iC3hn n‘bmcn3h3i. i(ri6n36x bm63i>3o &i3h(j3C3&X 9|}36ih30ii yn63ii>i3dp3m{)i
bb3igibb3i i (Larcher & Bauer, 1981)
- 345 -
3036663060 8^ & n 8660 n 6630
Ijbhncn 6.4 ]gsi>6rob C a 9u35sh33bnb (Atriplex sabulosa, Tidestromia oblongifolia) qjmoieD&'bD

9s(ns(xi (jyWjhsvj^hnb (®C) 10 6owii6n ‘bjapiJSjejfenb SjgDfê gj]6ijnrn6nh36nb 1 0 % -ai6n
n63n6nhQ6i (Berry & Raison, 1981)
Atriplex sabulosa Tidestromia oblongifolia
ajmoicnb 0336^50060
63^m-qjrTyjmbn6on3'bn 43 51
0n^mjm6ghnxn b]6onJ3i 50 55
930&hi63&nb 6ib33hig5i9^)ihmdi 52 56
JCf’imSpsbinb qgSjisnibo
gjm^mbnb^39i 1 > 55 > 55
q3m^nbnbj)39i II 42 49
RuBP îhdmjbnQibi 49 56
PEP ishbrrjbnci'bi 46 54
3-PGA yi6ib i 51 51
ip ^ n c ^ ^n6ib i 47 49
cgmtxrfm33;)brn‘b:> 44 52
6 .2 .2 .2 b fllib O
gih(p333360 gi git>ni636nb 3hmu3bob 8n3go6ih3m6i

Qbipipn U30Oi6 6om8386hi636ob cgn'bn^h-jnQn^h Qg^mQih^-
m6ib gi (jriQob 8m0333Q36ob b ^h^^hib. 3b Uhm^bo 333Jb33igni. bnubobigSo
g)i 6 b i i 3 cnh3 6 n o i 0 5 h d 6 m 6 o ih 3 i (j o 0 i 3 m o g 3 6 o b 0 3 0 6 h i 63 &o. b 6 m h 3 g i 0 n ^ m 0 , ,336 -
33hi^3hxn 9iboi636ob 3hcn-3hcnn 3oh330o 8^33636300^ qjm^mboBo^bob Qihp-
333^. oib3gi3nh33Qig ob(”3g36i qjm^mbob&jSi ll-nb qj36juoi, 838935 30 3ncmh-
93&i 6 ib 9 oh6 igob 83^>6m0n'b9nb Qih(j>33336n (ubh. 6 .4 ). jQmhm3psbl53&nb g>
boi63&ib 3m3y336i cgri^piboeo^bob g33h3boi Qi bmpmb - 33h3gob brujmijbpn-
b36ihoi6ni&nb 9^3300136^. hig5i6 cgm^mbnb^Qi Il-boi3ob gi3ibiboioi3630n:>
cgmli)mn63n6 nuoi, bnijbobi gi 3bob 0i(j>:>0o higoiôob 30106060(136300 801^8303-
6 i bi3b3dncn h3 i 03 hoi. ^ h n in ^ n 6i0ibm3i6o 3i(i3mb636 ob qjmcn0360 n 4 2 "C-
gi6 nBy^&b bn^b3<b3 gi0m3Og3&30O 03^^1063060150* bo663030o qjmcn03&nb q>
'bn>636ib ig5o0o ij3b 48 °C-ob ^33001 [ 149 ]. 013 y330i<33h i8ib 3 8 ^ 3 6 ^ 30933
3honn b^h3bxn qjij^mhn (8^5., 53i0 3i), 0m^)mbn6cn3'bnb g33h3bnnb 6036360
oru>3b 063636 3333 30 °C-‘b3.
bnijbnb i8(ji6rri&ob 3(10,1)03300 bibp3(i36ob 9^9536^ 83bid03&30Oi in vivo jpm-
hmqjo0ob qj03 mhm33 (jh3cn 5ibm83ob 83833016001 (6ib. 6 .2 3 ). hmg3biu 0hi3i0n
oi3h3m0 i 6 n03ho 36^080 b^mpmmg 5i0mgob 86ym6hogi6, 6om8386hi636nb haij-
pn b^h^j^hi oh(j>333i, 6n30in3h3&i0ii i0mhfi33ooin ^hiSbSmhô g i So^mjmBgho-
300 b36oij3i 6yg36i g i 33(1390 OQ>3336i.
5igihfi36ob 36iho bo(jbnb b^^bnb 3nhm&3&dn

0i(T>i0 li)3833hi^)3h36b 8(336^3360 3&hd30i6 boubob ^3330001 (33hi6oh36ooi),
336501136001 g i 3hmj)(Ti30i'b8ob 8g5higm6ob i0i(rC3ôoi.
cbm503hoio 8 5 3 6 ^ 3 boubob b i 3 o3 0 m j8 3 g 3 6 ib 0 0 30 9 ^ 6 o(jo036b 0 b o b dgo3ho
h igni^ oob 8m J839 36ob 33h6o03&ob 6 yi0 m 6 o o i. b i6dhob 6 o 6 i i p 3 g 3 5 8^1^839366
booi6mb obm gi^m ho - J3hjn ( 0 i 5 „ Sequoia-b i g i Sequoiadendron-ck b j3 0 o 6rn^-
- 346 -
67)6066030 U66flli'as=:0 36d6r)6060
6^b. 6.23 bnubnbsgSn bb3i@ibb3i hj'bnl)-

ii!)6ilX,T* nl> 9Jm63 3(335ihoo>i jpmhmojno-
nb qpynhjbujf^jni (j^SSjhi^^hnb
bhpiboii6 3hou>g (Schreiber & Berry, 1977).
8^64(533360 ^n Sn ix0 Q* 'bmSnjhn
bihijypnb 3uo6:>hoou>oi3nb nb. Smillie &
Nott (1979)
in3i6n ShoQbn; 0hi3iQO b] 0i536ihnb bmhicni6n, 5ii>pih336x n Johjn 5i6bii3cv

h^dncn b30 nihngx e i ihngx ’bmSQftSn), 5i6ibQ3dnb i3nh^6nb bj^pn bi0 mb30n
(6i0ibm3i6 91336^33680) gi 0n6nb;J330i nh5i6ri36nb (&mcj336n, ^bjhobn) 6ih-
3mj96 i. bnbp&hn 3haiQ3&6ni63&n, hr^mhnijii 3iQ036 n g i b^6^163611
(Xanthorrhoea) bi6dhnb 03g35ig Sbmpmg n^hobQbniS. 83bidc3630ni, bi6dhnb5i6
gHJ3nb bi8oi03&i ihnb bnpn(5n30nb Qipipn 330153x^ 61 0ioin jjhog^nb $39-
0363n. ij3g i6 5i3m0gn6ih3, 3hoip3&6ni6n qj^hmqjn^bnbioioob p^hmb 6 ^ 3 5 1-
6n gi063hm6i ^qjhm 6ij)p36ig bi5 n3 ni, 3rtgh3 SQhnqQhnopn J>i0&n30nb 8hnb 0 jm-
63 nhQ366ni636nbicn3nb. qjmoi03&ob îgibohQ&i oii3ngi6 10(500361 '■lhi6b3ohi(50-
nb 5181(50336300 3<33jl;>nb biSîoobnoi. bi 6 i 0 6y0ob 9ihi50 bi*)3 ihoboi, x ^
6mb 015361)13011 q)moi036n 4 -6 °C-ncn (‘601503)101 030cnb333i 8 o 10 - 15 °C-no>) 3cghm
fchnpoi, 3ngh3 5ih30m0i5330o 3i3ho; iboom ‘bm^&nb 0n("3&nb 51)1393, nbn6n
DD^3X i e gi'bni6go6mg636 (61b. 6 .24 ).
3hmli,m3c i <b3 nb boubnbig9n ipiogfefiiynte ihnb dgo3ho bôuoojooho 01130-
b3&3h36i. 3)1010 g i 00133 &3ihnb ibpm0m6iot3bi33 bibombQbnb 5i9mypiqji 83-
bidQ363pni b6m h x 1 8 oi3ob3&ob b iq o d 3x't>D- 3hcnn 0y36ihnb bt>3igibb3i mh-
J>i6m8ni3 y i bo(5bnbig0n ^mpohiB^rabi 516 1^ 313 36 3 0 0 1 (nb. 61b. 6.18 g i 61b.
6.19). h3<bnb^36l£X'm&nb gi8ibibnioi363po 5i6bb3i336i6n, hm0036n(5 83060361
91331*138ohmoi 81536^3011 5ih8m8m6obi g i 5 1 3 ) 1 1 5 x 3 * ^ 19500360b 01130-
b363h3&ib, fii8myipobg636 33rr»03(5onb 3hmu3b8o. 15030 h35om636nb (i£)6ghi,
3i(pi00oni) 8(536^3360 0ioio <336^503)10 0g5m8ih3m6ob (3Qi'b0 ob giboi63&i, hm-
5fnh(5 363h53ôj)Xn iho’bnbob 608160 g i (50iyn-fim6Rbnb g361^3)10)1361; 61b. 6 .25)
g i CDiWD^n bi'b(j>3h36nb (ob. ybh. 6.2 g i ybh. 6 .3) 0obog3ooi oqjhro 95hd6m-
601)13 ihni6, 3ogh3 <bm0n3hn bih^ypnb qjpmhob 6ih0m0ig536o36n, 3b 331-
6ibi)63c6n y i jqjhm 05hd6m6oiho6o - 3ogho 0 i i!>hm3oi)36nb 815361-
h33&o. ij3 g i6 5i0m0go6ih3, 0ih(53pm363&o, obQ3&o g i C4-bn6ghm9ob 0jm6o
61536^3360 bnybnbigan 5i6bi2)3cnh36001 l>)ri03hi6(!)XO6n 36 9 1 093636.
boubnb 3033^° gi0fnj)ng363QOi Qobn 8mj0x36nb bi65hdQn3Pi6i<b3; 3.0. ob
833bi6i036i gm'bnb jp 6m6b [15]: 8xiho6noi b^b^n gi bi65hdQ030 boybo ob333
gi0ilboi6363COi, hf^mhy - dpoohn g i bi 60 miC3- 10 0 ^ 0 , 03oii6b036ob biqj^d-
3X ^*3 bo(5bnbig0n h3'bobli3 6 ^ x rn^r'*J 8^1611530360 0m3yi3Oi 0i536ih3<b3 boybnb
- 347 -
8006669950 I 6 fn0 b'3c5 8 eJ&0 9 6 fn90&690
Time of day
6ib. 6.24 3gi65mb Snhmbjban j^n^bnb j)hi6b3nhiuni'bo n3 I33hnmg8n, bi6i6 8(5363113013

Byonoi SmSihi^jbi 6mh33Q3hn3. 03^10^3611, hm33Qoii ^hi6b3nhi(5nnb n6,j36bn3m6i 3i(pi(yii, Qrrpy
93339 0i(ni0n n6bm0Si5nnbi, ihn>5 53^036(101 3qjhm fchncn, 3ngh3 53h33m3(533pn Sijhn. 013
qjmenpnb y366n 539311^363 gi 33bi6i0nbig, ^hi6b3nhioni(5 d(yi3h 93315383 6s 3o5yg36i, cgmoicnb
(j3033hiij3hi b6h3qj3g n6y3&b “bhgib gi b3p 8493 53g33£>h63&b 53h38tn3(5330n i>3hnb ,$3333h3vy
3hib. Supipn ijhi6b3nhiynnb 3jm63 8(536^33611 353361136 (5639 5i 6nî,j36b Sbmpmg n3
8380163333311, hm(5i 8301 Byonb n6(j36bn3hn imhoiJc36nb b333303&3 5336611301 (Lange, 1959)
Heat inhibition of protoplasmic streaming
6ab. 6.25 ipjdnhn 8(536^3013 bnybnbsgSn Ii3,bnb(j36<j30fn6ib3 g> 3iOi 6>3hi5X3&il> Smhnb. fchrxyi
li&n^j&nb 8(536^3360, hmj,mh(5 631*1, bn^bnbigSn 63503611 h3<bnb|)36^30(n6noi 5331*10^33036 93
Snhnjnoi (Kislyuk et al., 1977). b]6oij3nboi3nb ihn^nj30fi ^3853(13^3113603: 35-40®C (3hiyiyib
9(5363h30i30i3fib), 40-45®C (‘bmSnjhn b3h^ycrb 6303bm336(1 3(5363h336nbai3nb) 93 53°C-3p3 (3936-
6mb 3(5363h30i30i3nb) (Semikhatova et al., 1992)
6ib33ihbiicnni6n 5330361^ 339353606 606393001. 3hoibsicnni5 n ,b39mj03936nb

939oib333i9n 9s'bn5636nb gs6y36nb 'bQîhn 1-2 °C-ooi 9360b 9^06339^.
ihb36m6b boubob:>g0o hjbob1j3 6 ln c fr|^n' > bb9 n ^n3n:
bn(jbnbigdn dfthdBmbnshj bsbjm bibn. sd 363qj0o 3hcnn:>6g3&;> y330i nb bib-
3m6i, hrn93Cnl5 ‘60*6936:. 32,33 30 -40 ®C (3i093h3b 036016333^90, 45 % -b 3 ). 3b]-
- 348 -
6-36066030 b&6 0 bT)c?0 36:1606060
6ns: ^ i h n n ^ x 0 t>C'3ob 6yiQ9y36ihp36n g:> 6y:>03n fisdnhx*! (j^hm-qjmoioobBo

8536:^3360, 6ycnoi 6^3 3h36300 00^6360 (hoi9036o(5 53 m83 hb0 8g50i8:>h30i6390
^^338^36360:0 gs b0xocnnb 63^03010101136 85363113013 333 ,^ 0 16 3. 95363113013
bb3ipibb3i 5^0101536^0 &3j£)ho36o g> 3nh3b36n 351130133 ‘bn36g36o36 33gih3-
6ncn 93630 ^3033hi(i)3h36'b3 (6^5., 3m9oQ3hob ^>)6oi6ob 3nh3bn 0(^3336^ 40-45°C-
b3). 3b bib30)636n 533115303630(1 ihns6 nb3cn 3^60^^3600, bigi(j 3h 5^60500-
36 dQ03h 5^gib3h36ib g> 3<33toho ti)h:>6b30h3500b 039330160(71 333d0O3Oi 330-
63)1636016 bb33onb ^3333hi^3hi 6oih3iQ3h q)3li503 &3 n.
djgshjdnoi bn/jbnb $d&>6n jjtjihnm ^jbn. &bnb6n gi 89h30n 3i 6 nî6 p6 nb 853-
6sh33&o, hoijoihu 63bo, b ^ 0ôig bo5b330^3636o3. nbn6o 3dc3&36 50 -60 ”C 113833-
hi^hob 6ib33ihbiiCnoi6 3oij03g36sb. 85363(13013 3b(j>iogn<33h36i5oh36xo 33h3-
P36oboi3ob 60 -70 °C 53g 3X 3b33d <b(j'33ho3.
bnybjfr^ddcj 3hm^hnm^jdn. ^0150311010 oi3h8mcgo03ho Shmjphnm^n o^>6b
dQ H 3h 3ip b Q O j) 3 0 3 3 h i ^ 3 h 3 6 n b 5330363b. 5 3 (^ 3 1 1 3 6 (1 6 |jb 3 0 6yo363n, b3g35 ^>38-
^3hî5Dhi ip633b 75 °C, b3bgg36o36 5r>i6m6i^3hn36n. bb33 6ij^3ho36o 11^.636
9 0 ^ C - 3 g 3 j, 3 8 33h s ^ 3h b b , bm Q m p h 3 i 'b ( T 3 3 ^ n b 3 o 3 3 h c n 3h 8 m q j o 0 3 h o 3 ^ 3 6 3 ^ 3 1 1 0 3 6 0
(835..Pyrobaculum, Pyrococcus, Pyrodictium ) ^ 3 6 0 16 3 6 110 °C
[235 ], y330b 3b 0^53601380 3[?£)h3O0O3 536b3i30ih3&ooi Sg^hbgo 33h3g30O
8 3 8 6 ( 1 3 6 0 0 1 , 63103060b 8 9 3 3 3 6 0 0 1 3 g3 1 3 0 0 3 6 0 0 1 .
boybnbig0o 8 35336 3 bBhiogbg 3n3gn6ih3m6b. 33^363 bg3&3
gC336ob 5360330016300, <bm5j3h 3gjhoi 6^3(135. (jb x gp3360o bojjbobig0o h3lbnb-
vj)36lj)XfT|î 33 >g(T'nboi3nb 3cghm 3 i(pi0 Oi, 3ngh3 gooncn. 1303 30 o6 g9 o 538016(1-
01016b 8o0go6ih3m6b h30g36o03 gpob 5368330016300. 538016(101016116 ghoib 3hb3-
&xo (j3833hblj3h36o 3693 093636 o8g36bg 830300, hoi8 83010 80^839363 33I13-
gob 3hnlj)oi305<b8oboi3ob 6^(136300 oyoib. b8303onnb 85363113360b 303l|>3boi6i9o
i 3 80130363b sg5n0o ij3b 35 ^C-ob ^33001, 93(153001363600 - 38 -40 °C, boi0oi
*J33X36\33&o o6hoioi6 o36 p 30ob 03(^300 ^ 3033hij)3h36nb ghoib (63b. 6 .26 ).
boi5bnb5i6 3033^13^39 9385330 3hcr>-3hoi 6383303639 8(16633003 bnybnb dm^nb
ynQjbn (HSPs; 1300360, (101830013 801031303(10 60163 03hy3Oi6b 15-110 kDa 0 oihob;
30oi33h3big: HSP90 , HSP 70, HSP60 , HSP 20 ) [260 ]. obo6o b6h3cg3g ioigohg36o36
3jh3gnb 6ohcn30n, bo6oi3'bohg36oi6 130^01^01000, hob 330g353(3 5(1013936036 jgoi-
hm303blj360n g i 0nôijoi6ghn369n [ 160 ]. 3b 1500360 0633336 Jhm3 bô6 nb b^h^J-
6ib. tS b au36bh33&nb bnyb-

nbigSn g i 3fnyiQ;)&xD&> • NH
'fcOyW&yy'' 9u36ihnb bnijbnbipdn
0h>6|imSii (p(nnb bbdîpm
30°C; 9jn^(Tii9nb biSîpni - 20®C); H3
h b h g i lijcD^hi^jhnb
3nj93g36nei6 3 gpnb 339935 (50/40°C);
Hi j h3'bnbj)36(iX(Ti6nb 3^336363011 21
40 50 60 70
9Cnb 339g36 (15033 (50/40®C)
(Nobel, 1988) Stress temperature [°C]
3 49 -
9036663050 UStfOUTJ^ 3 ^ 2 0 3 6 6 3 0 6 6 3 0
^^hoftnbi g i 639611*63606 6li*6opn'b*(50*b 0* b ^ b 3 6 1 ^ 3 6 h^Sihiynx Shm^Q-

b^6b. 6 0 (56 0 6 b ^ h ^ b n b 0 * 6 0 1* 3 ( 1 3 6 0 6 930036, h *9p36o33 b * * o in b 5* 69* 30^ 6 * 9 (1,
n b n 6n n b 3 3 jh 3 6 n * 6 .
0h*3*c 9(j36ih39n bo{jbo6 *p0 n 8g5h*gm6* (h3'bo6li)36tj3Cfn6*) nlS3CD®*:> 60nb
5 * 68 * 30m6 *9 n (6*b. 6 .27 ). 3b 3hr*H53bo g*i*30nh3630n* 5*63(101*113606 hnlj 9 cni6
g* 5*h38mb ^SS^h i^h n b 153000363601*6. dnhnoi*gn ‘bhgnb qj*b*9 n y 330 *
9(536sh3 60(5606*960 dgn3h 95hd6m6n*h3*. 6<u6*hn 6y036nb* g* bc^nb 6 y*o 9 -
I536*h336nb 60(5606*960 h3‘bnblj36|i3(V'i6* 933b*6*936* Bypob lj3933hij,3h*b;
530*6 <b*cgb3p3n 9 *oin h3'bnb^36lj x f,'î 39*pc3bn*, bmpm b* 9 oi*h9 n - y s x ^ D
6(50113. 60n3ho 9 noi jqjhm qphcnm*, h*y 3qghm gngn* 6ypn6 ^3633(1*-
^3(13606 5*6663*336* b*qjb3pb* g* b* 9 oi*h9 n. b0303«nnb 9(536ih336nb 6*6d09n
A-Type Rhizoclonkjm^/ °c
46
/ \ 44
42
/ X 40
\ /
38
!
S -Type! 58 *
56 §
\ 54 S
:
52 o
w 50 8
W-Type Abies 52 §
o>
50 o
!P ) “8 i
i / ' new shoots 46 x
r 44
W/S-Tyf)e ,___ _ Artemisia . 54

52
50
■ '7 ^ A / - 48
\ / 46
Spring Summer Autumn Winter J F M A M J J A S O N D
Bib. 6.27 bb3igibb3i ^nSnb 9(j;)6ah;joi> brnjbnbipdn h^nb^jS^Qmdnl) b3'bm6]hn pn6i3nî (Lange.
1967; Larcher, 1973a; Kappen, 1981)
bn(5bnb*g9n h3‘bnb(j 36 (j>xrn&i n9*(j)36b b*qjb309n; *ho*6 nb30n36nu, hm930 on* I13-
<bnb(i)36l!)3Qfn6* 9*^3pfn6b b* 9 oihnb 0nb33636nb 33hnmg9n. 3b 3impnj)n3h*g 3>
h*gmjbxn 9n9gn6*h3m6nb ^n^n 5*63001*113606 3hro(5360n gm6n6*5^3ho*. g*6m-
pmb, *hb36m&36 6(536*113360, hm93poi* bn(5bnb*g6n h3bob,j36^30016* 6gnb
5 * 66 *30r>i6 *9 n *h n(5XD&^>-
b*6dhnb 836035 5*g*hR36* p*6myi93&xn;> *b*gn ypmh^bob *9 myhob 36*^63,
9 x ^ 3^no1 J>*h5*0 0 * b X ° 6*b*C3hn J>3ohlj36ob *hb36m6*b3 0* 60606)336* nh5>
6013606 5*6300i*h36*'b3. bei5n3hoin 636job* [253 ] 0* b3 9(536*hnb*oi3ob [ 159 ]
- 350 -
&3636 (n030 U&fiO bÔ g 6 dSH(fi050
g:>3ibiboioi3&30ni pohmbi gi 0 ^ 3 3 6 0 9 16 i 9 ni6 iyihnb 06,1136^3110 5163001111361.

bi6dhnb 039935 Pinus canariensis 51911163600 (^3110136^3 06^36^31119 0300111136b
:>0<n6iyihb; 33iPCn3(i36b 1^031119 5i9nii;|3on i 9 m6 iyiho jp 03 hj636x n
6ngi6. jjhminjjx ibyD3^n i 9 oi6 iyihob 5i6bi2)3aih36001 06^36^3110 5163001111361
a^jdooioi 3ip936bi 91 150)5031101 3ihj>mbi6 9u36ih3b {Inga-b bib3m636o).
Lecithidaceae-b bib3(n636oi5 biboioig36oi6 nj>g636nb îh&ig 5190161^300 36ihooi
[113]. Snhnqjn^bnb U3ijb0j)i0 d03 0 ^ 0 3 6 0 91 61901(3360 9g5higoi 33hnoigxn
bi6dh3&ob 9 o9 ihoi. nbn6n 9bmpmg (16 32>30i3big) bnybnb 'b35i3036ob 330935
061^3636 i(j'9m(33636ib. 10 35303b 51632)30136361: bb3igibb3i <30^30,
Cupressaceae-b 91 Protea-b bib30i&36o, 332)1003^36(1, ^0150311010 3i09i, 0hi3iQO
91 6ibi3hig 6361)0 (Cistus), 151130133 bi3i636ob 6i0iboi3i6o 015361113360
{Lantana camara gi <bm5fi3hoin 91111530013160).
hi oij0i 3691, bi6dh36n 1^11163636 9(536ih3X'T'6ib, 0i5hi0 9b3g3x<T'6ogi6
ih 3691 5i9m53hK3b i 9 90139360b 3m'bn^)n3hn 5130361 9|536ih3X(Ti6ob 51630-
01^36^3 g i 9oi0ni6ig 32>mbnb(539i cb3 : bi6dhnb ghmb 195000 ij3b 91511013000
bj^hmOibnbi 9* biqj36ob h30o63hi0o‘bn5mb 91 9 ôghoi 6iQib6ih36ob 5103R-
by^&ib, hiy, 01530b 9bhn3, b3Qb 36ym6b ib i0 5i‘bhgi 0690309360b 51630011113-
bib. y330i 'b39 i 3ohxn bi6diho 06333b b3ju3bo36ob uo2)0b ("bi6dhob 130^00")
[252],
6.2.2.3 bfll5n3D K> yfl6 3 i
oihr>r m 6 n gi 9^^03636360
l!)3033h3lj3hob 2)d£ik0,i6 3^0119 joOo^ho 3hoii53b3&ob 9o9go6ih30i6i 6309361
Qi 6m6ib6oihm6nb h35^130360 nbh360^6 3631150 ob 5i9mou>3ob3cg036ob2)36 (03 9 >
(jx^Db 3ho6t5o3o). 601503390 33^1601Qot^n ipih 060331 bi2>0ilinb 36311501b, bijr
3360 6n3cno3h3636nbi 93 Bygob 930)3nb36i 5 KJ6 X 3^DCr*i' 6ooibn6oi3'bn ihi3hm-
Q3iin3Cni. ibo9o0n3oi obp3936i gi bhgi 699361. 163010 <brn5igo 93<bp393ob 513-
C3&> 015361113^3 9 ooi oqjhoi 90901, hiy 3qjhm b9 nho 91 bi65hdco30i gi6iQ^3333h-
ii!)3hni6n 33110019360 g i 90901 51(5033606 bihobbo.
bb3igibb3i biboum^bom 3hoii53bob i2)3933hi^3hobig9o 95hd6oi6oihoi6i 5i6bb-
313363001 (61b. 6 .28a). oii39i 3oh3x i 9 bg36i 3hoilyri3oi<b9 ob (hm9 oob bo^myb-
Cnl)36ihoi6ri6i 393^9019 ihob Qi9 rn2>nQ36x n b36oij3ob 3 x 3610 5i9myniq3(i0n 363I1-
&nnoi 3'bh3633oyfTK3i'b3 gi 3631150001 9 gogiho grnbcgivi^nb 9 ^ 1 5 ^ 3 ) g36igpi6ob
9j9l3nh36i. b x 9^303 ghm9 n 313391 qjm,joibo6oi3'bob 1^030161(5, hn5 2)^619 Ri6b
ipboii (530ob 5 1^ 9 3 0 0 1 gi jomhmcgooob qjQ30ihm33ljhnnb 90161(53936001 (61b.
6.28b). bi6rn0fnmg, b30biyh30n 3ohm636ob gig50i9ob 939g35, 0(5361(13 o930i0iig
013 nb33636b 39iC3D; 9 bob 06^366031111 (11901(501 bo(503ob 9 mj9 x D 6 ob 33homg3o
303i0ng 9nb 939g35 id0O3h36b J0nhm30ibij36ob gi'bni636ob bihobbb, hiy
^X36b gi, 93bi6i9obig, 'bp3gi3b gib33636ob 3hm(53bb. b0i503nbig9n 516b!-
ijnh36001 95hd6m6nih3 01336^33630 ihb36m6b Jonhm3cibli36nb qjmtjrnmJxigiuo-
Dhn Qi'boi636nb bi 9 o9 hm36i. <bm533h 2>holyi2)Xn d>3033hi^3h36ob <3 ih5P369o ^39-
ty'i0>3h3on 2^303015036^0 ihi6mh9 i 03hig 015x361, hiy ih36o3bob goi5hi9 ib 3
6^061^361 ehqjob 5i9hx36ob ig5O0ib (61b. 6 .29 ). hm5mh(5 63bn, b0(50306 1^13-
^ ^30936 b36cnJ3ob o6(j36bo3fn6i 9 i^ 30m6 b.
9i6i0o lj)3933hi(33h36ob 0O3h 33h3gnb gi'boi636nb bihobbo 919012^93630(11 o9 >
- 351 -
0006660060 U6 6 0 U7R 80j&0 9 6 6 0 0 6 6 a 0
tiQ, 013 hs (jnMj 90333013636* 9^36^3 b^h3b3pn qjn'bnmof’vyiob 5b33i j 9n. bn-
ynjnbsgdn dfrhdBmdnshj dyjBihjjdn 13056936056 33b 50933 85806, hm(j5 63353^-
i^ h * 1158936003 &h593booi 35piC05 5590630b 63hlyiQ<b3. nb333, hm^mhy bn^bncn
5560(1633300 QiCO^snb 938016333590, 6045033 06333b 6nm0336hi636nb 9513056365b
95 3jh3gob 363hj)3ôt,3co 3t)h3B330ymgnb 83j56nl33ob 8m9pib. 35365(13360, hmd-
03605 33b 30)03636 0*650 *3833h5^3h36b, ^056936056 55906306
5hb3&m6nbib: 55yo63nb5g3o 85hd6m6n5h3 Jim300360 0(^365 8501 3056001 yn63cob
Cucumis s a t i v u s Chilling at + 2 °C
P ro top lasm ic s tre a m in g
P h o to syn th esis
C hloroplast flu o rescen ce
C hlo ro plast s t r u c t u r e
Plasmolysis
Vital staining
0 60 120 180mm
L i Normal EZ3 Disturbance H I |nhib«tion and injury
Q) 2 0 #C 10 s
>
J2
a>
o>
o
c VMJ
a>
u
o
a LI
A
u
a. 10 s
u
o
u. / - LU-UI|IIU-U4444-t-
o
.c
o
o 1 °c
>
'>
c
1 min
Time of induction
feb. 6.88 Vje* bohiom: Ijh^nb 006*5036(1 fcigiur^nbieen h3'bnbvJ36^ x 8(536^336311 95
bnuninb lbi3m«Te3&nl} bi66hc(pn3(n6nb 'bhps (Kislyuk, 1964). brihsoin: bmnj3n jcmhmojncnb
q3CT>hab|536isn5 l 0^ : e» 2 1ffC-t3 (Larcher & Nobel, 1989)
- 352 -
67)6066030 U6 6 3 li7)^n ■36 d6 r )6060
a b c
1/T-104 1/T-104 1/T-104
fob. 6£ 9a-c &ii5n3D6nlx>G9 n b]6bn^3hn (36y33(jn bibn) gs ijmc3hi6lj x n (6y3!tonCn bibn) 9^36*-
h]36nb 03(ji6riC3hn gj36J(jn36nb 9s9myi93& x 3&b (jsQS^hiii^hi'bj &s0mb:>b3pn ih36n3bnb
eni5hs336noi. a gmcncnb 63^rn-qjmljfnbn6oi3'bn (MeWilliam & Ferrar, 1974); b g)3b3nbi gi ^ ^ h n b
35h3936nb 9n,jmjm6ghn3&nb b36oij3now i^jn3m&i (Lyons & Raison, 1970); c qjfvnâiSn
6n3<nn3h36icni î&jSmh^nb bnfijih3 (Giaquinta & Geiger, 1973)
6ih9mj96 nbib. yn63ij>i3d03 (yr»63nbig3n 9l535ih33&n n6>

636 Jbm3no3i)0n biiSbmp gngn h^g^m&ncn yn6xnb 6 ih3mj36 ib.
g»'brtt63&nb dndgnGihortbi bnun3ntae9 n a&hdem&niho 3(j36 ih3369n

OhmSnjjxn 6 ih9 m9 rndnb 'bm&n^hom 6 ^ 3 6 ^ 3 gi ^ 3 0 ) 3 3 <bmg,n3hcnn 9(536ihnb
(hm330cni 3 3 6 3 6 ^ 3 ^ 0 6i6n(V)6nih ihni6 bn(yi3nbip9n 3f,hd6m&nih3) 0rn06ncg3&nb
3hrn53b9n Symcgn 6iymq)3&n np333&ni6 0-1 O^C-nb q)ih5Q3&3n. bnun3noi
5i3pi533xn gi‘bni63&3&nb bnph03 g i 3ndgn6ih3mdi 3hoibi g i n0i33 0i536ilv)3n
QiOmyig^&xni 6il5n33^n^ bihnbbbj, biSfchdiyvjn&ibi g i (J3933hi^3hnb &ioii-
6i&h3&nb bnRjihj^ ( 5 ^ 3 3 6 1 ^ g i nb^3 fcioxbmbnb). j>i6bii3cnh36non hagpni
3diojhn (b^D hi^jhxn &igib3C3&nb ((ioQSah^hxn 3rny>) f c i g i ^ i . gi‘bni636nb
bihnbbn Qi^jQmbb lj,3933hilj3hnb ^pg&ibi g i b,jh3bnb bi6j)hdQn3fn6nb 9cn0i^3-
6ibo»i6 3hoiig (6ib. 6 .30 ).
9(j36ih3369n 3hmlyn3pi'b0 nb bn(jn3noi gi‘bni636i 3nou>hg35i 3hm5 h3b x i& (Bib.
6.31): oni3g i 3nh3X i g n‘bc,3g3&i i6 63993!!^ 'bm&r\3hcnn <5336j(jni, 933935 cni3b
nfi36b 333ju33ign 3hm(53b3&n. hmfcmhu 63bn, bhjgn gih(i>333nb 6096360 5 i 9 mR6g 3&i
hi3g36n03 gpnb, <bm{)33h 33nhnb, 3 ^ g 35 - bnyn3nb 3 mj03936nb 3nh330ign 3 0 ) 3 ^
5i3rrinbil!)36i n9 i 9 n, hm3 pn3ng3hn yn33m636l!)3&n otb33ig-j)hnb(!>iC3hn 0g5m0ih3nv
6ngi6 &>gigni6 &3cnb q)mh3idn g i ig&npn ij3b 3306hi630n 3hm^3n636nb 93(bmhi>
□eî^rtib. 3b 1530(1936360 0633336 33h3gnb 33p633igm6nb 93b3bli)36ib
gi 3sh3gnb 3n5cni3bnb j)m33n636l!)36nb gncg^bnib 3nb j>ih3cn. 6n3cnn3li36ioni (J3(yib
3hmi53b3&nb gih(p333nb 8 39 35^ g i b36or)3nb i6i3hro63pn qp'bnb n6^36bn3m6nb
9n3i^ 36nb 5i0n QB^h^nnb S h m g jj^ 0tfnhg36i. 3jh3g3 n j,hm3936i 33vî6 n 0n<b3nb
93iQ3g3hn g i bb3i tynjbn^n 3hmg3ij36n. bljh3jlj3h30n 3homi6fn6nb giîh^nb
339936 35h39n fiCO^Dbi.
bntjn3nbig0n S^hdGm&iuhmMb 806393001 djndQybs îSmnymb bm&>Q5p
d&hd6mbru>hj (3. n. Siam y33C^ 6i6non 3hot6inhig 05hd6m&nih3i) g i 6s6ncmdhn3
dfthd&ndnshj (Qift., y3i3nc3&n gi 3 m36 nq336nb 3hm(j3b9 n Symqjn 6iymcg36n) 3(336>
- 353 -
90066^0060
6*b. 6 3 0 3360b b^hnbbnb^ 93

bb65hdco3fT^f)b 6*3006* qjmcnQob
9^ 0^656^ 3. 80C-ftb b 33ncn 2^ 0336*
*h*6*nh Q^^bn^S^bx Dqnfeb *h 06333b
Duration of low temperature [h] (Larcher & Bodner, 1980)
Phase transition in biomembranes
Liquid-crystalline Solid gel
Consequences of changed membrane structure

. Increased permeability
jg Reduced selectivity in membrane transport
e Increased activation energy of
membrane-bound enzymes
Chloroplasts Mitochondria Protoplasmic

membranes
Inhibition of Increase ol respiration Solute leakage
photosynthesis RQ<1 Impaired
L e ss metabolites less ATP, N AD H . active transport
Le ss A TP
and N A D P K
Lo ss or ion6
Accumulation of
stress metabolites 6*b. 6 3 1 &btyi33&ntagdn b^Gbn^^h
and loxic catabolites 8^36^336*63 gbdiQn ^333Qh^3hnb
5^3036^ (Lyons, 1973; Levitt, 1980a,
Injury and death of cells \y\$. Larcher (1985) 606393001)
h;ft6n. bb3i©ibb3> nh&i6nbi gi Jjr^ n fb ij3b bn^nlwgQn

9 j)hd6 rn6 fu>hm6 i (6bb. 6 .32 ). &ihi3330n bibnurnjbcm qjbtb (orrjbcD&nb
bn&Dh]) 5 i 9 mnhfi]3i fcbtjbbiijonh^xn 9 &hd6 m6 n:>hm&ncn.
- 354 -
57)6356030 1>&6 3 U^ 0 96d6 f )6 0 5 0
5*yo63fi> 3hn<£)bo 9* drib dngh a xIfflS m n n 9*1x1^636360.

3(536*11380 <^06390 ou>39*3nh3x*9 6*h0moj8636* n0 *95093680, hm093&m5 1)33-
Ci'b] Rjah* 9* *9309*9 (503936*. 3b36o*: ^ h n q ^ o x o 5*0^*ho 601^501360 9*
33(13980111063680 501693660(136300 6y*pn. '601503(101 6*j^3ho*b (Erwiniab 9*
Pseudomonas-rh 6*63016360) 5**fi6o*cn '639*30(1390 3(101^306360, (101803600 6y0r>b
9f'CDiX32,f>'i 9*5*380(13606 83933016001 5*1*06306 631^5006 (*196*3 9*69*
50-100%
10-20 % 0-50%
40-60%
0%
10- 20 %
c 100%
Sib. 6.32 ei&spn (+1°C ) 5*3036* Coffea arabica-by s o x * ^ 3 <bn*6e36:>
(ei'bni636i J>i9mnbi^36i %-noi) 13303360, isS&rg&i, j^ysnonc^xf1 pi qjmcnc36n, 6 * ^
6ncn j 3oh^3&n gs 0130005*6 ip 9m6i^36o (Sakai & Larcher, 1987)
^63336 9*, *9 0bhn3, 9330)90*01 6(536*1-106*01306 uqjuinhn 63503*^01(106 hoi0ob 83b-

hxD&i- 6*yo63ob o9330bo 5 * 6 ^ 601^,013690 9* 3m9 mj,363^1 jbm3nc369n *9309*9
3^(539936*, 601901 3m9mj)363hni6ob g*h(p333ob *95093680 (3*3hob 638(535360,
9 ^nghm 5 *03 hj63& x n *5 53(jn6n'b3630o 03mh*9O 5390360) *8(50(13636 yn63Qo6 6*h-
9mJ06nb bofij*h3b.
6yono) 9go0*hn, 01*309*633 916*90 ^3033(1*153113606 9mj03Q36nb 83935*9 5*150-
336300 3hnl5m30*bv536nb 5*yo63ob 3(1015360 0o9go6*h3ri6b jsh jgn b dn&6ncn. *9
phmb 33(13980 y o 6 x n^ 5(^(5*9360 6*h0oioj0636* 3p3nb3&3ho bob6h*03001 (6*b.
6 ,33), hob 150^0130*^9* ^0*6936*. b8oh*g, 1J06390 01*309*6 6*h0mnj963-
6* *h* 3hoil5oi30*b,5368n1 *h*039 33h3g9mhnb369n 9* 838935 - 33(i3gob 53939b* 9*
3hm^rri30*b(i)b Smhob. *0*b 3601936* jjh jg fr ih j yn6jQnb 6*h9 m8 oi6 *. 5*9015(106,5*03-
- 355 -
8 0 066 6336 0 1 6 6 3 6 3 ^ 3 ^ aD 8^6 a n 6^30
6c>b. 633 3j h 3e 9n e i ( 3ih(jb 36s b^hiowi) 95 3j h 3e& 3h3 (0 ^ 3 33 6 1 b^hicrm) jiy n 6 3 i. 3n h 3x '
3 3 6 0 *13 3 3 ^ 9 0 6ih 3mJ36non yn63cnb ih n l^ e o d * 1 3cnens5>g ^ ^ 0 4 6 36 36 Shryjm SpibSnb b ^ h rjj^ ^ h ib .
35(13053(13 5iyn 63rtb ib 33h3pn 6y(vib gij>ihb3nb 53801 g i 53(13 6 6 3 3 ^ 5 * 6 <143366363.
933m b i'b n h 3c n i yn63cnb ihnb)$ac3&ncn g i 3nbn 5ih bn b qjm hSi 3ou3pn(yii (Moor, 1964)
6390 yr»5 x n 600^639366 (101501(1(5 68(1390 ^33(10, (139536 1)063906 639330(163 6ypnb
mhoiJcnb 66333 ^qjhm 9363903, 309(13 5 H5n33&DCn bb63hob <63g33oh'b3 . 30 nb
6393539 69363 3(101(5013936(53606 533&y90i363 93 nbn6n dpn3h39 (80i(5390i6nb 2 /3 -
693) 0^636636036; 53666090 6030103(1363606 ^6(336(^13(303 630160 63(5390166. 5 3 y-
06306 3(101(5360 5(10309363 0J3893, 6363d yo63Qob <336363 93 33h39nb 6336b 3 mhnb
3h 93893(19363 013)16019063005390 601636601(10163. 601636601(101606 63^4090 930005093-
63903 ,53833113(53(1363: -5°C 9 ^ 9 3 6 3 -6 MPa-'b3, 601901 -10°C — 12 M Pa-^.
636^013(^6006390936 53611169063113, 936390 (53633(13(5311360 0633336 ihm^m-
3936806 539m6hn636. 6015030b 6(5h3bnb3 93 3hot(50i39368o6 53361)9013606 3 ohm-
63660, bb63hob 5331^0633 6360960 83(5390166 83(10936063 93 0111536390 83333606
3016(536(5(13(503, h3(j 06333b 36608360b 93345033(5036. 01680163(10 d3036ob:> 93
80x5390160010 ioi6u36^h3ijoob 80183^3606 53dm 60018386(136360 0)903(1 0036363,
(1363(5 0136 69336 8386(1363606 9030936063 93 3(101(53063606 93093 (63b. 6 .3 4 ) 93
6386(13606 3(jq)-3<b36nb 636(5011363. 53^630636 6306(1363606 9360363606 31100-3(1010
6063603 (509350109360936 3936(501(50360606 536mymap (ob. 63b. 6 .37 a). 63601901019,
3hoy5o»393<b8 ob 5336y9oi36ob b3ho6bo (6011538390, (101500(1(5 6663(1063 93 6yshn 033^33-
6ob 630(3(19363) 603(^6336 08 901636, (1019363(5 395090 3^36 3(101(5013936806 b(5h3j-
(53hnb 633*53339 93(1^333366.
53yo63o6 6(5(13606536 013393(533

06 1360(53(53600, 6393(5 53^6306 63008(101363 33(100193903, 6(5363(133603 530 rr>-
00363336 00393(5306 6h33390 636339363 (63b. 6 .35 ). 363603 3h3 66019019 531)06306
- 356 -
5360&6030 b 6 (f> a u ^ 0 ^ddnftOftO
(n b . 634 qjmcnpnb Jpyn63i3n fo ho ixn SHmu^bj&n (Sikorska & Kaspeiska, 1982)
Frost survival
* + +
Prevention | Surviving of ireezing events Surviving of
of freezing secondary effects due to frost
i I
I
Spatial frosl
Freezing avoidance
1
Persistent
Freezing tolerance
I
Tolerance to
1
Protection against
exclusion supercooling cell shrinkage photoinhibition
(life forms) during winter
Freezing point Tolerance to
depression freezing Protection against
Temporal frost dehydration winter draught
exclusion Extraorgan freezing
(seasonal timing) and translocated Tolerance to
freezing in tissue hypoxia under
spaces ice sheets and
compressed snow
Resistance to
mechanical strain
caused by frost,
ice and snow load,
and snow pressure
6ab. 635 S ujfohjoto 93633^ yn63ibcni6 g i ‘tadonhrib
l^h^bnb bi6n6iip9p35rng dncbhoixn ppi6n6dn3&3&a ihi03g <bi9crhr»b brnyv

3nbiOi3nb Qb&tabfuogbxn 01^68933(1 9*11303636116 (grwinSSndnuni, lbb9onhnb bnd-
9 hu>03 , oim30nb bn9 dn93 g i bb3_) j>i63nljhi03&i. 01536^36 a^jcJgni ou»3ngi6 in-
ly^xnb, Sîhbngnb i6 33^630016 ^063001 5i6m6333pn gi'bni&j&Q&n. 5 iyn63fl6ig 9 n
lyT'0Dhi6iX(T* nl3 bihnbbn ib3ai 3i 6nly>^363fi 5igihfr)6n6 (yrtgqjibni.
5 iy n 63nb ih n g3 &i6 9^ 3 6 ^ 3 i 6 3h63& 6 or*3h9mn'bfn0i(jnr(6i g i 6no*6rob gij>sh5 3 ftb
039 l5nh36n6 6 i 9] i 036noi: h3j)363hi(yin6ioi3nb 06*133630013i6n 330^3606 Qi3 i 03 i
3^nghm 6i9pib300n (6i 0 nc3i 81536^ 36(1, bhgob 6i 0 O0 n636hn cgmh0i);
- 357 -
0036663360 8^80866306630
5 ig i(bi0aih36nb 01(1516013606 516015361 03010103606 bsq}ihj330, J3O<335n03n 16

60191500 (53^1030^360); 05hd6oi6nsh3 01(15160136(16 61901(33361 dpo3ho yo63nb gi6y3-
6nb 606 (5iyn63nb 6i9o9hoi36ob gh111360010 1(109361 0301010100(5336 915361)13363(1).
li)hfTi3nj)36nb 0oioi6 h35nm6369o 5051663(10 hm'b3630n 6(53611133606 0901010360
p i 9 noi 93601363331 0(5361(106 bb3i 61600366, hiy 06333b 630^3(116:^300 51601 bb-
03360b 330^(13616.
5iyo63fib B^h^yicnb 01^33001361 1033(16366 6y0ob 5iyn63nb Shm^bb 9(j36ihnb
5x03003690. 5ibb6o0o 6i3hoi36nb 63933016001 5iyo63ob 63(16090 303)1ro 9 16 0 1
0633b. 33h3gob 63360, 9ib 9 o 5ibb6r>0O 613^1013606 3016(5366(11(5006 906393001,
01^06361 -1°C - -5 °C-nb 031(1503690. y,yn 6jn b Sjh^ngnb g id jji ihob 906306516
gH53ob 03gih36ooiAb3bô, 615(119 36y3iho bi33i036i. 5ihgi i 9 obi, 33(i3gob 6336b
ij3b ‘63963619 511503360b 361)10; 3.0. 33(5hig ih oyo636i 51130630b 63hlyi0Ib3 9 16 10
^3933)116^11^3. 5 1 ^ 6 3 1 6 1 gi ‘63963619 5H50336i'b3 91330)13361 6 3 ^ 0 3 6 1
6(5361)1390 ^3933)11^3)106 916330b 5i'bm93ob 3hro(53t>0o. 01139131-1)133010 Jbm3n036n
(5039361 3i3hob ^3933)1163110b gi633ob 3ihi0X3hig, 939935 ^hob^x'n'bxsnnb
3hro(53b9o 519016011306309036300 363(15006 bihjt^, 63033hi6Dhi 91(1)300166.
i 9 n^m9 , ^D^ÎDhi^hnb ^3010 601616361 C^boKrqhOi") ihnb 511306306 3hoi(33bob
gibi6yn6n6 91633636390 (61b. 6 .36 ).
J6 o039o6 601^3013600 g i 6y0ooi 9gogih, gngn 33h3936nb 6)0163 3ih36jo6i9o
5191(3033606 695016^30161 ghnig 916003)101 (^hg^d^Q n ^ g ^ n jjd ^ ), 1150
003010110 5)^309361 hi8936o93 biioib3 936 bi6b 91 ih ihob 3033^3)10 6165(^9030
906306 30(10163660. ^013093690, hoi9036bi(5 511660101 633031(5006 61)103)1360 (bj30o
g i 9 ^nghoi 33(13906 3390360), 936199363001 33h9 i 6 366300 5191(503361, (101630015
93 nd03&i 9361)1636936 gngn 6606 5i 69 i 30oi6 i 9 o, 6 i 6 i 9 ih 51910916361 h ip 1^5
'b(j'3h30n 63933)1163)1300 6iho3ho. 33h6 i 6366X 5igi(5n336ib boi5n3hcn cgoKnoi0 9 n
195090 ij3b -10-12^C, ‘601603(10 bi(i6y0nb 63 9(536ih336nbi 91 636^3606 y0oih6369n
91 93hji69 n (633(10 6yob 63 6(5361(13, 636090) 30 050 -30 - -50 °C 1^6336. i 0 nb
636936 936ib 6 i 6 o03ho 9950101(130161 0)1(^3331 g i 35h3gnb 51(1301 51691536300 yn6-
300 5 i 9 oi3hnb6 i 0 o'b03 &i (916190 63933hi63hob 35‘boioi3h9i: 616. 6 .36 ).
’601503)101 0136960, y0oih6bi gi j3hjnb J6oi3n09o 195090 ij3b 03bi93 ^n3 nb
gH53nb (J>oi6o6do36ib - 6^^66901(501136x0 yn630n6 6 ih 9oij96ib, hoi60ob ihbo nbn,
hoi6 6yi0o Jboi3n036ngi6 gi 33h3g9oihnbo 01336360016 5(10139361 (51)1030
195093690, bignj 050 030g35 906390b gob3h3630 6163619 51910^5331. i 9 nb
0393519 33(i3gob 63360b 3016(536^1(501 016390166, (11(5, 011306 06(103, 063336 5iy -
n63nb 63(160906 9163316. 6y9n6 91313360b 619319361 ij3b 15(130133
033m3m0obijihn036b, (10109360(5 5(1013936016 <6 i 3 oihoi6noi. 015., Opuntia humifusa-
9 n [ 145],
0(5361)13366, hoi0036n(5 d0n3hn 90633606 30(10163660 1)163601636, 3691 511669301
511/163061980 695(1190 3hoi6oi30i'b5i. 5113063061990 6rlCDhi6630oi6i 900^6331
60(503061990 6616093(10 q3oibqjoi0o3o036ob 06635(10(136001 60019306(1163600 (n6.
616. 6 .35 ) gi 666190 6i69oh6yi06i036n6 (9 jihn g i 019050161^1)100360; 616. 6 .3 7 ),
3oi9noi036n6, 91619001933x3(10 iboi6oi3i6o 613(1013606 (i9o6oi9gi336n, 3oiqoi-
606360) gi 6yi00n 666190 (5093606 915(10133606 61631036001. i 9 901393606 010(533-
00 03ji6n'b0ni nb, hoi0, 3(1om 0bhn3, (5o6oi30ib6i9n 61600133606 30x5366^11(5006
80181636001, 6y0ob 303(101 8ipi0n 8391533001606 3nhoi63&9n, 600(^6331 013(160190-
618033(10 601616601(10161 gi, 8301(136 06(103, 63669361 gôghiyoooi 510 01633x0
gi63nhoi3i. 51)191 i0n6i, 91610801033x3^0, 9306hi6o6ig0n 63n6hi93ho 613(1013606
915(1013361 063336 6(,’jkn33hn 0016360b 3016(5366(11(5006 830150113616 (i^Cndîijhn
jq}jibn) e> 303j6hfT1Cni!0&n,J 6^0*1ic)333&^ ) 0306h i 6nb 66h3i33h369n 03013190
- 358 -
67)6066030 US60b7)E50 36d6D6060
9
©
2
course
fbb. 636&-C a 5^yn63nb gjife^bn gi

A j,iyn63nb Bjh^ncrtb
eiiBDno'36i; SC dpnahn 5^^631; NT
ynBxnb 6ih8mJ96nb ET
od'bmoohdn; FP jpyn63nb Bjh^ncn; MP
iy>g6n6nb e^tyjnea b 3sh;)6jn9nb Sîcn
9D9ll3DCfn^n°1 6i 9mhfi3x Pih{>i6mcni (i9
930oib333i9n anbmb) fjiyr^nb
0i{pCnom: Tf jpyn63nb c
[fOhmb 5pyn53nb 91551x 1010: THE
36bmcn3h0nb 3 ip ^ n LTE
3j)lbmoi3h6nb gi6sen
(Larcher, 1985) Temperature [ ° C ]
(300360956.
5*yn53nb*e0n lyn(V)h55vjx r ,K!)i 8(3365(106 55630015(13606 3hoi(3363o 5h5on56565hn5:
06^36603(10 6031x3300 ‘bhgob 3hox33b0 o 8(3365113015 383^360165 5h 0300166 55-
8oi6hoim6n6 365116 9 5 580,1)018 956500 j)3833h5^3h36065980 qjho59 851x^016(^35.
63^0163110 50085^06 8^0163 11350016360b 9^365(13360 33(13906 55h3 yn630ob 65(19ov
J86o6 36511b nd3636 33801950180601306 (65b. 6 .36 ). 8(5365h33&nb 558016(10101606
035^58(1 559563006 30(13390 3ohm655 bhgob 3(10x33606 9580153(1365 (b3 8(33-
65(133600) 5 6 0363(1365 (6505601356 8(3365(133690). 013 558016(10101606501301) 3bse3-
65 9*bh3095, 069365 655301(103 558oi6hoim6 ob 3110x3360. 3b 3(101(3360 b5033b3h36hn3-
- 359 -
auae-âa&o usrtauac? aân aartam aao
® 40
Stress temperature [°C] 0 40

Concentration of protective
substances [mOsmol]
fob. 6.37a,b nb3i5ibnb qjmoic3i>9ri bnndjdhiSjbnb g4bni63&> j>iyn63nbi gs j>:>c(ymdnb ghmb gs

bb5ign ^>h6m3ngh^36nb 9^6^3300 3(33^ (Hincha, 1989; Hincha et ai., 1989)
^♦Cooi to warmr— * High temperature Very low „

'< temperature ^
temperatures
Cambial Cessation of growth /^ S a

activity -------- Leaf ~ Leaf fall |
Cell structure
“ Flush ~
of growth
yellowinB in winter
i.Frost resistance 5 Frost resistance
a decreasing J5 increasinQ
Break
■Onset of •»
dormancy
Early Late
; Rapid Maximal
summer cell summer cell
i dehardening hardening
Carbohydrate
■ —Sugar—starch— » assimilation Starch-sugar
N . conversion conversion
proteins. RNA
phosoholipides
6sb. 6.38 3l536ihnb yn63:>5s3dc3m6;>9n Rihcn^pn qj^BrnQr^n^hn, yn^mQm^n^hn, ^n^rijnSn^hn g>

O]n:bnm 0n5n^hn Q ^JiG rrbS ^n. pi6^hnb3on shncn 6^^336360^ 9:>jbn9ip3hn b3CnC3^DGn
yn63if,i9dpyn6i3 n (Siminovich, 1981)
- 360 -
67)606(f )030 ^ d 6 D ( f .a b O
fii [254] gi y3X > li^qjQb^hb 9nbg33b 9^9933601

9(536ih3363n 530m6hcnm&nb 3hn\j3bn n 5 y j6 i 01133390 gpnb 36 Qhonn jôhob
5s69^30m6s9 n 0°C-b3 F19633 93p300 Smja^g^bnb 3393539 (63b.
6 .39 ). 30, 5i9m6hcnm6nb 6063 bMjQb^hbj 03j 3ho 93 bb33 93915330 633horj6n
5hm3g36i 3hm^m5c ;>b833n, 6y3Cn 3311393600 iOO&XfT'&l} Qb l3D6ibh;>Cîn 33i T r'Cn
nynqp 0h 3330 330)^3 3 3 *1x 10 3 6 3 9 . 30 i h r u jj b Q b n b 9 3 0 3 3 ^ 1 6 0 0 1 ih m ,!)P i 303,b 03 Sfb^g-
9363 0309350 qj3‘bob3cn3ob, hm030ou 30 0133631133690 9o9go63h3n6b b ^ b ^ yo63ob
3nhm6360n (-3 - -5°C). 30 ghmb 6om0396h3636ob 93 3 6 ^ 9 3 6 0 53hg3nj9-
Months
Months
Bsb. 639a,b 93hj6ns6 9u]5 ih30u> yn63iîddp3fn6nb l^'bmdjhn gn6:>dny>. a 3^3 cnb 336>^>(y'3hn
i 3nh|ij6ob Ii3lbnb1j 36,j;xyn6:> (Tyurina & Gogoleva, 1975). b Rhododendron ferrugineum-rh
qjmoic^&nb yn63ijp9dp3m6i. D&CD&n ^336360 bb^mtab 9n5n9i(vjh h3%nb^36lyxmb>b:>
pi 3m,j36(jn3h h3'bnb^36lj3Qm6ilj 3mhnb 0i 6<<nha(y'ih n x 3nhm63&9n. 63636^ 03(1 inhmbjbnbso^nb
an63nbip8n h3fenb^36lj x ,,’&;> &s3nba#]oni 9]Jn 3*30 bitncn (Pisek & Schiessl, 1947)
- 361 -
9006660060 U & tfO U ^ 3 t?aD 8 6 6 0 n 6690
636* ob3CT6*nh*g, hm3 331139b S^jdpmb 6990b 5*h53nb 5*g*(!)*6* yn630nb qjmhanh^-
60b 3hm|53b3n. 5*3m6hcnm6nb y 3 X ^ D 3 *e>2>0 bihobbb 81536*113360 ie'63336 yn63nb
33h0 *536li)30n 0 fnJ03 Q36nb 3ohm6363 o, b x ly1^ -5 - -15*C. 3q}3j^3hn (53833h*l53h3-
60b <3*11603680 bb3*g*bb3* b*b3m&* 5*6^3*336300*. 013 d0O3hn 9063360 8369936*,
3hmii)fn3pi'b9 i 36)136936* 5*3m6hoim6ob 3oh3x <3*b*b, 3*5h*3 15030 ^ o m g n b 9*9-
5m9nb*b h3(bob^36v!)30m6* ob33 3 *li,30m6b, b*6*8 81536*113 *h 5*9*3* 8fnb3363&ob
qj*b*8o. hm^mhu 50 b*8cnhnb 3mb33636ob cg*b* 5*O30nb, 6*8m6hoim6nb 36*110
05*11536*.
b*8oihnb 30(10163680 6936* yo63ob*90o b3bm63ho 6*8m6hcnm6ob* 9* b*60mj)C3
06931501136390 *g*3ii)*(503&ob (hm0036nb 83833016001*15 6936* h3<bob^,35l>)3Qrr>6nb
b6h*ogo h35300(136*) 5*9*qj*h3*. 5*3m6hoim6*b 5*6^5301(136001 639b 36ym6b
b* 3 oihob 9*95^16*393 *hb3630O 90630b $31100)9360. *3 ghmb h3bobl536l530m6*3
93 od03&* h*3936o03 g(j>nb 5*68*39016*30 8n*(n6omb dspodjdb. 530*6 “b* 3 oi*h3 n
8380163330010 016090 *8o6gob 9*95018* 3156*9 *8150)1366 yo63*5*8d03oi6*b, *51130133
83* b*9 oi*h8 o h*6936083 gcpnb 5*68*30016*80 +10 - + 20 ®C l53833h*l53h36nb
801^83936* 3l536*h3b 355(15*3b yo63ob*gdo 5*8d03oi6ob 36*hb.
6*0*bm3*6 81536*1133630 5*3m6hoim6nb 3hm(53bn 808906*h3016b 5*8*11(50336x^9
[ 109 ]. 6*8m6hoim6* 06936* +5-9*6 - 2 °C (53833h*d)3hob 3oij33036ob ghmb. *8 838-
0^333*80 6 *3 m6haim6 ob 3hoi(53bob 8m3b*g3&*b *95090 *h* *j3b, ggmb b*6£>hd-
0O3m6nb 86083639016* 83mh3b*hobbfn3*6o* (*6 b xX 5 *11*3001*110*). h3<bobl536(5 x n'v
6*b. 6.40 *6936911 9m'b*hgn ^363110160 6153-

6ihnb Solamrn commersonii-b
yn63*5i3dc3m6nb Shmu^bjbnb gn6i 9fv>i
ghmSn. 916*011 ^^hi^jhnbsgSn (2°C)
9353361 (0130*111 nb*hn) 9n9gn6*h3m6b 20
gpYib 5*59*3cm6*3n, bmpm 938935 hjiyyiSv
lyi’b*(}n*b Sipxyi ^3033(1^3(13&nbip9n (10 91
20°C; 9*311 nbh36n) b^nhg3&* 24 PP3 (Chen
& Li, 1980)
60b *9 *(?03&nb 36083630013*60 3ohm6 ** 6*b3oh6y*06*936ob 9*51101336*. obo6n

5110139360*6 o8ob 5*801, hm8 6*g*boi3*6o 81536*113360 b*8oi*h3oi5, (101501(115 63bn,
gml5mbo6ai3(53h*9 *^03(1360*, d*5h*d *3 ghmb 6*h9m36390 *^809*^360 *h
6*801093636* bhgob 3hm(53bob*oi3ob. 6*9*bm3*6n 8i536*h336ob h3bnb^36^30m6*
9*l530ri6 b 90630b 33hnmg36nb (5*6^530^36001 6o*g*5ob 5*yo63ob) b*65hd0o3m6ob
0m3*(j(36*bo»*6 3han*g. o3 ob 6o8*6b, hm3 5*8m6hon6ob 3hon53bo 3hoigj*bo*6o*,
6*h3m*g536b b* 3 cnhob b0i503nb*g30 b6h*qjo 835336*: 8(536*113 3(536 n3ooi*h36b
3*Jbn3*03h h3bobtf)6ti30fn6*b, 3*5h*8 *b333 31536 5*h5*3b 3 *b (6*b. 6. 4 0 ).
- 362 -
67)606 eo030 6 6 6 3 6 3 ^ 0 ^ d a n c o O & O
6ib. 6.41 bb3igibb3i ipoQi^jho cbn6nb 6063^13^60 9(}36ihob 3(^,536(30^0 yo63i 5 >9d03m6i (°C)
(Sakai. 1983; Sakai & Larcher, 1987)
6nun3nb*e8o h^bo6.ifl6^-)noi6o6 30166^)0,531503(10 ,303360

y33Qi 8(5365(136 ih diQ^db Q^bbQn ,33033(15^3(13606, 9ncn 363^36, 906306 5,^565
95 y33c^ yn53 S5 i 9 dcD 8(506^3^ 5(1 833dQni 558016(10101606 93395 ojifenb 55305.
SbJjnS^hn (i3'bo6(i36j)30oi6nb 0ob3p3ocn (nb. (jbh. 6. 3 . pi ybh. 6. 3 ) 830(^0365
550oinyoib 95 n3 3 6 3i6n 35303360, briQn (i3'6obv536lj30oi6o6 1>3dIS<>-
qjn^tfhn 015306363(136560 556530(10163636 8(5365(133606 553(1(5303656 6^33330
q)bh5C3&9n (656. 6 .41 ). 9063555880301606 9g5hbQm5nb bb'brr>8ig 5P 3&30 O5 lU ^ A "
5^3(15, hm9 xt> 0l5 8(5365(13015 608383606 61163(16 656335(10 0^33365 feWPCJho ,538-
30h^]hb; TLjo).
bnynjnbspdn dfihdSmdnihj d^j6Jijjd n : 50 35303b 333013606 y330^ nb 0(5365(13,
h(n0Q36n(5 5>yn63nb 63h,yi0'b3 309(101 05(p50 3 ° 655hd6ot605p
'6056936056: ai6(i0O ‘bc'33606 6y508(5365(i33&o, ‘601503(1010 601301 p5 (11(1013033606
'6015(13(1010 ^3h^Cfn3i ^n 8(5365(13.
g^yn6nbigdn d&hd6mbnshj 8(5365(1336(1: 3b 353030 86019019 906306
3nj33g3&nb 930ibjbvJ3630n 6583503636001. 60fib (503 33(100193680, 33(13906 6 336 ^
gs 3h(T^m30b<b9i9n 05^300166 oib8oib3h5g i ^ y ^ o 653(1013606 3016036^1^15(1*. 601300
60nb 5568530016580 85(18601605(13360 ihru>6 15030 <6q>336o6 69598(5365(13360 95 6^3-
65(10 6993606 "601503(1010 69598(5365(13, ,4(101303300 95 636^101303390 £3(1^0013560
61536^3360 pi 1301803(159 016000 I13500163606 6635956635 65630165.
yn6j i i >^ddQj d(jj6±hjjdn: 63<6oi63h5g 9016355588935 8^365(10 69036065 p5 80^3-
3>8mJ(533ob "601906 '601503(10111 6950815365(13, y3305 30085(33(10 “601606 (8501 3 mhnb,
^hm3ojj]0n) 6536360 p i (5030 'bi8cnho6 8^0163 h3500063606 8(15350600560 8(5365(133-
60. '601503(101 69508(5365(136, 633(1 0OJ366 p5 6635956635 63 8(5365(136 565605013601
3^6^(138503(10 906355588930165, 06060 6565(18003 pi d0O3h 9063366 (>35636 p5tbn>
6363606 55(1383 (3^633(10836^0 30)93636 0163359(1 5‘bn(yi6 0)3833(15^31156). 05050-
6oinb '601503(101 6595601356 815365(136 8338905 yo63i5i8d03oi6o6 5580183853365 8013-
- 363 -
8006660060 bS 6 0 bc)cN 8 ^ a n 8 6 6 0 n 5690
Abies alba Acer pseudoplatanus
b b . 6.42 Abies alba-b p i Acer pseudoplatarms-rfo b b ^ ib b ^ m h^m bi p i J>m3ii(Y>b yn 6 3>

5^9d(X)m6i bi3cru>h3n g i 30M<!>50nnb SQhnmpcJn (Larcher, 1985)
pn 6pnb 5560530016590 (3.0. '6503630005) - 59 938016333500 9 o6 #*>13003600 6>h-

9010^66365 33113906 5 ^ 3 yn53cn.
yn635550do3oi6ob i 95 5595^3606 ^Bahnb 35(1056300165

6635956635 011155601 95 J6oi3o0n ^3033hi^3h3co h3'bnb^35vj30ni6 ob 9bho3 3(101-
85630106556 65385019 06083630013659 556663539365 (65b. 6. 42 ). b0oh5g, (133(101-
e3J6n3Qn 01(155601360 55665330*36001 ©5(^601605(135 1^063065990, 563 955., 1)3530-
03606 6565656360 b 59oiho6 y0mhj)369o 56 65ymgjo6 13366360 53353003680 (656.
6 .43 ). 65385019 ©5(^601605(135 55h30i33 00606^3385 01(155601360. 6010^336065 95 ho-
'bn036o6 60(503065990 h3‘bo6lj36^30^656 5301030^3606 shb3601606501306 5595961333-
(jo 060836300165 5^36. 63 85365(133606 0010056 h3'bn6^36^3001656 5566i'b(ri3h536
<3363015 6o6^30o6 5583(11)636300 656003606, 556653301(136001 033630b y30nb, (i3<bnl>
^36^300165; 50 656003606 95(3*33356 6011^3365 015365I1O6 60339000.
y330i'b3 65303659 05hd6oi6o5h35 0o6o6‘b305 65600360. ("3(1013606 35660380
6I13Q0 559oi6hoioi6o6 30(10163680 6635 #>01300360156 93g5h36ooi 558010(16335 y33-
05^3 05c»5oo h3'bob^36(j30r)6ooi. 5565603606 330(1^366 5j3cn 0ooi 3qjhm 05C 5oo
h 3‘bo 6 v;,36 ^ 30 Oi6 5 , (15(5 30(101 b 565 hdQ03 0 oyoi 1)06306 801^639365 (65b. 6 .4 4 ) . 330(1-
^360, hrn0o36ou 0*6580*13636 65083901 958(3530 65035(106 5511383, 06301033 8 v
(n5ch3,bob(336(33coi65 565605013601, (101501(1)5 £>3(101366 (*301330 9380*6333500 3qjhoi
9i(p5QO, 309(13 0301010366). 6059550156 5b0o>6 99365(13 550oib5<b50oih3630 330(1,3366
30 9565(^1 h3<6o6lj)36vj3coi65 555660501. h3<6o6&)6&j0o 33010015006 5633ij0o yooih-
,336b 9090 860936300165 ij30i: 00 0390*5333590, 013 330(1,3360 J563hoi30fu>. 653-
- 364 -
57)6056030 l»6 6 3 b 7ic ?0 3 6 d6 P)6050
6lb. 6.43 B^Snb y3^3ri0nb J 3nh^36nb

yn53i{,i8dcom6i (9ih(jbo6i b^hicnn) gi
3s3pnb i3nhlj36nb bnyr^nbigdn hj'bnb^B-
jp'bsqjbx'to (8^53361 b]hicnn).
<fc>3n ojjhntn i(n6n96x(v> i 3nh,j36nb y3j-
Cib3 3ghm b]6bn^3hn 3663611 (Pisek, 1958;
Larcher, 1970)
Therophyte P h an ero p h y te C h am ae p h yte s H em icryptophytes C ryp to p h y tes
Bib. 6.44 (yi3'bi8cinhn}6n h3j,rv'>63&ngs6 'b59cnhnbs@9n 63^336300 3yn9mhqj3&nb yn63^9d^3n6i

(>3^rnhnb33(y» 9m6nj 3836(1). 9^16^39360 53n9h36nbiOi3r»b nb. Sato (1982). ^3h9nbn: *bs9onhnb:>p9n
^063D^Xn 3y T'9mhq)3&n" 933b>ijy3nb3?>i h>36ifV)hnb (Raunkiar, 1910) *b>bfHjm(}bom qjmh93&b"
6bhnbcn3nb qjricnmcnj335i ih ihnb nbj Qi9 i<bfUi636 x n* 3 ^6 ^ 3 qjmcn036ou>6 3)10159

530 )1^ 36 0 6 g&xv&ybs 3533 5 5 9 5 8 6 (^ 3 3 ^ 0 5 8 ( 5 3 6 ih o 6 6m h8503h o ^ 6 3 6 0 16 0 6 5 0 13 0 6 .
0138(55, 5b3cn 3 n h m 5 3& 3n (5 50 633R 3(536 sh 3b 8 33(^ 9 0 5 5 5 6 0 3 0 0 15 ) 10 16 5 3 m 6 5 y 5h o
b5h3lb 3 h 3 P i, 3i c * x ? n h 3 ‘b o b a 3 6 (5 3 0 m 6 o 6 3 J m 6 3 5 30 )1,5 3 6 0 9 5 6 . b 3 3 ( 5 3 6 5 ) 13 3 6 6 , h m 3 0 3 6 -
bb(j b3nh 5g 5 6 5 6 0 5 0 13 6 6 66m h3g 5630 *1 (13536 3)15(50 5, 3 o 6 o 6 'b 3 g 5 65600360
2>3fijnb3&3hn 95 9585606336300 5J301.
6635956635 565580 3(5365h3b 5558605 5566635336390 h3<bobvi)36(i)30fn65. 8(53-
65)13015 653h(j303&ob 65500160156 9535300)136001 556655301)1 36390 ^ 5 9 ^ 3 6 5 3695
803)1536 56 59 55^ 9 5 5 6 556 , (159556 i 3 6,5590580 8(5365)13 y3305“b3 35hd6m6o5h35 95
6601)139 856*635 9581^150936390 585 013 o3 656301606 5595(18365 95 553111530365. 36
5^1559 Ri6b b030on5035<bp33oiob 956590 (jyob 35550ocn‘b3 (65b. 6 .45 ). 3333 -4®C
- 365 -
8006660060 a e ja n a ^ a m a a o
yn63i 06333b 801300 600b 3b3cn 0 1 3 0 1 6 0 b 93(313533^ ^0019300 - 8 - -10°C yn&3D-

60 5 3 8 0 1 ( 1 0 1 5 6 3 3 3 6 ^yob 63636(103 536360363b, ( 1 3 9 5 3 6 ^301 ^ 3 9 3 3 h i ^ 3 h 3 6 b 3*^03636
abmpmg Sm'bhgnQn 0690309360. -20 - -25 °C 1/1633 3333 ^36^01033003 j3>03bob
^ y o b ic r q o b , 5 3 6 b 3 £ )o ih 3 & o o i 013 050 Q ogb36b 5 (^ 3 0 9 3 6 3 6^ 9 3 X P 6 0 3 ^>
P3hm36ob 3056001. 3 3 1 1 3 0 b 5 3 (1 3 0 1 3 63600360 030013(13636 3qjho» 9b(pbQ h3<bnbvi)36-
6300163b (obo6o o0ynq)36oi6 y300i<t 0 ZKBhm 03p 3oo iE)033(i3O3h3On 530mbbn336nb
3ohm63&3o), 3119(13 303(101 93(3300, 33hjob 3056001 53693536300 36305313(193 b^pgn-
-4-6
-6 - 8
-5 -8
Below g o - 8 ' | |- 1 5° j 1-20° [ ^ > 25° S g e
6sb. 6.45 9]b5ihn ^ynb yn63ij,i9dp3m6i. Rihfirn9n 9mou>3b36x3b
n8 ^393^ ^ ^ 366, hm930cni ^333001 ps'bni63&36n 31,33 {>s0mnb:nj3&:> (Larcher & Mair,
1969)
360. 300(1)010, 33 od03&i ooij3^b, hoi0 331130b 153033x0 3010301636^360 835336300

3(1036 395oob30yoiq)30<b3 3(^36300 (*1(503001 93^30(101306 013306363(136366.
3hcno 3oi3x3(50ob 153033X n59030936b^13 533660301 53613633336x0 538016(10101-
60b 363(10 93 h3<bob^)3 6 ^ 30 ^ 3. h3'boblJ36vi)30n6ob 0133^363(1363013 qgihoim b^^j-
^ho 631100139536b bb^poni^ 03(j>3oo h3<bo6&)6v!0coi6ob 9 Jm63 iQnQb^^ho 3j,m-
jjo336ob 63h8oi3 oi6ob b3qod33ob; obo6o boôiubcob 36ihb 833063(3 y 063(16363636,
hri93b3(3 30083^11 b365hd0O3 93 0603363001336 1530003636b 53601390b 93 b3b3mdob
633IH530363 i 3 3i 6 o ^ i, 9 o b33^3m 69363.
i]h^Cm3s6n 815363113360b yo63ob3g 0 o 005 h3goi6 ob 3301031303

bo(3o3ob3g0o h 3 'b o b j)3 6^ 30o 3301,5033606 6 3 h 0 m j0 6 3 93 h3<bobli3 6 ^ 3 0 m 6 o b 3h >
3300 b b 3 3 Q 3 b b p 9m 6o b 3MJ360163 3o 30V K rt36b 0 8 3 ^ 3 , hm 0 yo 6 p 530d c 3m 6 n b 36V
ho 0(5363(133630 30013(19360193 b3< 33b 3h 36(i0339. 6o(503ob3g8o 33m o3(in3hn 3935^3130-
- 366 -
67J636 (n030 g^d 6 Hff)360
ob 3nh3xn 6560505 ob, hm3 016000 95 '5360560 ,5(101305300 (13500163600 00(5030059-

3n 05^60)605(13 8(5365(13360 9330)90)01 600)8336(1563600 9565(^1 ,1,3833(15^3(1005980
9353360b 5580183853365 95 36bo336nb 83056580059 55(195^865 (65b. 6 .46 ). 8389350
656050 50650) 0^360195 ^>3983^59 551503360b ^Bshnb 5530133953365 95 5590630b
63hôcob 956(15 956335, h5(5 8(5365h3b 053350365b 3011153395 95^0563600 55(1383
55953,1)565 h6o(yi, 33o<bni93ho 9063360 (955bp. -10°C-393). 5b3cnn 59085,500 8^0163
h3500163680 (55605530*36001 b36^h(T)2o5360n) 5590630b 01530956 5(5093600 b^li5l|)3-
505 65(180)595360 05353013001 yo83hm3nbb boijpi^bcnb 305qj(iai0oi36o0 3<bh3633cy-
0103505 95 6pob 1503 33(1001980 b5j)35hob g)fTX5fnbo6oi3‘bob 365hb 8mho0: 05585(1000
551503360b 365hob 33333016001 0301019360 95 03(101360 55h559 5hob 95(5390 b o ^ o b -
Annual mean tem perature [°C]
Electrolyte leakage [%] Duration of cooling
6sb. 6.46 3ih133001316 9(336^300 yn63nbip0n b36bn^,3hm6ne56 ,jmp3h56£X}Pi6i 9n dêsbxi

bn3ipo<ib J1hipn361yib dnb3@3noi (Bern 3nc3x 9n. ibi^n 4,30635). 3511136365 bjhienn: 2600 3 -
9p3 em3 n6nh363B C4 3ih(S3cm3636n. 3qjhm 35(n5p bo3sc<Q336'b3 C3 015365(13360 ^5h&mt>36. T
bnaipcnenn 5^^9036^. 35(133365 bjhioin: C4 3|}365h336nb ic ilx ^ U ^ 3280 6-^3 53>^>h5e 30jhci
yn635{,i3d035, 309(13 05033 du365h336nb 2600 3 V ) (Eamshaw et al., 1990)
556, bmpm 030,10 9063360 5h 0633336 3511390b 55369001365b. 3hoi,i)oi305<b8ob 365ho,

006803933113059 5505(1010 558016(10101600 3hoi(53bo, b3pb 36ym6b 8(5365)100 9505693-
65O 3jb^h335C3h (801593 335365(5oob 33(1001900 3JR63) 1560*55^3630. 0601(139 580,5-
m9 , 9 3 X ^ 3 Q0^ 90635553(1030165 3695 55801933533630090) 8(5365113015 o3 goih-
83690, hoi303&obon3ob5(5 9595b5b0501363^05 5jô3m6ob5 95 80*03363600 33(100193-
60O 5363^0531159 95090^0(136300 80165(53030165 56 O30 (50^55, bhgob 8363(13600 365-
ho (ob. 01530 5 .3 .2 ).
935300013560 3(5365h336ob 9o635553do30)6ob 55801838533600 3hoi(530o 596501
80390651130)695 mho (0530956 5*6(55053336300) 330103(503(10 5<booi: 0593(159036015,
ho»8 lbp3ob 90160956 On8500oO 65^36500056 3(10159 o'bhp365 5590630O556 0153-
95(5300 365ho, bmom 53015(15030300 556390b 85^36500156 3(10059 - 55yo63ob5g8o ^01-
03h56vyn65. ,jlioi3o536oO 35050801056 (13500163680, hm3o36o(5 830(^036* 556306000101
- 367 -
9006660060 usrtObTi^ o ^ a rm rto n w a o
Ubhncn 63 3363633006 1693600 91 536011b 0oiit3 0ie>ic bndipc335'b3 Sm'bshen pi pidob yo6-
336n0ig9n i@i31yih3&3(yi 50516^3(10 hm'b3(j3cn 65361(13360b yn63i{)i9dc3m6i (Beck et al., 1982,
1984; Goldstein et al., 1985)
qapKnQfth 50%-m6n
9i'bns636ob ^jSSjhStjoKi
(CDâhn iKJ83ahieohi) ° C
Espeletia atropurpurea 169360, 2850 0 -6.1

3100 0 -8.1
Espeletia schultzii 169360. 3560 0 -10.0
4200 0 -11.2
Dendrosenecio brassica 9oai 536(11, 41 00 3 -10-nb j339mon
Dendrosenecio keniodendron 3on 53601 4200 8 -14
Lobelia teleb'i 3aii ^3601 4500 9 -20-ob j330mcn
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363(10 06301 gm 63lb 3 33 0 9 3 , hm3 8(1333060036 8(5363(13366 638330 36 3 803(5301 g 3 3$-
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0ob 63033(106 6 ih3 mj86 oai. 3b <31^501(1360 (60393506 53yo633, 010130061 93 yo6-
- 368 -
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30nb 6565(^0030 b^qjihn) Sihib n ^ Q brib i 9 5 5g53n 56ob “6 3 3 0 1 J339 365b cn 56 3(10159
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n b i(j36i 3(10193^030016065 g5 “bhgnb 33(1001906 838(50(136530, 556b 5j)30i(i36nan
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yo63nb 6o33h503
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3>"-
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hnb 606110093501606 55“bhgnoi g5 033633606 “bhgnb 83'b03g3Oai. 6ycnb 830130-
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5hb36m6b bsSoihnb 53503nb 3n3gn65h3m6nb oiho ^n3n:
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055(15365 303)101 63,50 6 y50n, 3ngh3 8306013365 6059550956 56 5595595099361
Bycnb b505h55oi Jboi3o036ng56. (5(116630(11(50001 55801633390 33860130 digo6 55801
69361 558,55(10 ^3(1^93606 556y33^5 95 386019036(16 65h3 oi3 oi6 i. h5g 556 38601-
90360 “60395336 6y0nb 5191195093656 ^0038580, 1301010360 g 5 03(1013606 $36-
63hm36n b 056g 3&056.
pibn ^Sjdjdb jhm6njjjp dndgn6^hjm6ib 5g50gn 5^36 n3 8(3365h3363n, hoi3 -
036015 lb 58oihn6 8016336360b 3ohm&368 n 655336b 5h (56 3 i5nh3g) 5(313636, hn6
5580115 85010130b g535b56d5Oi3630n5 dohnoi5g 5g i3tiniXD^n 56 33hog3h3 x n (ih5&-
b3oh5(305, h5(5 oi53oboi535g 6yoob 65^03659 j)5h535b 53006680166. 56301 ôSb,
33oh3303b yoi30 ob5, 3^30136056 6063013560 g 5 “658015(180 8330301010530 b3 315365-
h336o. yn63nb 6n89h503 3(3365hob5ai3ob 3 ooi 3q3hoi 6580805, (15(5 3qjhoi 06^36-
bo3ho5 8nbo ^h56b3oh5(505 g5 651)03605 6ygo6 6585(15501 603(11533606 301(5300165.
03(1013606 j3hjob 801(500365 j 5hnb, 8 ^ 3 6 9 3 6 0 6 5 95 35h5'bnli X n b0i])0i36nb 8393-
559 56h3ai33 bhgnb 3(5365hnb 553oi3hoi6nb 658n8hm3656 (656. 6 .4 9 ).
013365(133606501306, (1018036013 010130001 5h oq35h36n56, “b53oi5ho, 6 x n 65gnb
- 369 -
9 0 3 6 ^ 3 3 5 0 bdrtObTJC? 9 t?aD 3660 n 6^30
Water content
[ gH2O■g-' DW]
[....1 0.31 - 0.40
M l 0.41 - 0.50
H 0 .5 1 -0 .6 0
O 0 61 - 0 .7 0
r ~ i 0.71 - 0.80
S 0.81 - 0 . 9 0
r= i 0.9 1 - 1.00
■ i 101 - 110
H 1-11 - >-20
» «' ‘
6ib. 6.47 ^ynb t>;)Qs bi’bîh'b] Rhododendronjerrugineumi-nb qjmai03&b:> g i (rrjhmj&Sn 6ypnb
3o3b33Cm*nb eij33ncn3&s <bs3cn:>h3n. A cnm3pncn 83960339 giqjihxn; B ^393 ycmh^^bn
CD363h3i03n cnm3Qb b^mcriii; C 3^36^3360 onm3Qnb bigihnb 5^383 <y>yn6x Einipij,^ 0ih^3n.
3hmi536^36n i(^6n36i3b ajihgm&ncn 5i8m3hm6ib (Larcher, 1963c)
♦ 330m
• 1000m
» 1950m
* 2100m
FMAMJ J A S O N D J FMAMJ
Months
6k). 6.48a,b 'bsScnhob {>i6ds3e<,>6i3o 6063^1316 3^36:^3013 6063360b ihm£h3bnh3&ipn J>i9m3hm6i.
a Pinus cembra (jynb Vjg* bb'bcîh^ 93 3nb ^>330011 U35^hic3*1 ^C^3^9n. • <bhp:>bh3cn 63360; 0
1-2 3-ob bnSij'cnb b336n; * rfbmcnh36300 (3hoi3xn 36‘b303Qsh3&ob bibncn) 93^60636x0 b33&n
^ynb 1339s bi'b(p3hob (b30moi. b Picea abies bb3igibb3i bnSijxj'bj. mho33 6sbiVb3 3hx36n
$83363636 6y(Vib 3o3u33C(T'6bb 6063360b 93,bmq)ncob 10%-ni6o g:,'bni63&nb ghmb (Larcher, 1963 b;
Michael, 1967; Tranquillini, 1982)
- 370 -
5 3 6 3 66030 UftrtOUTIÔ 3 6 d&r)6050
ihnb ihi 9ih^m 13030, i h ^ g 99hign ghm. fihgogmQcn j^SQgob'b], ob^

0^^090)3630 o)m3cnb bicrphob bi65hdQn3m6b, bo9i(T>Q3 Q* 6oig:>5ob 5iyn 63i ob
5iQs96y33(i)n qjij^mhjbns, hm0p3i>nu ^(^36^336 b3 01536:11133606 6i3h(j3p36ob ^ 3 -
ifb. n0 195003600, bsgiu dgo3hoi 6nigij,nb yn63i 9 1 ih ihnb cnm3pnb biogihr),
bj 81336^3360 ocfrp360^6. ^ 3 0 1 3ohm6369o (b^bihjôjjob ^35ghi, 0j>p:>o0aii g i
^ 3 -) iWD ^>g6nCb 3O)0m6b 6ig:>bm 3i6 ou>6ib:><bm 5igm 3636b.
0U36ihnb g^b3b&)6i cnm3cnbi g i yn6x°b b*6&hd(Yi3n taqphnb &b3m

on3gocn giO}sh30n 61536^3360 ihm 6 91(5300 gs6igo (i)30;33hi£)h36nb, ;)ihobi 91
5i0m9hm6ob5i6, bi3i503hrr\g 5160159016 oim3(Y>b bn0dn6ob 66331b g i
5161013630(116^ 3i0bhobm6sb. o0ob 006393001, 013 hm5mhm ovn3gob bo0j>3ho33,
01536^3093 n?633b bo6xngob bb3igsbb3:> o6&)6bo3Pi6i, ib 3 0i5igncnig, 20 b9
Incomplete development
of cuticle
Abrasion of wax and cuticular layers

by snow-blast
1'
Reduced resistance to
cuticular transpiration
High atmospheric evaporative demand

during winter (strong radiation,
high leaf temperature, low atmospheric
water vapour pressure, wind)
High water loss
through cuticle
in winter
Limited water uptake (frozen soil,
cold soil, high conductive resistance)
Water content falls below

critical level
J
Desiccation damage
6sb. 6M >c33&3n 611631*1316 33hj6n>6 91536^33630 (bi3onhnb î3m3hm6nb 6ih33shoi33pn qj>j^mh3-

&n (Tranquillini, 1979)
- 371 -
91306^0360 bGtfObTJC? 9£? a n 9 6 6 0 n 6630
bnQiC’cnb om3onb J3q3 3(^6336 b^g^nh'b] 3(1636300 6363cn3&30m6ob 1-15% .

cnm30ob b3qj3hob 6363611030 6365(1800301606 63001363(139 5313(193 6301600383(101
(13500163680 06333b 6939013606 3(10193^03001606 bi93scng 20-30% , 1301533(1 y i
70% 93^3300136^ [7 6 ].
5i6bi])30)h363p b38o8hm363b 63(180139536b 936636090 010139063 9b yn6;j0nb
03360b 3n3h CC>2-ob3 93 C^-b [5390 539133(10163, hnb 539m(5 0(1(313333 9u36bhnb
53lxn3 15393. 815363(133606 b36onj3ob 53601 CCôb 3016(536^3(503 ^ ^ o o ib b , bm-
pm 33656bgnbb - 303630016b. ^ 3 0 1 30(10163680 63(18010^86363 8m9639p33n 63133-
60103(10 3(10103^360, h01930013 9oihob 536633301(136001 633633 301360190 (COo-nb
93p30 j)m6i}36jjhiuniboii6 3(10139). 150,301001503(1 36 33^ 8 0 ih36mh9ip3hn 6 9 5 m-
93(130163 j)39mnb3^363 360oi303,b93ho 8386(1363606 3110100033(13(50363 93 6001938-
6(1363606 3016(536,5(1300 3016360b 6bh9m9 rn6i 9 n. 93636,536300 6(5363(13360 6 33 0 3-
639 3hni6 yo63o63g9o 99511390 93 b3 nhsg 3339936036 qjbojhoicgo03ho b 0130136001
(635., 0101300b m6n).
6.2.3. 9^666^9nb 6^C D 6n6^ 6 n ^ 6 9 f l
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b9oh3g 538019036 6330(1360936 9 3 9(5363(1330016001 93033(130 603935133 539013^301
90i9n, J3o9i; (3601(1360 don3h 3^360363636 ,5369(136, 93 9 3 6 0 0 1 6 8 0 5 3 6 9 3 5 3 6 3 0
bb33 395ocb39ymq33036b (536633301(136001 aim300b 9601606 939935); 3 9 3 8 0 3 6 0 6
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qj3b 336ob , 6 0 39 350 b (560133036063 gi. 33hm 630o 9 o j)hmmh5b6o'b936ob 9 n3h. 93,3-
3336 0 0 , d30O36 9 3 6 3 3 9 3 6 3 9 36 6yoooi 930331130 60 39358 0 336 56 39 0 8903(1 6 3 0 3
90033690(1366 (6 330 1360 6 5369330016380 h 39 g 36 o 9 3 801(5. % h3mg36m6non), h ijj
0 6 33 3b 9 ob 8 6 3 3 3 3 6339360163b qj3b33& ob ^ 3 (1(5 3 8 0 . b 3630 336 56 39 0 b 3016(536-
,5(13(503 6 0 39 358 0 9 36 39 0 3, 3 j(jn3hg36o36 3633(1016390 9o2)hmmh536o<b 0 3 6 n , hnb
53801(5 5(10139363 Fe2+, M n 2+, H 2S , 6300309360, hdob 8 33336 0 93, 0301 So ih n b ,
80186389330 6030103(136360. 53hg3 39ob3, 6 0 39 358 0 6 35(^ 6 0 16 9 39 3 3 h 3 b g 3 6 3 3^301-
,50b 6h363ob (133080. 33656390b 6339360163 5 3 6 6 3 ^ 3 ^ 3 6 0 0 1 g39 3b 3b 0 3O i36 30 0 3
63(1090301(1360360, ainbm 336o 6o3g3536ob 3ai3o b . 66(1303 89063(13013 6 3 50 (136 0 13 6
3(1636390 638300 03 ^3083 603935360 Byoob 90160b 9 m93^ 36ob ghni63(5 j,n 33(1-
5 3 9 3ho36 3bh3633oym q3o0o 3365639001.
6365(^90339 93^360363639 60393636133 b3bo036o36 86019019 53(1333300
9(5363(13360, 635., 63036013360 ^3901<3303360, 63360936 - 3030b 63, ,50(10030, 63(1-
y36o, Taxodium distichum, Nyssa-b 636301&360, 9365hob 915363(13300163, 330936b
8mhob - Nypa jruticans. 633(10 8(5363(106301306 6yonoi 93,5601(133 63^1801395366 909
63808(1013636, (130936063 0(P38n 36 330(1380 050 06333b 9(5363h336ob 93(3133336.
(101501(1(5 63bo, ‘63983^0 93^360363606536 8(5363(13360 0^3336036 303(101 6^3(13,
309(13 6039350b 530n 8hoi6ob3b.
- 372 -
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6 .2 .3 .1. °3:6jl3n3hn B}h(T333D&n Q M \X X > X (\ 9 ^ 6 6 6 ien b 6WC3-

fcPlbMOb
0U36ih3ou> qj3b33&b i;)3Ci i6 i3 h m 6 x n b ^ o ifo n b ;)6ihn, 0:>&hi0 gi6&6ignb

6iiC36m6nbib hb9g;)6 n03 bsicnSn ig&nQn ij3 b fcigibh^bb. hmg^biu
gi6&6ignb Sshunicohn 66331 1-5 kPa-bQ gi6Qi 315331 (3n3mjbm), h^bSnhi^rnhx0
301303015036^(1 - RQ l-t o 3>jn 69361, nhoi33bi b36onJ3nb i0^3h6i^n;)0n fc'bD&n
(3 ioin 3 6 ^ 8 3 6 0 gi5)6 ig !b3 3ghm 6 1^0 3619 ihni6 g i3myig3&30n, 3ngh3 un^mjhm-
3 mjbngi'bs) 9 1 ig36ngi0)nb bnb^Qnb 363h{)3^nt,30n b ^ i^ b n d0O3h 315331. 033b-
33&nb bhgnb 3hm(53bn rfbjTi^gjds (65b. 6 .50 ); 33166619m qj36i3n 3 mb33ghnpn
gj3b336nb &m0R3&n b036i 91 n ^ S j b i g i 3nou>hg36i g i3 1^360010 03^3360. (33b-
336nb 3<ghm 1 6 1 ^ 3 1 6 0m6ij)33oip6(b3 b3 nhig 3ncnihg36i ^mhSnboft^hn &$3pi6-
&ib. 650 bn0n5gnb Snh^pign qubj^&nb

bhgi gs 6niesj,an 3165,61^^ Sih^jAic^hn
66331 (Saglio et al., 1984) 02 partial pressure [kPa]
well-aerated soli Anaerobic wet so il
lAA normal lAA increased
Ethylene low Ethylene high
Normal leaf angles Epinastic leaf angles
ABA levels low ABA level increased

Stomata open Stomata narrow
GA+CK levels
normal in xylem GA+CK levels
reduced In xylem
ACC absent
in xylem ACC accumulates
High rooting density Death of small roots
Bib. 6.51 sp6j>&ipnb 6i 5C3&m6nb 6^30^ SujGshnb ‘bhgb-^snooihjbit] pi qDBjynmSnh^bi'b]

(Bradford & Yang, 1981)
- 373 -
9006660060 bSÔUT)^ 9^ S n 9660 D 6630
I'bihg^ftn (n6^]03bi3p6j)36n) 9 1 mbSnbQirjhn 33h3g3&o.

£P>6j)6ignb 6i$036m6ob 9 m8i£ ) 6nb 16 8obo W130O ihihb36m6ob (^ 6m jb m )
933oib333i9n, ^ 6 0 1^ 31 011013361 i6i3hm 63p go bo Sogiô i^. i>)3h3o6i 03ho mjbngi-
Ijnnb i8m3ihg6ob 518m 0 ^ ^ ^ H536i CP33ngobi g i 3cm6m0ob i>m6(536^hn5ni.
3Oii6m0nb îh6n yn 6(536,^1(501 ihnb 31656190b 6ii036m6nbcn3ob g i8 ib ib n i-
01363011 bn33j>m3n. îh6o himg36m6nai bn5on3'bnhg36i i6b^nbn6nb 33133611, 3010-
P36n g i 30100360b 6n6i biqj3b3hnb 3hmg3j^36n; i 3 ghmb cgmoi0369o 8 3 0 6 0 36 3 6 1
6153360b dgn3hn 333n6hm36i, g i9ibo633630o bhgnb agmhSi (j^nG sb^ns) g i
qgmCTimQ5336i (61b. 6.51 ). 3jh3g36nb 3336h i636n 1*1169361, 3ntjm-|m6glin36n g i
35h3gnb bb3i 30li)hib(jh3;j^3h36o h3g3(5ohg3&oi6 g i 3iaio 36^08360 6i6npm6hn3
001(1536361.
6 .2 .3 .2 . 6iW hR06^ 3^666ênb 6 ^ C D 6 n 6 n b SflhFlbD&an
3165619001 pihnb 60191580 8(5361(13360 I(j'3mi536g36ni61 516030011(13636 033b-

336b 91 j,igihR36m 6 3bmgmg o3 330cnb333i3n, hmg3bi(5 333dpmcn b i5 i6 5 3 6 m
lbm336nb 3 np36i.
91656190b 6i^Q36n6nbig3n 0336^03110 8 36 336 1 3gj,m3ih3rn6b n3i 8 n, hm3
i6i3hm6nmcbnb ghmb ih n3i^36b i0im3mQg33nghmf)36 i<bnb (AD H ) him g36m 6i.
18 13 3 ghmb, hi3g36r»83 buoinb 5i 63i 30m6i 9 o 51(1910^86361 (5090b 8 3li>i6mpn'b-
8n. bn3n6gnb ib ig ^ ib h g i o6go3og369o, 93bi6i3obn 536360b 6 i i J6 n336nb
8 39 35 19 bn6cn3'bnhg36i 20-3g3 i6i3hm6nm‘bnb Sm go ^^n go , hm3g 36015 hm^nnhu
n‘bm36'bn336n (530016 3oh3i 6g 30 (815., i3hm630O A D H g i qjmbcgiôb f,ig i3 ^p6 n
36^08360) 36bo336b [88]. bi6m0mm 3hmg3jîg 3^15333006361 ih i 3m 86>30i3n
3on6mpo, ih i3 3 g hdob 8 3131, 802,080b 16 31800b 89131. ADH-ob ij^n3m6nb
5 i6 b i‘b(j'3hocn SoSrnjboob 3ohm6369o 83od036i i 3 i 013 o3 3(536ihob 3 1 6 5 6 1 9 1 ^
6 ij)036fTi6nbig8n 85hd6m6oihm6ob himg36m6ho3ig gibiboicn36i (61b. 6 .52 ).
giG^&ignb 6i^036m6nbig3o 8mhqjm0m5o3ho 3 3 5 3 3 6 1 3g5m8ih3m6b Siohnb
5i3îh n jbm3o03&ob 33j36i 3o, hm8036bi(5 ifooi qjihcnm 35h3g9mhobo b o b ^ S i
{ i j h j 6jndi)\ 81010 b1831036001 S ^ b ih o b 8o6ob'b3gi 61600360 i6 3g o i6 3i3h b
qg3b336bi g i 6yig9 o Ridoh30 bb3i mh5i6m3&b (61b. 6 .53). 313(1001 j>ih5ig
3lbh36330ymc3O0 qj3b336b 833d0oioi 3i6 56ig o b 5ih3cn 5i8mymqji g i h3g3jynnb
36ihob 0jm63 b;)6b^hi^36nb 5i3363630ym cgi, 815., Fe2+-ob Fe-III mjbogig 51(1-
g ij96i. 3i3hob 8o6mg36ib b3pb 36ym6b i5h3oi33 ^3833(11^3(130° 6^19^36^0.
îm6ob 81536^3011 gj3b336ob 3ih36jn8ob 20-60% 3^1301 33h3g8mhnb bo3h(5336b,
8i 9 o6, hmg3bi(5 School iih 5 ig 3'bh36330ymqjO0 6oigi58o 93bi6i3obo bn3h(533-
6o 8bmpmg 10% ip 6 33b [37]. 516^^3011136001 8ôghm g i 0^3(103 6 m g i5 (b3
8(5361(13360 030011(13636 qj3b33&ob ^339130(130 bob^38ib, gi^6mho0 ig5O0369n -
(P3hm36ogi6 5i3m ‘bhgo0 6y0ob qj3b336ob bob(j33ib (815., 1030b b3, ^nhnqjn,
83hyi6o, 003160, A cer rubum, Eucaliptus-nb bib3m&36o g i bb 3igib b3i 60631^3160.
hm5mh(5 3^ (13 8 19 3110 b33i5oi0o'bi(5oob 815100010, 83od036i 3m3oy3i6moi 8165(13-
60b 0n3h b33(5oi03ho bib36oijo qj3b336ob ( ttjjd ^ m q jm h jb n ) 5i6300iih 36i 16
îm6368o Cupressaceae- obi g i Taxodiaceae- ob 6y0ob ^33001 iSmbhgogo o)3b33-
60.
- 374 -
100
• Oryza
^ Laduca
80
Seneclo vulgaris
S.rupestris
Carsx arenaria
| 60 63 S. aquaticus
& V ^ V ic ia Caltha palustfis - ►
g 40
L> Ranunculus fiammula
Juncus effusus
©
£L Glyceria maxima if
20 Iris pseudacorus
Pisum \
0 --------- 1 r --------1---------- 1---------- 1— I
20 40 60 80 100 -102 -10' 0 +101 +102 +103
Ethanolcontent of seeds Changeinactivityof alcohol

llunol-0'1 DM ) dehydrogenase
6 ab. 6 5 2 a ,b 6ypnb 6if}6 b nb b]6l)n^3hm6i p i 3^6{,6ignb 6iiC ]6m6s (Crawford, 1977; McManmon
& Crawford, 1978)
6«b. 653 Nissa aquatica-b

(P3hmb:> gi qobjnb i]hb(jnnb
bnbooQi 84 gsB&bignb mh3ij,n
pnqn'bn^hn {,'b>: j^Smbqnhrn-
9^6 hn'bmbo)ohm3n pi <33^33-
6ngb6 ij)9mbq}3hrn3 n (Hook & zone
Scholtens, 1978) roots
6.2.4. 5 3 X 3 :)
{)3bC3nb b^h^bn 6 i h 3 mnJ36 o6 i 9 i 9 n6 , h m g o b iu 9 u 3 6 b h 3 3 6 n & b 6 n u g n i6 83-

b:>6 * 9 nb o i3h 9m gn 6:>9n £)Q SQ bn Q ih Q m bbSn b h b j 6 x n B ygn b 6 ^ 0 3 6 m6:>b: 6 n ig > -
{,n b b n 9 9 h iQ 3, d g r^ h n im h o r j0 3 6 i, B ycnb mb 9 m b]hn
69 :> 9 ihr\Q ni 6 b^ b b ^ h b ^ Sn ,
6 n iQ iftn b y > y n 6 3 b g i > 6 ^ 3 0 1 3 3 3n 6h m ^ h ^ ^ n b 9 Jm 6 Q 6 0 ^ 5 3 6 0 , h m 0 c 36 n (j 
9 ^ 3 3 6 q j3 b 3 3 6 n b b n 3 h u x 6 ^ 6 3 0 0 1 ^ 3 6 ^ , B y p n b 0 3 6 m 3 i b 9 } s 9 nb j>i 9 fn 0 6 3 3 3 3 6
B ycnb 6 iQ i6 b n b Shm ^h^bx 9 3 6 3 3 6 ib 5sh 30 m 3n h m 636n b :>g9n . bb 3i b^h^bxn
g ^ 3nhoi336nb5i 6 5^6^3^336001 53^c3nb bvi»h3bn £)h^0m3;>6 9u36bh3363n 3156*9
- 375 -
0 0 0 6 ^ 0 0 6 0 U&tfOUT)^ 9^ a n 9^ A 0n 6^30
ih 6ih3m oj3636i - 050 cni6gicni6m6ncn 3i (J X fT,^ ) 6^ 3nJ33g36nb bi65hdQ03m6nb

6 i‘bhgibou>6 ^hcjiiQ dpn3h93&i. b6mh3g i 9 o^m9, 63^03^ b^h^bnb Snhmb^bSn
ghmnb qji^mhb 360^36^ gogo 36o93630m6i.
6 .2 .4 .1. 6 3 X 3 ^ hn&nhl3 M h D b -o JW lh n
53^03^ ihnb 6ip3j3bncn (j>ihn6n ^homgo, hri30ob ghmbiij 6y0ob 933 (333Cm£>i
6niQi53n 08936^9 31538^, hm8 3(536ih 336b Byigo di0Oi6 ij)0g3&icn. bQohig, 6^5-
hi8 ihi ym33ccn3nb, 6oigij>nb bo39 hi03 gi»,i39nh3630ni h>3hnb bn39 hi 03 bcni6
g i Sbnb 8iQ>i0 hignii50ibou>6, hi(j nB333b imhoij036ob dpn^h 3i^36ib. 9 bmomg
6b03j3&nb bn86nh3 ih 6 r»96 i 3b ihng30m&ib: 130311 3m0ih30O h3^0016360 6^03-
j3&ncn (pihn&ni, 9i5hi9 nbn6n ih ihnb6 ihog3036o, higftiB ij 33i3m^hi6b3nhi^n-
nb n5^36bn3m6ii5 gi6i0ni; 99hipn h353m636nb 9u36ih30ii (331)360 ic'6 3336 33(1-
0^636^30^9 j)36ni6 911336b 6i ^ i g x 3&nb 93(bm60i g 6^60^53630 igj,o0360o g i
j,h36^nb 6y036nb ib0mb (9i&., 6i 03h3on iiy33&n, gs6pm6'b3 6i 6 ci 63&3c n £>DRjm-
3i6n qjmh0ii3n36n).
bnddhspj shnb 6^ p jjjd n b bndynhnbs g s ds(?i>pn smhonjpjbnb jhcnm bpnjn
dm jdjgjdnb djgj& n. 39hip h35nm6369n 63bC3n^ 33homg3pri6i 9^> bi65hd0n3m 6i
ob3cnni, hm3 6y0nb 6003110 imhoij036i 33^0^ 6i03j36nb 60n3h ji9 'b 3. i 9 9 36 -
cnb333b3n bij03 &3iJ3b ih n g x Jj0o9iâii6, 330ng3hn jjgnSi^nbcr^nb ^n, 3nhnjncn,
g i 9ibibnicn3630Oi 6i 03j3&nb bnîh63, hm030ou ^("38 ^36 ^ imhcnj036ib [2 4 9 ],
b8303Oinb tb3g i3nhnb 0iib0m3&noi 1 /3~*b3 ig^non ij3b 63n8nb g3<3nun^b, bd
qjihcnm&nb 6ib33ihn (12% -0 g 3) 08936^9 39hi0nb, hm3 6i03j3&nb Bonjhn j i 9 n
250 93 gi6i0Oi g i ig5n0n i ^ b 31^36(303110 imhoij036ob 3bmpmg 3hcn 930101-
b3gb (6ib. 6 .54). 39hi0o h3j,nm63&nb gngo qjihcnrr^&n d33b Rhgo0m3oi g i b i 3-
bh3cn 5i63g3&nb 150 g i 300 9mhob (6ib. 6 .55) 9ipi09cnnb Jihni6 o)3hgm63-
6 ^ 3. 'bp336ngi6 g i9 mh36x h35nm6369n S^ n gjh o ^0n3i^n £,igignb n93niau>g i6
33hnmg30ig 99hi0, b39oihog3p i0n3i^9n g i 6m0mb, ih o p x J>0n3i^9n, hm9-
0nbcD3nbii5 gi3ibibniO)3630ni 6 3i0 3i g i 6nigij>nb g i S g ^ & i (6ib. 6 .5 6 ).
B0n3hn 6i03j3&nbi g i imhcnj036nb oii6i<3ihgrn6i 9bm0m0 “bm îgig 903010-
0136b 3i 6nî^nb 339ng3hm6ibi 013 ihng30m6ib3 9i336ih336nbcn3ob 6y0ooi 3<bh3&-
330ym gib 5i6bii30ih3bocn gogn 96o93630m6i ij3 b 3353iiibnob 33hong9n. B0nb
J)i69i 30rn6i 9 o >)0n0i^nb q3ihgm6noiig 3^9093bn 013 ihog30n biboioinb 93b>-
q3ib360ig 93od036i 5i 9m3ny36moi 93^ 3mhm0fn5 O3hn big53h36nb 0m6i(33936n -
oi3h0m3033om2,hi036n [75,120, 265] (6ib. 6 .57). ^3933hi^3h30n 9h3go 6 3330 0 63-
6i, hmj,mhij 60ob 5i 69i 30m6i 9n i^ 9mbqj3hnib imhoijQ36nb di0nb 9 ^ 3 3 6 3 6 3 0 0 .
60ob nb ghm, hmg3biij 6i03j36nb 9h3gn 3qjhm g i6 g i ihnb 6ib 0i 63^ X n- 3ngh3
- ^3033hij)3hobi, 9u36ih33&nb 393^3bm6nboi3nb (hm3036n(3 33)1 1^ 36 36 36 5^136-
ôb 6y03&b i6 ih oh6y336rii6) 63i03fib 33hnmgb 6 n96i 3b. 3b 3ho6(3o3o 93nd03&i
J,i0my363630n nymb nb30i bigj)3h36'b3, bigi(3 ih o'bm036i mhoijc3bi g i 9 bob
higniuni. 0 139 ^ , ib3oin i 0n9ignij,hi936ob j>i9 my3636i y3306^>6 ih 93nd036i,
9ij,., bn9i(p03‘b3 g i 8m|)ng36x n imhoij036ob 9m 9i^36i g i 3cnh36ooi 339-
I536ih3m6ibi g i 93iiy333'T>6i3n, 5i9rriny3636i b b 3igib b3i jho^3ho39n, hm9036ou
- 376 -
536366030 U&63U'3C^0 ^d6D 6360
6>b. 6.54 p;)g:>3n6nb 39hi(Y> h^mG^bn. jQibnajnj^ni giqndBj&xni 6pn]hn 6i(v)£)&ftbi gi

h^&nmtyjftnb 3m,);)6(yr)hn 33i3m^hi6b3nhiunnb oiiSiqjihgmdi'bj (P/ET^,): bhnfrxyi icn0i^n - 0.3-
b] JSjSmon; bjSnihngxn - 0.SO.6; b^Jjdrtg^hn - 0.6-0.9; S^Sng^hn - 0.9-1.5; dpnjh S^Sng^hn -
I S-'bo \>33noi (UNESCO, 1979; Box & Meentemeyer, 1991)
90
Arctic tundra
80 Cold desert
ao 6»b. 635 Bpn^hn 6so3b-

6nb 5 i 6 i 6 n c 3 6 i ^ itije j& n b
70"
3n b 3 9 3 0 0 1 g i 3fn(J06(}n]hn
so 33i3mv!)hi5b3nhiurL> (ET.,,,
5iid>5ihn33&3eni <® a'1)
gi giSibibftiOTjSixn
o BOO lobo isEo 2000 iimi
SuoBihjx0 qjmhaujnjbn
Annual precipitation [mm] (Schultz, 1988)
- 377 -
aoae^fnaabn bfttfrouac? a ^ a n a a A a n M a o
<# jgOPoW
<250- < , 0 0 era*'*’0 aeas^_
90 " venn##*'0"
6>b. 656 j£nd*^nb, 9u 36ih 3x m 6nb:> P* 6 r u e ^ n b (J3C*'CD&4 s h n g x m&n*J & h ie°3t^yib dnb]p3rK n.
6;>c3j3&nbi e* 3f^J36(yohn ]3i 3m^hs6b3nhi(jnnb 0h]K)6nb {^9^^3300 ip6n96b3b bi'bp>3ihb shnpx
g s j ] 9ng]h jc o 9 ij)3 6 b 9mhnb. (»i0mb>3O3on 33hm3nbiOt3r>b fihgnpm-
9bl>b303on9^6 bi9bh30i-iC,9fTibi3C30u>3p] ^>b 3nnb p>£>c'md:>9p 3 (Vtysotskov & Morozov, cit. Walter
(1968) 9rilyje3noi); j 33( » b^Kscnn: i(p9mbi303O > eib i 3C 3cnnb 3hmojn(yi Wig. id^h n yib ih jh rvj& n b o sn b
300 ieb6 1500 aieo (Walter, 1968)
- 378 -
636066030 U6 (*)0b'3C?0 ^ d 6 f)(f.060
a b d e
K isa n g a n i 14 15 m ) 2 5 13 ° 10 4 2 S a n F r a n c is c o (16m ) 13.6° 5 6 1
f 199
g 178
O d e s s a (70m ) 9.9° 3 9 2 A n k a r a (6 9 5 m ) 11 .7 ° 3 4 1 C a ir o (2 0 m ) 20.8° 26
A b e r y s t w y t h (Am) 9 .5° 125£ A r c h a n g e ls k (10m) 0.4° 4 66

B a r r o w 14 m ) T2.2° 104
6 *. 6.57
bbjigibb3i
h3jnm6nb
5Qn3ipn>
5hs93&n
p i q j 3 d 6 3 6 x n:> 6 n ig i& n b ^ 36 n i6 m & n b g>j>'bm93 ^>bi> g i 6 i 0 3 j 3 6 n b J ,i 6 i 6 n 0 3 6 n b i 6 i 0 -

n 'b'b], ^^mqjn'bnmQrn^n^h i b S j j ^ S n & 3 i0 3 ( ib 6 i 3 0 3 6 n b b ih n b b n b 0 i b i g 5 3 6 i g & ih 3 -
an q p j^ m h o & n b u m g 6 i ih ih n b b ij)0 ih n b n . 0131101 3 1 5 3 6 ^ 3 J ,3 nfi3363<!)bl b i g g i
hm gnb 6 i h 9 m nj 35 i 5 3 i C 3 nb 5 i 6 9 ^ 3 1 1 ( 1 0 1 3 1 . 0 1 3 h i 9 g 3 6 i g ih n b 6 5 3 6 ^ 3 o ii3(ib n
5 i3 h (3 3 P 3 6 n b ig & n p ^ bn 9 9 h i0 n cn g i ^ n h o i x n (bn00 hiQ no) g i ^ 3 n h a i 3 n b n 6 g 3 jb n
RDI [ 96 ]), 0 3n d 0 3& i g i3 ig & n 6 m o i i h b 3& x n B ypnai 5 i5 3 h 3 & n b g3qjni5n^nb
(WSD^j^) 93gih3&no> i0 3ih i03^hnb ih n ^ n ix 9m6nj30oii6 ( W S D ^ n^).
RDI = W S D ^ / WSDihn0) ( 6 .3 )
g^jn h cn jn b ^hn^nj ,jq Bjh^npsg, n9nb 9nb3g3noi, 013 hi 3hm&030ib 306(101301,

03ndp36i 9 n3nR6nmcn 531031161516 5101*163330(1 3nh330n 03b i9R633n 6 n9636n g i
^>6^30133 qj36jun3hn gih(p3333bnb gibi6ynbn. 6y0noi 6i33h3&nb g30]nun^nb
6ii53pig 93ndc3&i 5i9m3ny;)6moi 6ycnb 6ioi6bnb nb3oin ^0103(^636^3611, hm^nh-
nuii - 5ycnb qjihgm&noin 330U330m&i, 3pi,536151131111 mb9mb3hn 3(^3615(110(1 g i
6ycnb 3m^36unicn (03^1313930(1 ,jhi6b3nhn5nnb 3nhm6369n). pibipn g3^n9iQ3hn
9m6iy30n (RDI 15(130133 5i0m nbi^3bi 3hrn536l!)3&9n) 0n3Oina>3&b 0U36ihnb
306093630^ gi^)3nhoi3i‘b3. gi^ n h o ^n b ig5d 0 b i g i ghrnt^ g i9ming36x n 5i6bb-
3i33&i6n 93nd03&i 919506936 3hoin bib3m&nb bb3igibb3i 11690309^3, 915361(133-
0m6nboi3db g i0 ib ib m o i36 x n 91^311(1013116 01131^363(13616(1 9 3 1^ 0 3 6 1 j)i6nbi'b(n.
- 379 -
8oa6 6 6 3 3 an a ^ a r m tfa m .b a o
3hmb 505 orj n0 h35om6obo*3nb g505b5bo5oi3&3go b^bjmftnb RDI-nb 0m65(5303-

6ngi6 (isbh. 6.6). 6 0 5 9 5 5 3 0 6ygob 05h55nbi 9 5 igôQbiOymqjx'bD imhovJc^bnb
3ohm&3&ob U35C3&igm6ib gngn 0 6 0 3 3 6 x ^ 6 5 5 jjb 95, 5^3956 550m3go65h3,
bn33hig3 ihnb d t j^ i h Q c n i 556i6ng3&nbi 9 5 6i 3^bXD^f|b 3ih35x n^0^ X n 6^D*
3 m qjij^mho.
Ubhnpn 6.6 bn99hspnb b^h^bn ({p3fnb:>£)0n {pSmShmbnb qjshgmbnon o6g3jbdo RDI %) 9uo6ih33<vn-
6ob bb3igibb3i ^n33n (083115010 0ijbnd^9n) (Larcher, 1973b; Bannister, 1976; Sveshnikova, 1979;
Bobrovskaya, 1985)
a: qjhRbncjdSn 9fH5393Cni 08305010 SiJxtdiQ^hn 9m6:>u3d36n
3e36jh3r>*5 6 36 36 ^ 3 !Jnhm6)5an 5yc*nb

9(536ih3x<T>&<^> iifi3n
&ij3h36nb e30jniyy>nb b i3 > c m
' '' 0/ 3ijbn3odn (WSDao)
(% WSDcrit -gi6 5-10%
ei'bn5636nbjcn3nb)
35hip063j63 &3f5j65ho bnSShxyib 0o0nl»(,b3 80 • 85
b9xo'i9Di'bc,3nb3oh30i3o (Sijjjnbn) Rhpncm^cinb 0080^3 40 - 70
6ib33higd3fij5ihn b03coii93b‘b(p3nb3nh3cnn 90 - 105
Ericaceae-Z>nb&J5 3355 :>^>6vyiyib m*,3>6nb 5>y>- 10 - 15 (88) a
6]fij6iho b>5i5nhm
<3mcnrlp9l53D6n 63360 95 6^360 RhgfXynô 33hm3b 10 - 40 (50)

:)33 (jynb 6iC5bfn3636n Khpncn^cn 6 - 25 (50)
Jb3hmoi3h9x n 5160^5^3606 mho366n- 33hm5i 40 - 85
560 65051x1136360
9p30mb 35611 6(j36ih33pm6i [335lihx?3hn 33hm3i 20 - 40 (75)

b& )3nb 3(336ih 336n 1336^ ^ 3^° 50 - 90 (108)
U36jyi^C3hn Vbni 60 - 80
3g566mb 63^360 Q5 63360 ^>hi>ymh39nb 3Qi>56m 30 - 50
39566111b 6ib33hs9&3R;)36n g> 65oh6>c- l536lihx?Dhn 50 - 65
sb3&n
6.2.4 .2. qD6jl5rDhfl eihC'3D3D6f1 Qb G^cbn^6D6i6fl
Bygnb 6i i 03&m&5 06333b ^jhogob Omu^gmbob 333i5oh36ib, 3 obo 63360b jjfn6-

l536^h5(jnob 0m 9i^36ib 9 5 3hm^m3g icb0nb 55]6ycm 365b. 5h 5hb3<bm6b ob^cno bi-
boism^bgm 3hmi53bo, hm33c‘bDl5 5h 0mj0393bg3b Bypnb 3m^36uo5Cob 3 3 0 ^ 3 6 5 .
6 i b . 6 .5 8 -^ 3 0 mi5303 On;i D 3h3gnb q }3 6 ju o 3 & o b 0 0 13 9 0 I1050, hm0Q36ni5 0 ^ 3 0 3 6 0 5 6
B y g o b 3m^36ôi0ob 3ihg6obib. O0g365g, h50g36igi(3 5 3 5 9 3 0 b b ^ b o 3m0mom-
- 380 -
b'36a&A030 bSffjOUDÔ 3 ^ 6 0 6 3 5 0
a____________
CeH growth inhibition
Turgor-regulated
PeVsomt reduction
ion transport
Nitrate reductase Inhibition
Osmoregulation
Abseisic add increase
Sugar accumulation
C/tokinine decrease
Proline
Stomata ctosun
Photosynthesis inhibition Glycine betaine
Photooxidative disturbance Synthesis of

8l/ess proieins
Os motea accumulation
10' 10’ 10s 10®

Water potential decrease
Time |s]
5sb. 658 a,b brt93hi(Vib b^h^bnb 'bjdrrja^gQi'i 8uo6*h33&'b3. a jjh^jgnb ojjbjunnb b36bn£jhm6i gi
jjhjgx11 ShnHjjbjftn 6ycnb 93030iyi(jnb 3nhrn6369n. 3rT)hntMTi6^«Q3hn bi'bj&n i(n6n36i336 n8
bigib iC'r'5f'9536i 6(536ih3cni 3j)iqjnm h3ifoni; 6y33^npn 33h^nj,ip3hn bibn ^3363^
&ij,33&nb gib3h3ib (Hsiao, 1973; Bradford & Hsiao, 1982). b ghmSn 5i9mbi^xn 75^36Xn hDi}-
503611 ijhjmhnb b3CnCD&i<bO (WynJons & Pritchard, 1989). 9^36ihnb h3ijun3&n 3mc3iX 3^ gt'G'foj
nb. Bray (1993)
]nph^c 3i536*h3360n 3ooi*hg3&* oi*6g*oi*6m6ooi, 3hoi0*63oinb 0oi0g336oi 8013-

0363611b oi*60o0g33hoi6*0o i*h5*g 05330136* 0obg*0o 03b*6*0nbn <ZD6Jljn3&nb
cg*hgoi6ooio 05hd6oi6n*hm6*.
3nh33pn g* y 3 3 C ^ 3 85hd6m6nih3 h3yj(50i 6ygnb 6*j)036oi6ob 8o0*hot *hob
j^hjjmhnb 9*1538* g* *8001 5*001633300 bhgnb 3hoi(33b36ob (5*6b*i,30ih6ooi br»3h-
15X n bhgnb) 5*b*65hd0O336*. y 3 x ^ 3 *gh33QO 5*3036* 030600636* (500360b
33^*6^100^0^3 g* *0n6rn09*336ob bo6oi3<b<b3. 6yonb 6*i036m6ob 5*801 3©bo036b
Smhnb 9330*^3 3qjhm 0011153636* 6o^h*^)h3g3J^*<b*. 3*hymqgocnn 6*o*6bob b*6-
601503 5*3Q36ob*b 6n^h*^h3g3J^*'bnb *^ 0 30 16 1 31330* 20%-ooi, bm0m 63*03°^
5*6hdx3&nb*b - 50% [9 8 ], bf>oih3g *0o^m0 , b*b3jncn 5*6*9010303113630 0(536*113-
3680 53*03ob 33(1001980 6n^h*^36nb 030(5330016* 0 * ^ X rn^,b. s'bmôb qj0jb*(500b
3hni(53bo 3*hj)fnb636nb qj3b3ob 5019(13680 *5h30i33 dco3h 05hd6m6o*h3* 53*0-
3nb*g0n, d*5h*0 bo06om‘bn VA-8o50iho'b*boi*6 *0 3qj3j^b *h6o036b. (300360b
3>!i*6fnoncb0ob 03'b(p3g 3;> 06333b g*yoiqjooi bhgob 3(101(5303606 8 3 6 3 0 3 6 *^ 3333
8(50(13 53*03<ib 30(10163680 630036* 8olyn‘b36o. *0 ghmb 8-09*13* y330*"b3 05hd6oi-
60*113*. 83001^33680 (8,53(10*63606 5*63ooi*h36ob ghmb) 030608636* Jhoi0mboi03-
6nb *6010*00360, 0 3 ^ *g *‘bob* g* *6*q)*<bob g*hp33336o, ob3, hm0 63*03ob Unhm-
63680 8^3ho*636ob qj3hl5o03hoi6* 3(338* [ 164].
33(1 50033 b3b^n &3*03ob 30(10163680 0 ^ 3 3 6 8 0 5*hoi,yi6oiog36ob g*3gob
03g36*g bo6oi3<bnhg36* *6b(3o'bo6ob 83*3360 g* obo6o, (101501(1(5 b*bo5&*0o 6*3(1-
01360 ("o33b3nb bo56*036o"), ^h*6b3mhôhg36* 0(536*hob bb3*g*bb3* 6i6oQ0o,
b*g*(3 8h*3*o6*oh 3qj3J^b 0633336 (6*b. 6 .59). <3010103680 obo6o 5*6*30(10163636
6153360b g*b;jh3*b. 033b 33600 g* <3010103680 53*03001 5*801633300 301(180163606
bn6oi3'bo g* 8*oio 930950180 5*3(1(53036* 06333b *bo0o0*»j36nb 5*9*6*60036*6,
bhgob 06,53660301606 oi*6*qj*hgoi6* 8o6nb<b3g* g * 8060^)338* 6*6oo36b 8oiho6
0(33036*, 6*h0noj0636* g*0*b*bo*oi3630O 001(10301001503110 15300036360 Q*. hov-
50ih(5 63bo. d*0o*6 *gh3 n6y36* (133(10193^0300 3(101(536360. 013 5 3 *0 3 * gogb*6b
5*5hd3og*, 03*5*6oi0o(b88o 8*£)0oi6b j»*^*6oi03ho 3hoi(53b36ob 6o0o, g*63h36nb
3hoi(53b36o Rj*hg36* g * *0*5013*60 <301010360 930010036*.
y33Q* 3b 3hoi(53bo 53*03ob ghoi30*g 036y33ôb 030cnb333*8o 80100*6*9 83-
- 381 -
900 66^00 60 b 6 r t O b ^ a ^ a n a ^ a m a a o
D ro u g h t s t r e s s
I
R e g u la tio n b y
A b s a s ic a d d E th y le n e
( in c re a se ) ( in c r e a s e )
Stomatal closure
Compatible
O © Leaf ab scissio n
Increase in
osmotica
Debydrines
© © stem diam eter
Senescence © © S e n e sc e n c e
S ig n ific a n c e for th e plan t:
► T ra n sp ira tio n d e c r e a s e d
M o re w a t e r s t o r a g e
W ate r tra n slo c a tio n
fob. 6.59 5mh9m6 i 03hn J>i9 m9hni6nb ■ In c r e a se d d e s ic c a tio n to le r a n c e

b^h^Lxib phmb: (+) hQiJ^ni nthpa&s; (-) h^pjiyii • W ithdraw al to s u r v iv in g o r g a n s
aynhejfti (Tietz & Tietz, 1982) a n d d o rm a n t s t a g e s
lril5D3i 9ni. eob^h^jijn^Qn gs 91(330010 h31;)(30360 009906111301636 3hcn9 i 63cnnb

j,33h9n-j,33hg, hi(3 tphftig fii6b i 3 9m3036ob phm3n 5 16 3 0 0 1^ 3 6 1^ 3 Q ij^ n f^ d o -
bib (ob. 61b. 6 .58). ^h^mhob 9i(539iboii6 3110119 610(1013361 m bQ m h^^ci^m hxn
93^160^3360. bn6oi3'bob, bibi036Qnb 5ih g ij96ob 9 1 ^hiSbQm^njoob 03833016001
33h39nb b ^ j ^ f ^ b i 9 1 (jn^m'bmpOn 5,110139361 9i 6ip 3mQ3j)303ho 6i3hcn36o
(bb6i9o 6ib3nh6yip6i936n 91 ibmôb 01(1516300 6131101360). 3b 3hmu3bn b3Qb
36ym6b Bycob 93b30ib (jolyrt30i'b9i9n, 33h3gob 9m(33pm6nb bî6npn<bn3nib 91
I0(5oh36b ^h^mhnb 3ihg6ib 93'boiqjo00n (61b. 6 .60). i 9ob J,i9 m 6 15 336 0 3qjhn
Osmotic adjustment
Leaves Roots
I
M a in te n a n c e of M a in te n a n c e o r
le a f turgor root tu r g o r
I
— L e a f g ro w th P h o to sy n th e sis T ra n sp ira tio n R o o t g ro w th
. M o re ■ • M o re so lu b le R e q u ir e s • E nhances
d ry m a tte r c a rb o h y d ra te s w a te r s u p p ly w a te r u p ta k e
\ D e la y s p ro d u ctio n D e la y s s lo w d o w n
of a b stisic a d d in cyto k in in e s y n t h e s i s
6bb. 6.60 qj3b3o6bi q* qjrooipobSn mb0 mbnb U3Qn(v)dncji 5^80153330° D<33iin (Turner, 1986)
- 382 -
536066030 U6 (?i3b7)C?n ,3^dS(T)ro360
5bb. 6.61 Pinus strobus-nb 6nf>3nb ^hfiQn. Irjhiatt: 6yQnoi 5^53*13630 Qg^mSih^mbbSn; 1)3391
b^hiom: 9nbn 63900 6m6nb 55%-ncn 5i6m8hm6nb 838936 (Parker, 1952)
6>b. 6.6S 3386hi63&3n (jbn3m3i6n 94 unom3i6n j)m33m636^36nb

03bid[yn mhn35>ji(jni g^nphiûjnnb bihnbbnb 3nb393noi: ByQncn
6i30^36x 3g&md:>h3m6i9n (3ih|jb6n3) 9s 933n£hnh3&x
3pj,m3ih3m6i3n (8^33603) (Bewley & Krochko, 1982)
0 O 0 b i6 b 11 6 3 6 0 1 6 fcibbB n pn, h i^3 (j36i h 336o b Qn^h 6 ib 8 n h n h -

3 'b h 3 6 3 3 0 y m q jb
316 50 b 3]o i3 n b o 6 ib 69 n.
3 3 h i0 h3j,nm 6 ]
^j)6m6nb Shm^bnb Qi6y36nbib jjh^g^bn 3(10181630^ 338m 3^n036oi6 (61b.
6.61). 3311300b ^c^jhib^h^^h^bn, 3mi530m&ob 336(5(1(1361^ 8 3b i6 i3o b ig , n^Q^b-
636(116 g i g^cgmhQnhg^bniS. i 9 3hmiJ3boii6 g i i i 33nh36x n 33h3g8fTilinbob 303-
336^3606 3(^61536,5(11(5006 îbhgi (516^3301(136001 nm63&ob) 933Ci'b3 3qjhm 83-
^ ig 3nj03g36b q}mvj)mbn5oi3(bnb 33m hig 3hFH53b3b‘b3 g i 3 nj>mjm6g h n x 636-
0^ 31^3. bibmpmm cgib^n , 33h3gnb g iC D ^ S e D - I536^hic3hn 3i33r>i0o nymcgi
8(11310 3i^ ih i 3ij,3mpig, j0mhm3Qiblj36nb ,50 0 130 ^36 0 g i 3n^mjm6ghn36nb
3^6,5360 33b 063(136016 g i 333g36 08036016, 60(101306 51(160 063(115361 g i 3m-
pnhn6mbm336n 080361. i 3 cgi'bnb 10(1330 bîgniSn 'bmftr^hcn 8(5361(136 (516-
6133011136001 jjiSmBhmbnbigQn Sg^hig 6ib3m636b g i 013603606 Ri6ibib36b)
Sycncn b30ibgi 5133(13606 83801633313(1 833800101 gihc>33336ob 1 ^ 5 3 6 1 . 33(13-
gnb 333g5m 3 &3gb 6933,56 6om8336hi636o6 g i‘bni636r»6 bihn6bn (61b. 6. 62)
g i (1331(11(5006 36ihn g33nghivi>iunn6 333036- 60018386(1163606 3hcn3i63^'b3
336 3336 1 0 i 0 i3 q i 6i6m0mni0, 063336 33(13006 60330006.
- 383 -
90a6^ 03&n b&6 0 li7)C? 8^ & n 9^(f.0n 6^3 l)
6 .2.4 .3. 6 ^ h fiD 6 ^ 63iC 3flb 3n h n 6 353fl
3(536ilinb 36ihb 55950,5560^ bo83h5onb 33(10019360 36019365 53ip 3nb:»Q0 n

h3<bnb^36^ x ,r* i - 63i C3n'-) 3ohoi6363o 3u36bhnb 55g5hfi36ob 3i 6bn 3ocn ^qjhm
gogo5, hi.^5 jqjhm gogb56b 33d03&b 050 3hm^m3p i‘b0nb 550m3hoi6nb Sh m ^b n b
33^3h36ib (frsdmdhmdnb d j 6jp jd s ) 95 (15(5 jqjhm 33^59 n,556b 3hm^)m3pi'b0 i 5 5 -
3m3hm6ib giboi63636ob jp63norish36nb 65^083 (frsdmdhmdnb j 6*hn 5 6 5 5 -
dmdhmdnbspdn ^mpjh^b^jpmb^). 301801001309(130 8(5365(133630 5i8oi3hm6nb 365(10
65585019 9565005; 535035boii6 6hdpipnb 3<33j^3hoi&5 3(3(101 jp 3 oi3hoi6 nb 8 3-
8563036300 33^56(1^83600155 55630(101636300. 33h50 (13500163680 hoi3 OihbQimb,
3(5365(i3b ih bôhgjbi i3yn06pi9 535030b 58^560 nyoi: 535030b 01530956 5131103-
65 3301^0365 55h3oi33 b5bouniub0oi ijoj,0ob bBhiojn 3o00o6ih3oi6ob i 6 801^03
b3hoi03juooi 33homgob 83833016001. 83hi0 ig5o0bi3yoiqj30363o 3 1 5 3 6 ^ 3 3 6 ^ 55 -
0ihR36nb 93bid03& x bv|)hi^)35036b 5306336366 6ib. 6.63 (jb3hoi(30^360).
535030b 50hnp3&3on JyjhmqjntfJ&n

33hi0o 33(100103606 559565,55659 531309363005 ghoi30ig 3 m36o<393b 5 5 0 m-
3hm6ob5g3 o 895(1590 01360360 56 53503obi5i 6 95(3300 b33(joi03ho rnhîBm^bn.
3393001013(101<30,5360 ihoi6 656801503 bibnôiijbooi 33hooi9ob 8joi63 ^3(1^0013560
815365(13360, (10180360(5 £>03936056 d0O3hn 6308360b 8 389 35 g i b6h5(359 50-
0153(13636 5i6300iih36ib. 3iOi, 33oh3303b yni30obi, 00353013636056 330oip5 m0 nb
3hoi60oi6o 8(5365(13360. 63080b 3<3303h36o gpob bi65hd0n3oi6obi93o 630,5(1503(10
36 g i oy3636, 63080b 8 389 35 y3i3O0Oi6i hoi8 33d0oi6. 8ioi 330(15^ 3b 0165 6yc>nb
8ihi5obi93o bBhiojo g i b^bô 335336 55. 535030b 33hooigb obo6o 5,55(13636 55-
3 oi3hoi6nbig8o 3g5higo oi3b0ob b,5590530. 5301(30,5366 5iiR6nioi 6y0ooi Ogngihn
8o6nbj333i 01(155601360 (hn‘boi336o, 6010^3360), hoi3o36oi5 6 o ig i53o 951530360
5(1056 d0o3ho 553oi3hoi6obi5i6. obn6o b6hiqjig o(bhg36oi6r y3i3o0oi636 Qi nbb-
i3 3 6 6iyoiqjb 63080b 33hooig3o gi5hoi30on 6ib8nh6yi06i936nb 65(13133.
Xerophytes
+
[ Drought sensitive] [ Drought resistant ]
* +
Drought evading Desiccation Desiccation
avoidant tolerant
— by by
pKenologlcal improvedwater protoplasmic
plasticity: uptake tolerance to
desiccation:
Pluviotherophytes _ efficient Poikilohydric
Geophytes water conduction species and
dormant stages
restrictionof duringdrought
transpiration
water storage
6jb. 6A3 93hjc h^nmBjbSn 8036^3015 j>ip>hR;)6nb 93ji6n'b9o&n (Shantz, 1927; Evenari et al., 1975;
Turner, 1979; Ludlow, 1989)
- 384 -
6D605ff»030 36d6n63aO
5s3n0hm6nb 9o6x 3&*

5i9m9hm6nb 930i 6 x 3&3Cni y3DC^ nb e>rr)6obdo36i, hmOQnb 93933m6ooii(5
01536^3, Si^hnbi g i Brogi^nb bo09hi0ob 3ohm6369o, ib^hb^ftb Jbro3o0369o
6ypnb bi^Sihnbo 930y33Qm6nb 9 3 6 ^ 8 3 6 3 6 ^ . ^ 30 10 C'rn6nbdn3&3&ro: 6oigij,o-
g i6 Bygnb 936m3ob &i303m63b36i, Byonb j,i(530ob 338(501136^ (gnqjj'bn^hn 6o6i-
ip0g35m6ob 2 ^ 3 3 0 0 g i 3<33Jii3hn fcibhgi, bi^h i6b3nhiunm t^giSnhob qjihoim-
6nb 939(jnh3&i), Bypnb jjihfcig 5iîh3&nb 36iho g i 9nbn 9m 9ih i536i. 63:>C3nlJ
936x 32>nbioi3nb g i9ibibnioi3&3Qn 633hn oii3ob363h36i 9mhqjm0m5 n3hni (6ib.
6 .64).
Sypnb 5i303m63b363Qn 936m3i 9nn(p633i g)3b336nb o B ^B bn }!™ ‘bhgob 63333-
mbnon 6oigi{>ob b i d i n g ph0b qj3636i0Q3. b^3336nbi g i x ^ ^ m b 0(536ili336ob
qj3b336n 6oigi50o ^(^63336 2-5 0 bnph030g3 [247], i 9ncn ©(536^3360 ^(^63336
6oigij,ob ^360^6 03363690 g i 63^03^ 3<39hm gogbi6b 0 ^ 6 3 6 . 99hi0 h35om6369o
b3-0U36ih336ob ip0m6i(53636o 03001^3636 dohooiig <33b3b, hm030O(5 9u36ihob
bb3i 6i6o036'b3 i 0)33(1 gogoi; 0^(53^0136360 0300^(13636 d0O3hig gi^môQ
<33b3oii bob^30ib, hm0Qob (5iQiDXn &3ho0o 0 ^ 3 3 6 0 i(p63336 3(1010 93^hob
bo[ph930^3. <333b336ob 060036x^13^60 6i6o0o 5i03hJ63&3pOi i 6 6ygnb bi0ihi-
&mg ihob 0nBym6oQO. 0o6ob'b3gi g i 0o6obj339i 0ib3&ob 93q jih g36i 9 ooi 3q3hm
fcigibhopoi i 0 3j)i6ibi63(yibi36, hi(5 3q3hm 99hiQOi 3b i 013 ob h3j,om6o. bohoi3-
03360 6ih9moj963&i 9i 9 o6, hmg3bi(5 cg3b336ob bo3h[5xn 6i 630ou>h36oboi3ob
5ib. 6.64 81536^300 bb3igibb3i bibnumubpm cgrnh3 ;)6 n, hm903&ni5 3^13633091110)36 53^03nbib 9iai
5igbhfi36ib: a q)mcnrx3(i336n 6mcnQnb b3, hm9x nb p 3hm3n :>5hm336b gngn himg36m&ncn 6yi0b
(igs6brri6ns/bmhn'bnib ,jnin); b b3iX3îyi. hmSx^HJ 6yi0b i^hr^^db p3hm9n (y>j^3bn/33q)mh6nib
^n3n); c b3j)3036(jn, hn9xo|J 6yi0b :>5hm336b cpmoi0369n (i&î/ihsbx^b iin3n); d 9shig963s63
b3 s6 63fijn ph9ig j,}63ncn}h3&xft 9d0i3hn 0^3011 bnb^jSncn (bicohmajnQ^hn ,jn3n); e
qjm<jiri09(j336n gi bSnhig 31183300 63813611 (is3ihnbnb ,jn3n); f pjhmSn Jprnhmognonb 9o9l533cn
636^36(1 (h3î9ib ^n3n); g 9i3hn3>>mhgns6n 9}hu30Pi363&n dcnjhn qj3b3cni bnb^ncn gi J>i6ibc3-
6sgn i3nh^36noi (ihnb^ngib ^nUn); h 6i0o9i 91536^3360 (i6i6i'bnbnb iyi3n); i ^mojryjjbn gi9>
5hm33630n 0^ 336001 (iSfHjhxDbnb ^nin): j 6md3ns6n gi j,363hn>6n 53^103(1^360; k
q3C33nmoi3hmo3n^36n (3hot6cn>6n 9(536^3360); I {pdm3hm6nbig9n ^m03hj6^xo 9536^1336(1 (3mny
n0m1ngh30n ^n^n) (Troll, 1960; Stocker, 1970, 1971; 1974a; Sen, 1982)
- 385 -
9(5066^3050 u e x n o u ^ a ^ a n a ^ o n w a o
195090 ih 601110)366361. iSn^mS, 306(101^0103000160 6(1191500 0^0163 ig5O0bid-

90103303690 3(1(5300 03303011 bnO^SnO 6^0163 9(5361(133611 (5160153011136001 63
0(5361113360) 531030019811 516015301(136001 05hd6oi6nih3i. 8360163600 5i b i 96-
31636019 (S^ng^h h35nm6360rnj) 510010936361 0boi0oi9 5 310 315 10 ^ 0 3 0(5361(133-
6n, hm0036Onj umî 6yi0n 156190103003601.
ByonO 51,51(13600 ] 6ihn0 161(^0361 00006331 5i0îho OnO^30oO q3ihoim6ob
0oi0ij)36noi (03^0 jOo030i, 03010103600 303(101 06(536003(10 gidihjjvji) 9 1 69000
51O13030O 5t>n0 330uoh36ooi (^cghm 001503 o6(53h6oign36o). 013 1 0 1 3 3 ghmb
930|5nh0i Oi^hieOSohi^noi ^giôhoO q3ihoioi6o, 031(190160010 516,51(10 cb30i 5nhn
5036300160 (333(530300 i6Ooi03^3ho 5i0îho (b30i 3ohnb 3ohoi&363n).
^hi603ohniOo0 06^3600301600 333(50(1361 b30b 36yoi60 6yoo0 (13133(133600
35016010010. 0oi030iuo3hi0 3O ShoiiÔo o0ihoi36i 6153360O gh013919 gib3h3o0
03933016001. 0019003051(503(10 8 35336 1 51001061,5361 Oo09hiooO 3ohoi6369o 9 oi-
‘bihg 030101036^3 S i ^ h i g i 0ôghmg 516 0 153630 0 61533600 iSihiôO Fii0myi0n-
636190 (61b. 6 .6 5 ). 103010 0(3361113360 6153360O 03933016001 3qjhoi 51(1519 1(135-
300113636 j)hi603nhn3Oo0 06^3600301610. 53103001900 ‘^ r n q jo ^ h ig " 9 3 5 3 3 6 3 0 °
0(5361(133600 03010103600 33ng3h0oboO 53g036o 303(101 53^060130(1363001 g i 511ft-
60101 bj30O 'bhgnO 0936360. 6153360O i 3ihiô 5160153630 01 0boi0oig 030101000
J3301 b3gi3oh‘b3, 013001 6153360 S i^ ih n g i b9 ohig 516 0 153 6 30 0 1 63030360b
J330 (("1(1360, 5ho3(536o; 6ib. 6.66). OiOi<bp3hoi 1 3 3 9 1 ^ 3 6 0 0 6o6i i p 9 g 35m6 o0
51'bhgob 61(13133 qioioioigO 5ih0 3h^y0oO ,5360160 3i3hnO 03361. 6y0ob 51153000
93‘bp393oO 3hoi-3hoio 3<33ji53hn 0 19 310 36 1 ihoO 030101000 0 16 3 3 3 1 (nb. 61b. 4 .4 1 )
16 0i^hi603ohnjom "b3gi3ohnb 838(50(1361. 6y0oO 93030(50,500 30(10163600 516 30 -
onh3630n 0301010360, hoi5oih(5 63O0, 3i^ ih i g i dg03h gi6i633h330Oi (jbjhmdmh-
qjmbr).
61b. 6.65 Phaseolus vulgaris-nb ajmanprib gidihp3i gi bij^jbnb bn8^nghm33 Bygncn y>h&ig
3m8ihi536nbi CbQgi Irjhicnn) gi 6nigij,nb 5S]6acni363bnb (J33gs bjhicnn) 3nhmd3&3n (Tumanov.
1927). bn99hicnb b^hjbnb ghmb 6(536ih^Sn 3]bj8K633ig nVigj&i i&btfn'bnb 3gi3nb j)fn6(j36^hitfni
(Quarrie & Jones, 1977; Ristic & Cass, 1991)
- 386 -
67)5066030 b 6 6 3 b 3 C?0 ^ d 6 H(fi360
a b c
6sb. 6.66a-c jb^hmqjn^n 9^]6ih336nb qjmaipnb b^h^j^jhnb hi9g36n03 Si^bcnCTin. a Cerium olean-
der-rfo biQjhmqjnQ^hn gjmoipnb ^hnpn. 83088633^ bjjQn 3n3mp3h9xn 9h] ps 6hs3i03hn>6rt 63b-
hnb^bhn 3sh36;jn3i (6:i(jhnbqjhie Saqnhnon)- 6^533611 Qi(J3cni £>hnb t>36;>3C36iTb6i5; b j>senojmh6n-
3Cn Hemizorua luzulifolia ssp. rudis-cfo qjmoi0nb ^tinon. 3b 9(j36:>h3 5yipb ij)hm336b 33j,jn6nh3&x
3jh]g9mhnb363n (5^hnbqjh^) S^gj^hnpn); c icghn^nb 3gi66m3&nbcn3nb p^3ibibnM7i363^0 b^xs^-
ii3hn 9uj6ihnb Sygophyllum simplex-cfo unpn6eh:30n qjmhSnb gjmoicnb ^hnpn. j>ihj,ig 3;yi3f;6;]3i
Jcml135jn0nb 9^nghm 3h] (6i(jhnbq)h}{> 3jg3hrxyi), hmSxi^J &shb iihi3b (?i n(ji3b 6yc<ib qs3i-
dhmj^&x Jbm3ncb (Stocker, 1952; Morse, 1990). 3m6s(j3336n iij^3b36nb 33^^36 nb. Nobel et al.
(19926)
(jhi6b3nhii5nnb J>i6b:>i30th36001 bBhiqjig g i gngn 9ib9 ^>636001 030jjnh3&i

a^36ih3b qjmaifn(^336nb 03933^6001. 00hipn h35nm636nb bb3igibb3i
b 3 0^36ih3 63;>C3n^ ghmb j>i6nugnb 33hnmp30 q3<noim0U336ib. gngn bm3nb b3-
6(336^3360 (T>3hri36nb 33hnmg3h03QO (5hi6b3nhiyoob 03033m6ncn $ih 5i33 6 umî
Byipb. 6530nb36hn 636^360 g i <bmj,n3hcnn (j'3hm-b3j)3036ljo bi^nhm3&nb 330cnb3'-
3i9n 061(3636 cgmcnm0i3336ib, hnb 5i0fnu 0ioin b3gi3nhn 1/3-g i6 l/5-0g3 0(joh
3&i.
6ypnb 0m 9ihi{,36i ihnb &i0rn0hm6nb&i6 cni3eil53nb y33Ci»'t>3 bh3pypiqjoc
g)mh0i, j,i5bii3CJih36001 0i 3n6 , hmg3bi|j n&o giii30oh3&]poi b3gi3ohob 030150(13-
6iboii6 g i 0i(j>iQ lj)hi6b3nhii3n30 6o6ii(j>0g3j)fn6iboii6. ByQnb 0m0ihi53&nb 36V
hnb <bn0ig 6333Cn63&i b3j)3P36v53hm6nb bihnbbn (DS) [4 2 ]:
DS = Bypnb 930u33Qm6i {^33(13630 3g{>m0ih3m6i3n (j,h) / “b3gi3nhn (gQ2) (6 .4 )
b3j)3p36^,3h 0^36ih336b 033dgniOi bi6&hdpn3n 63^>C3r>b j,ig i^ i6 i; 6mpn 630-

0nb 836035 6ypnb 9ihij,n 0u36ih3b ymqg6nb hi0g36n03 33nhnb 5i60i30mQi0n,
hnb 030g3j,i(j 6i&336n njj^xw&i 33h 9i636(i)3pig (6ib. 6 .67). 6yi0n, hm5mh(j 63bn,
j)hm3g36i b33^niQn'bnh36x mhj)i6fn369n. b3j)X 3 6 li360n Bypnb bi0ihij,m jbm-
3np3&n (nb. 6ib. 6.66) dnhnoiigig 0mcni3b363cni cgmai036nbi i 6 p3hm36nb On^-
6ncn. 6y0nb 3m6bih3ii500b 5i6bii>30ih3X qjmh0ib 6ih0mig536b Bypnb 60i d0n3h
^n^hmbtjmSn^hn 6ib9nh6yi06ig36nb 0n3h 0mh6rri3i6 3 3 1 ^ 3 6 8 0 g i ^ 3 0 1 3 3 33-
h3g0mhnb bo3hu3369o. 63i03nb ghmb i0 ^ 3ih ig 0m0ihi53&30n ByiQO i6303&b
qjmcn036nb ^»>6rn6nb 3hmi53bb.
bi6j,hd0n3n 63^03°^ 3nhm6369n gngn 06n93630(ri6i 36n^3&i 6ycnb fcigiigg,-
- 387 -
80066(^0060 usrtObT)1^ aânaônw ao
*o
na
B 6bb. 6.67 ynpnhiemb 3g :>*l® m9r* B nig^n b
5i9m3hm6i (*Fsoj[ 10 b9 bnp+iS^'bQ) gi
Ferocactus acanthoides-cb bij/ib pnqn'bn-
^hn 6n6isc'0e3{>m&nb (rs gpnb 3n5n939n)
Months "bhgi (Nobel, 1977)
n036ib b i9ih i5m jbm3n036ng:>6 g i mh5i6m36ngb6 (b ^ ^ n b b h m ^ n g i gngn

^ m ^ n , b b 3 ig ib b 3 i b i9 ihij,m mhj>i6fn). 9 3 h i0 h32,nm6369n bm^nphonn b^ 9^3-
6shnb (T>3hm369n bg3& i b i9 ihi;>rri Byonb ^h n m gjon 6i G;>i>G6nC3^)-:>- gig 6 n 6 g i, hm9
2)^6ni9n 9m bihg 6im 6i6369n p3hfngi6 qjmcnQ369n yn 33cg p n ]h ig g)ig ii0 2 )n 0 g 36 i
400 pn^hn 6yi0n [6 5 ]. p^hmbi g i j^hjb Bypnb b i9ih i5m mhj>i6m 36nb q jjB jyn i
ij3 b 339ng3h h3&nm6369n 6^3^(3303^30 9(J36ih 3369n. 
3nhm6368n 4 0 6(yib qgn^336nb Sîgpinb ^h i5 b3nhiunnb 6yonb g i6 ib ih j3 6 n b 30-
5 0 % 9 mgnb (p^hrnQdSn ihb3630n h3b3h33&nb b ih jb 3 [ 160 ], hnb 6 ;>9 ml5 Byonb
6i0i6bnb gidi63Q m 6ib 60633690 igôQn ih i ij3b . bn99hi0nb ^hnmgBn 6yi0n
ou>3g i 3nh330i g 5 ig iig ^ n 0 g 3 6 i bhmb qj3dnpi6, bm0m 939936 9 nb b b 3i igj,-
n03&ngi6 g i dmpmb ^m^36ngi6. 6yonb 6 ig iig ftn 0 36 i n9 n^m9 ihnb 960 9363-
0m3i6n, hm9 6i&336nb gib3h3nb (3.0. Byonb 3im 6m 9nib3 &sgib30nb) 9 3 8 9 3 5
hfi36i Byonb 9ihi53& n b 3hcniQ3hcn 6yihmg. 9 ih ig 9 5 3 i6 3 9^36ih336nbiOi3nb,
hm903&ny h9 n yn63nb bn99 hi03b $i6n u gn i6, Byonb 5 ig iig & n 0 3 6 i (p3hmgi6
ihnb j,igihfi36nb 96n9363om 3i5n 93î6nb9n.
5i9m3hp»6ob ]6ih n
6i 9 m9 hm6 nb 36ihn ihnb bib3m6nboyjnb b33isnq)n2L)hn 3hrn^>m30ib9nb
93533&nb cri3nb36i. h3<bnb^35l5 x r r'^nb <bm0 i g ^ 3 6 3 0 0 $ 9y36ih 39n g i 9 nb
jbm3n0369n (R W C , W SD , Byonb 3^^36(30^011) Byonb 9ihi^nb bibib(p3hm 95n93-
630^6360, hm9036nb ghmbiU n6y3&i H o lto n 9 3 3 ^ 3 3 ^ 9 0 3hmu3b 36n ("i)hn^nj)30n
■b(p3ihn") i6 6 3 2 ^ ^ 3 0 ^ 9^(3*333360 (b^&ODi^CDhn - 5-10% Q ibni536nb ghmb,
- 388 -
<y360i>fn030 b & fn S b ^ O ■JtoddHfnS&O
53^03nb D L 50 - 50% 9^(1^63606^6). Sypob 938(j339m6ob 55'bm830b

9393559 8 0 0 3 6 x 0 0m6i(5303bo 00)0330^6 bi2»0ihnb 6ih0m g536ib 535930b 3o-
110163680 ihbQbxn Bypnb h3‘b3h336nb 8 3 6 ^ 3 6 , 655*150 85010 559 1^ 36 3 6 5 bib-
3m&36nb 83b5g5h3695g ih 93od036s, b ij0 3 obii, hm0 58 85R3363690b >6bmQ3-
^hn 660836390165 95801509363905 5 5 6 ( ^ 5 5 3 9 3 3 bo63b3, 855., 0ibt>3 5539365b
56936b jbm30Q360b 83^56053(10 39303615360b 60160010 6090. 553m9hm6ob:>g0o
b33(3oq)n53ho (5m03hi6l5m6nb 9 ^ ^ 9 5 36 ^ 9 83od93&5 550oi3oy36oioi 6y0ob 301(536-
I5n>0nb 80165(330360.
3hml5rr)30i <b9 ob 5i0m9hoi6obip0o h3'bnb(536l530Oi65 0(536:>h3ou> b i3yihm9 o
dpn^hig 03hy3m6b (ubh. 6.7).
5^9m8hm&nbân 95hd6m6nih3 taboctfgbn

015365113015 303,53b oi6nb 3(101,501395^05 5>9m9hm6obs90o dQ03h 05hd6m6o5h35.
3m0mnoi3nph39o £311^9013560 9i536ih336ob cgmai936nb 331139360 0^3336056 9ib
036935, hi(5 (156936003 b55ainb 5560539016580 @531533059 3:^30160(1936056 95-
96% (633b5658365 -11 95 -5,5 MPa) 3:>3hob 035(190160010 li,36o56ri6ob 3ohm6369o;
qj3b336n 50933 30jhm 05hd6m6o5h3i, bmpm yomhdô - 0395h36ocn 550d93. 55-
6oi6hoim6ob 83933016001 (1313011,536(5390160b b5‘b(p3h36o 035(1010139365 (65b. 6.68).
‘bhpnb 33(1001980 331139360 556018(1016065980 5^6b5530ih36ooi 85(^601605(13 5ho56,
boipoi 0mb33636ob 33(1001980 558oi8hoi&nb598o 85010 895(1590165 851539016b.
au>0fTiq)ol536b 9 mhob 3056jl5m6ob 6^99(5365113360 95 (ph05 6yp3bob 6 ^ 9 8 -
(5365(13360 (013 553ocn350ob6o636cn, hm0 6mh0503h 3ohm6369o obo6n SyQocn
5ho56 55h930mhl5y93Qo) 550m9hm6nb590o 556b553cnh36ocn 85hd6m6o5h3 5(1056.
3oju335-8mj|j33nb Id0160b 6959015365113360, h010936013 0oju33ob ghmb 33(100193959
61b. 6.68 Olea europea-b qjmoigj&nb

6i9m8hm6nbiC0n h3'bnb^36l>0Cm&5 6m,jnm (•)
pi 09hicn (0) “b igbx0^ 3nhm&3&3n. 1953
Bpnb 6308016 lbiq)bx9n qjmoipjbn 01)3636
6iic]5>sp ^mc3hi6(!)XD2>n 6ycnb 93(30^0^0-
big9n, 30^3 Qhorm gpncn igh3 33hip
inhmbjaSo (Larcher, 1963b) Water saturation deficit [%]
- 389 -
01336^0060 U&rtDL^ a^ arm rto m aao
libhncn 6.7 815361113011 3jh3P36nb {>i9m3 hm&nbie0 n t$rric3hi6(5m6i J ijh n b b b 3 i9 ib b 3 i o jih pmdnom
(536ni6m6nb Snhm ftjM n 12-48 bon Smcn^b^&nb 03093£> (8 h i3 ie n ^ ^ h n b 0nb3g3ncn; Larcher.
1973b; G aff, 1980)
a^shocns }63<3n ,yn(v)hiS,jmd>

Jyjhnb <5ihgm5rtcnn Sypnb
^36ni6m6s (%) Cn (MPa t
tnj^nb BipoaiKtfohxjbn
j339i bjbcnoimhic^hn 6yic9u36ih336n 99 - 97 -1.4-9*5 -4-9p3
b36cnonmhi03hn 6ybc9i336ih336n 95 - 86 -7-916 -20-063
33Cncnmhi03hn (33lj,n3l5n336xf>) 6yic0u36ih33&n 86 - 83 -2O-916 -25-0g3
HepaticaceaeA) 6*h9m9iGb36xn b»3l>36n
3n&hmojn^36n 95 - 90 -7-916 -14-893
93‘bmojnl536n 92 - 50 -11-916 -94-993
#J3hmo)nli|36n (36) - 0 (-140) co -6eo
bi3b3&n
5njhmq)n(|)36n 95 - 90 -7-916 -14-693
83'bmo3n1536n 90 - 50 (10) -I4-916 -93-3p3 (-310)
Jb3hmqjn1j36n 5 - 10 -4OO-916 <0-693
Hymenophvllaceae-b 6ih8m8fc>p{)36e3&n
Hymenopiryllaceae 90 - 75 -I4-916 -38-0g3
b3n9h3&n
Pteridium aquilinum > 90 -14-093
Cystopteris jrag ilis > 90 -14-893
Asplenium ruta-muraria 40 - 60 -7O-916 -120-aeo
3m9mnm3nph3pn ynhOmajnA^bnb
gmonyib 33ng3h8nbn 96 -6
83(bmqjncn 95 -7
g3b3nb j>mh,j3jbn 97 -4
UmnyiprOnehxo inhSngn^Ki
Bon'd nitida 85 - 90 -14-916 -22-093
Xerophyta villosa 66 -56
Myrothamnus flabellifolia (in situ) 11 -298
Myrothamnus flabellifolia (ex situ) 11 - 0 -298-916 co-093
bn90hi030n 639360^6, J,i 9 m9 hm6 nbig 0 n ^gjhm ,im03h:>6(j3C36n ^>hni6, gngh^ ph 9:>
6yQ3&nb 6yi00i536ih336n. 9hi3i 0n b ^ b n , ^Bbiiôih^bncn p3nd0(ib b ^ b jb n ,
ihb36m6Q6 3bQhnb qjihgn6noin l|)36ni6m6nb ObmQmg <3ihj,Q363n, i0n^-
m9 3b 8536^336(1 33nd036i J,i0m3ny36moi hm^mh^ n6gnj)i li)nih36n. bi3b3&3n
J,i9 m6 hcnm6 nb 03033m6noi J>i9 m3 hm6 nbip 9 n 0 gg,higni6 i bij,hd6m6eig 0^30^166
5b9 m3hmbnbê0n iyTK ?3^<!iX n 61336^336(1

CTii>0mgn^36b Smhnb ihni6 5i9m0hm6nbig9n bh30nig 0g&hign bib3m&36n.
6ijvj)3hn360nI I3nb6m6ij^3hn360n g i (^ 36 36 0 0 ihnb 601300 hn&n 6ih0m0ig536-
036n, hm0036bi(j 033d0nicn oi33(n6nor»i g i 60m6ncn 00hipn 3i3hnb 3nhm&369n
ymqj6i g i g i(j)36ni636nb 030oib333i0n 03î6m0n(b0nb ou>3ngi6
(bnj)n3hoi 0iOi5i6b 033d0ni ihb36m6i i 6 bm03lj 3hig 00hi0 3i 3h0 n j^nh^dnb
- 390 -
bDBObrtOaO Ii6 rt0l>7)c?0 g 6d6 r)(f)060
5i59i3Qfn6i9n. 5i9m9hm&nb:>g9n b h x ^ ig 9g5hig36rii ‘bm^nQhcnn bmjjmb 9riu x,v

39360, bb3igibb3i bi3bn, 5309*1360 g i 93p i5 n63QQ>b bib3mb36o.
5i9m3hm&nbig8n b h x niQ 9gj,hign bib3m&36n ihoi6 y3i3O0m 3i6 9^36^301^
3mhnbi(j: Gesneriaceae-cnijpB - Ram onda serbica g i H aberlea- b bib3m636o
biOi^D^nb h3£,nm6ngi6; 9hi3ign "9i3gh3oinCTi iQig5m8nb ^6ihnb 9jm63 9i536i-
h336n", hmftmhnu ihr>i6 Myrotamnaceae, Scrophuliceae, L am iaceae g i Velloziaceae
6ih9m9iQ636p3&n, hm9p36nu 5i 3h(j3CD&3Cn ^hni6 U36^hi0^hn i'bnnb 99hip
h35nm6360n, i3bj)hiQOS0n, bi9bh3cn i93hnj,ibi g i 5i6bii3cnh36ooi biQbhôn
icghni>s9n [71 ]. hm&mhu 63bn, b iJ93 53ifob 8hi3i060Oi6 338o2>ho3ljmcgn(|,36oii6,
hm030oniu ibibnioriQboi 3iîh i, b3 ohig g i b 3 3 X n gmcn03&n, 6300 b h g i g i h33hm-
93Jl5f)nb gi6ion 36iho; nbn6n 53^>03ob 3ohm6369o 5i9m 3hm 6nb5i6 o i^ g i^ n b
3033^3!™ 93j i 6nb03bnb 3jm6Qmebnb 5i 9 m (0ij,„ 6ij>33&nb 530^6 gij)3^3i C4-9ih-
I330ni3636nb 839016333^11, 6ypnb b i9 ihij>m jbm3o03&ob 013 mh5i6m36nb 3jm6-
Qmbi) 3u6ig J,i36ycm3g36ni6; ^ 3 3 ghmb, 9 icn ibiboioi36oi Bypnb 3m^36(5nipnb
3jb(i)h39i 03hig gifeipn 8^336360 360.
5s9m0hm6obig6n 8g&hign y330i 8 15 36 ^ 3 33h3gnb 3jb^h39i 03h gQ^nghi^jnib
nî6b i6i6nm‘b3h 8g5m8ih3m6i0n j,igib30ncn, hm00ob ghmbi^ 6n30in3h36iDii
I530i cnnoij8ob 03R3h3&xni- &i9m3 hm6 nb 33homg0n 3b 8g5m 3ih3m 6i 8nnp633i
53i 03nbig9 n b^&ng^hn 3hm^3n636nb bn6o>3'bnb g i qjfnbq)m0n3og36nb 8ib^n6n-
0nIbnh3630n 6ib0nh6yi06ig36nb (hiqjn6m'bi, ^h ^ io m b i) 8 3 8 6 ^ 6 3 6 6 0 nB^^hnhj-
6nb bi03i036ncn. Ibmj)n3hcn bib3m6i3n 3jh3gnb gi^&3b36nb 3hmjj3bnb 0 3 6 x 3 6 }
i6 03Rh3&i bmh(jn30936i 9nbn 63360b 5 X n'J qjmh8 i 3 n îgibgn b 83033^6001.
5i9rn3hm6nbig8n 9gj,hig 3hfn^m30i'b8ib ij3b ^ 3 0 1 3 3 6ygnb 6ihi5nb 9m8i^3-
6nbib i0gj,36nb 3hmu3b36nb oii6giou»6m6noi BihSihoqnb ^6iho (6ib. 6.69). i8
ghmb 33h3gob b ^ h ^ ^ b o b 1(^1536001 8 b ig g 36 i Spîftmgn'bSnb * 1 3 ^ 0 3 ^ 0 1
(33(1 b36oij3i g i 038935 <gm,!jrabn6oi3,bo) g i 6mh8i03hn 6b3030m 6i.
Time [d] Time after rehydration [h]

fbb. 6.69 a,b imnyiQfalnghx 3(5o6:>h3363n &>3fn3 hm6nbsb 9jjs6mo3hn Simu^b^dnb n6iijn3H5ni pi
Siovi Bycncn 0n9>hi536nb ipej^nb 938936 (Ried, 1953; Bewley, 1981). 8^16^530360
DOii^^h^^hnb 153(^1036360^ 9b qgm^mbriBoij'bnb S jb ib ^ nb. Tuba et al. (1993; 1994)
- 391 -
aooe^foooao a ^ a n a ^ fn a n w a o
6 .2 .5 . e ^ a c ^ 3 3 6 n 0 1 6 ^ an 6 3D 3C fl biihObfl
S^bidoQbxni, 9 3 9 ^ 8 0 6 ^ 3 brujmybcnb 65h8m3m6ob 33(1001080 05(10036001 55-

0 m 633X n 051530(10133 nym 30(13300 jo9o3ho 6^(136300 qpj^mho: y33Cb‘t|3 ^>9 ^330°
bibnymijbpm <3^1(10360 ny3636 b 0 3 ob 3nihmmhj,>6nb336n, 903b5U 9^60^336300
3i 6 nli)i^36nb 0ib ib 03&i 6936s b 0 3 ob (9* 3(13 8^363(10 6y03&ob) 9bhog36. 380^-
m3 , mhj,i6n‘b936b 0133093633 369s 538013938333606301 nm636nb (13530313006 93
6001001503(10 b(5(13^3(1360b b^p6n0nb^nnb 3qj3;ji!)3hn 03j36n'b036n. 9bm0m9 33113-
gob 93hn03&ob3g9 o 8g5h3gm6ob 03ji6n‘b036nb 5>9m033i336nbi 9b 801300 9^3-
6bhnb 0m0|33Xn 33bi6s3nbo pm6obdo3636nb ©00360b 03333m6ooi 0ijbn9bQ3hig
83533630 3ic<T>qgoli)36b 833dQmcn g3b3b03630oy3636 3bhno36ob îh6o 3^16(336^(13-
(300b 3ohm63&3o (83(1000560 6059550 80101553b b33C30inb 'b3@i3nhob 935600136001
6%).
6 .2 .5 .1. e ^ 0 C ^ 9 35 X n ^dfliWiWbfl
09583 ^95o0bb0 ymq33036ob b^hoim 01306363b 63(180139536b ^9309^9 bb63go

9iho03&ob 0s(pb0O 3016(536,5(131505. ^ 0 35, 99383 ,56360 9b 5363360 3333,53(10 83(1-
O0Oi6n 5360,53,53603; 6030301^3 530093(10 9s 3(109300 30n03(5nb 3nhm63&3n t>n -
0bhnoos6o 6059550. 6393;) 36001 0gng3ho *1350016360 - 9fT>jLS33nb bm 6i, 90 36360 ,
33^360 93, 53*193 30ob5, 5309063(13 6y036ob <bm63, 9ihno36nb 055(10133606 39 5-
093603. b03nb 83(100360 (^hm'bmoob bibooi) jihob 0O3h 5 3 9 5 0 ^ 6 3 6 3 653oho-
936 h39g36o03 3090183,51106 600(18380. yo63ob ghmb b333^m8m6oom 5 ^ 3 6 ^ 3 8016-
63390 93ho0o 0m53o3636ooi Byoooi 0(13156363 93 b56momg 6o3g353o obb6363.
83ho036ob 333i3330m65 6,536003(105 8bm0mg 003 b03ob 3ohm6368o (6.5333-
onig 480 80m0 o N a+, 560 09m0O Cl”) 8oj(5333-8onj(533o6 lbm638o 03hnoo36m63
d0O3h 03hy3m6b (8365h36ob (bm633o - 290-810 88m0n Na+). 00383 ^30163680
N a+ 3016(536,5113(505 30 6 33b 600-1000 88m0b [9 ,6 6 ].
0U363I1330 bscgshSn 3353(i3(3nob 33homgob 5368530m633n 3mhoijo36ob 8 39 3-
5 39 bg363 03ho036ob 0i56g30i36m6n0i 955I1P13365, (11^9936013 8|3365hob b o ^g n o o b
330036. 6303j36ob 03833m6ooi, ob33 60393580 6h36g36o36. 05ho036ob 6h 3635b 3
g3 9mdh5m65b 33901030,53(1 5360,53,53680 3^336366 63b. 6 .70. 6050350b 8360311
g30058365b 5g5o0o 5j3b 08h5o 5360,53,53680 (835., 03^3b 9,53,50b 80383 303660180
- 2400 88m0-8g3), b305(3 6030350b 301(10^0365 3qjhm 8303005, 300(13 83b3o R 3 -
b30o 65p3j36nb 60o3ho h5m036m63. 83ho036ob 556653301(136001 0000 30161536,3(13-
150360 0 3 8 3 b 3 b n 3 O i3 6 3 0 n 3 0 3 6 0 m 6 3 6 o b 3 C T i3 o b , 6595(5 5(136,506 6y036n 0 5 0 0 5 5 536-
Ci 63£>3Cn' 536336036 03h^5q3363o 05 ob3Ci 305003680, b393(5 06,536603(10 6593-
j3&nb 3ohm&363o 6yoob 9(136330 8003(1 g3&50O5.
338003(1 113500163680 89383 6n305536n 830135336 ^3(160 (1301036016001 N aCl.
3b30!0 ô3nb 630,5(1393(10 83hnco36o 605955360 5hob 35h30i33 88h30 (13500163680.
3qjhm b8 oh5g, b^3336nb5 g5 30566m36ob 89583 605955360 830(35336 ,53^3 h33j(5o-
nb 0jR63 b3oqj3^36b 03 Na, M g g5 Ca 33(160163^366. 58 838016333530 b 3j83
633j3b 038oi>8363boi56. bm0noi 0585hn0n56365b 505noo 5j3b (^5333680 ("bmooi-
63(3", 65,5(10380560 6030350). 63006300b ,536036 0hm8o Na-556 6 3(180 ^ 8 6 36 3
063010 63,b503ho 85hoo36o36n 338360b 3m83o3jb3&n, hm5mho(333 N aH C 0 3 ,
- 392 -
57)606fn030 liSrtOUTRO 3 6 d6 n(n060
N a -h a '1
6sb. 6.70 a NaCl-nb 9n3m(jxi Sipmqjn^^h S^Qng^hn 3nhm6]6nb ghmb (Steubing &
Dapper, 1964); b Na-nb 3n9mij3ps gi 9D9l)3DCfT'2);> Hscmqjn^h 3i6n^)9n shngxn Snhmijfenb ghmb
(Breckle, 1976; 1982)
Na2C 0 3, NaOH, hitf 116333b pH 0 m05^;)i>ib 8 ,5-11-693. 6ib33ih3g:>66m;)&nb jc?m-

hngnb bxgb^n o i pi0ihnQn:>63&i ("bm0m6Rij)nH) 116333b 0ihnQ3&nb $^161536^150-
nb dgr^h 0i^3&ib, Jomhng3dnb, bxoM D&ok e * Bi^hn^nb, 65666360^ y>ci3n3-
0nb &nj>bh&m65v!)3&nb J3hjm3i6n 03611b 6 ih 9 nij36ib; 3b 6nigig>36n 3 gngihni cni-
6i 9 nhncn, bmpm 5 i 3 m06h:>c 3g 6 fTi3:>h3m6i 3 n 0Jj3hr»3n g i ync^rîB ni.
0ihfi0fi56n6nb b ^ i g , 5:>6bii3cnh36ncn 6nigi{)0ijrng63m65b i g i bmqjcnb 033h-
63016560, xP3^ X n:> Bypnoi 6533(136300 6 nig5636nb bn6j36nb 3C3jiihrT'CrW hn 556-
(EC e) S 6 1 . ECp-bi g i mb6mb3h Sm^BtffiiQb 3mhnb ihb36mbb 6hcgn3n
gi0rnjjng363Q36i (1 b 3"' = -0,036 MPa), bmQm 0ihnQ3&nb 939(333Qm&i 6nig563n
6i6nbi'bc>3h36i 6fn6 ri6hi3nb 33633016001 (6ihnp36nb 6m6noin 3hm(336(530fi 63615-
O smotic potential [MPa]
6ib. 6.71 (Snigijyib ftisjh ad x 3jbvjh:>ij9n y.3Snhn

dihnpnb 9D9l53C3mbib, Bypnb qjihgm&ncn
30al}3Xfr* ^ e> X0i»Shm6âi>ih36C,,'&^ (ECe)
Smhnb (Maischner, 1986)
- 393 -
9 0 0 6 ^ 3 0 6 0 b6 (f)3U D ^ g ^ a r m r t o m ^ a o
330m6i 6m gi5nb bb6ih3n: 6ib. 6 .71). 6 m0 m5 hi0n9i 6 Ri6b *51130133 9 i-

hnQ36ncn gi^nhcr^nb 0m0i,j)36i 6nigij)9n Bygnb 030i533Qm6nb j)036iboii6 Qhoiig.
0ihnp3&nbig0n 05hd6m6nih3 9u36ih33&n 5160(59016 4 mS b9"' bngngnb g i ^ n h -
013:1b, bmpm 9mh6y3nb 936935 E E C e ih i(p30i^3&i 2 mS bd-1. 'be^nb 5 y i0 n b
3C3d»!)hn0n(!)3hn 5i 0(jihm6i ip633b 44 m S bQ-1 [5 7 ].
6 .2 .5 .2 . 0^hnCD6nb dX }X I\ in6l336i!)h^l5nD6nb 6^3006^ d U Jb ^ h Jb J
0ihnc3&nb 9i(piQn 9 3615330(^161 9(j36ih3b ^nh ou ^b 6y0nb mb0mb3hn 6 9 nbi

9 1 nm636nb b33(5ncgn3hn 9mj9393&nb 03933^16001. 9ihn0n.i6 bb6ih369n ByiQn
mb0mb3hi0 dQn3hig ihnb 6030n 9 i nm66nb jjm B^Sîijnnb 0i^3&iboii6 3horiip
H5n b30 30)hm 6i]j036i9 b309nbi639m0ni 0^36ihnbi3nb (nb. 6ib. 6 .7 1 ). 3hm-
l5m30i'b0i9n Na+ g i C l' bnîh&3 n6333b nm636nb 6m6ib6mhm&nb 0 3j96 ib ( K ~
9 i Ca2+, Na+-nb b i3nhnb3nhmg) g i 3 6 ^ 0 3 6 ^ 3 , 0306hi63&b3 nm636nboi3nb b^^-
ynajn^h 5 1 3 0 3 6 ^ . 3b 6 i30 36 i 5i9m nbili)36i: 363h5nnb 0 i6 i0 5i0m bi3i03n cgm-
limq9nbqjnhn0nh36nb g i b36oij3nb 3i^33&nb qjnbqjmhngnhobnb 3hm(j3b9 n; i'bm-
tjnb ibn0n0iunnb g ih p 333ib i 9 1 (jn0nb 03î6m0n'b0nb ihi6nh0i03hn fct^bnb
6ih9mj96i 3n (3mQni9n636nb g i gni9n63&nb - 33^h3byn6nb g i J>ipi33hn6nb g i 5 -
hm336i) [6 5 ].
mb0mb3hn g i nm636nboi3nb b33i5nqjni3hn 6 i30 36 i, 0ioin bnîh6nb 8 3 3 0
b333i9n, n6333b < 3 3 6 ^ 3 1 1 gih(?3333&b g i g i‘bni63&3&b. 6i5336nb g i i 363nb, j(yn-
hm30ib(5369n 303j(jhm63&nb ^hi6b3nh^bi g i 03mh3g 3hm(53b3&‘b3 0ihn03 bnb
b33[5nognj)3hn 5i3036nb 03033^16001 6 9 36 1 qjmlimbn6oi3'bnb 93<b(P3g 3i. b36oij3nb
3hmu3bnb n6^36bn3m6i 9 (5i6bij)30ih3&n0i qj3b3369n) 9ihn03&nb 5i3036n b 5 i0 m
33nd03&i 0mn0ili)fTib i6 nj)0mb; hm5mh(5 Ri6b, 50n$m0n'bnbi g i lihnj)ih 6m 6ili)3Qn
0gi33bnb yn^prib 36‘bn03&n 0ihnp36nbig0n 3cghm 05hd6m6nih3 ihni6, 3ngh3 6113-
oin3h3&ioii (5300b 10^3(161^03011 5b3&n. N N aCl 0i(j>ipn 930b33Crn^)n'J 3nhm63&9n
bij)33&n nm636nb (5i6bij)3onh3&noi N C ^-, K + g i Ca2+) 03Oi3nb3&i j)03630mbb.
bhgnb 3hn(j3b3&n 9ihnp3&nb 5i3036nbig9n n9g36ig 05hd6m6n^h3i, hm0
bhgnb bnfijih3 g i 6nm0ibnb Shmg^J^ni 93nd03&i 5i9n3ny36moi hm5mhu 9 ihn-
03&nb b(j)h3bnb 5i6bi'bp>3hnb bin93gm 9*6 3363630 ^ m60n5i^3hn ^0^1030^360
(33î0mq3n(536n, 0 i5 „ Salicornia, Salsola, Suaeda, Halochemiri), hm00360(5 Q ijji3-
9 nh3&30n ihni6 0 0 i93 ig5n0bi0ymcg036aii61 0ihn03&nb 930i3nb36nb g i6 i0 n n6-
l!)36bn3n6nb 5i0m 6mh0i 03hig n'bhp36m6 (6ib. 6 .72), Obmgmg 0ihn036noi d^n3-
hn gi^nho^nb 030oib333i9n crii3b n63&b bhgnb 3hm|53bnb 93630 36i g i b^h^-
bnb 3nh330n 6006360 (i6,jmi5ni6nb 6ih9mj06i, j0mhmcgn0nb gi90i).
3i0mqgnlj36n (0i5-, 9hi3i0n Poaceae, Cyperaceae, Juncaceae, Glaux m aritim a,
Plantago maritima, Aster tripolium) oii3b J>ih5ig 5hd6m636 6nigi59n 9ihn03&nb
gi6i0n 3m6i536l!)hii5nnb 3nhm&369n, boi0m 0ioin 9m9ivi36nbib b6h10319 ncnh-
53636ni6. ihb36m6b i5h30i33 0ihn03&nbig0n n6gnqj3h36(i)30n 0U36ih336n, hm0-
036n)5 nbhg36ni6 dnhncmgig 30ihn0m 6 n ig i5lb3, mpim6g 933d0nion 0ihn036nb
gi6ign j)m6(536li)hii5n36nb i(ji6 i (9i5-, F estica rubra-s, Agrostis stolonifera-s,
Phragmites communis-nb 3 3 0 1^ 3 3 6 ^ Puccinellia- b g i Lotus-nb bib3m636n,
Artiplex hastata, i30ODh30n 9(536ih336ngi6 - 0i5., Vigna unguiculata g i Sijhnb
îhbign). 3b 0U36ih336n g i i5h30i33 ihi3i0mcgnli)36n (3i0mq3m636n) 0ihn036nb
930i3nb36nbib 5i6ni5gni6 thgnb 3hmu3bnb bi5hd6m6 03lb(p39336b.
9ihn0(ii6 3i6nli)ij)369n 6i3hl53036nbioi3nb 3i0fnqjn(|)36nb ic l|0mi53636nb b^hi^^-
- 394 -
6716066030 UftrnOliTlÔ StodftrWD&O
a b
100
4---
4 5
0
Salinity 0 0.1 0.2 0.3 0.4 0.5 [M]
NaCI-concentration In substrate
6*. 6.76a,b piSeiSo&nb &>3p:)6> 'bhe^'bj. a bD&tnjh^n NaCl-nb .yntju^^founnb jpthpnb ^36^303-
6i djjj&ihQj&'bj: 1 2£*o|,|9 niin; 2 <3iiXO^SrOhn i>CfT"3nibn; 3 pb6eb336nbip9n dijnhjp ^mc]hi6-
iXn ihi3bCm<3niin: 4 Jnyrojroin. b Wyjigibbji 3scmqjn,j;)6nb 803b 99hiQn 9ibnb ShmgjJiyis NaCl-
nb jm6u]6^hi(jnit)cni6 5i33nh3n: 1 Salicorma 2 tripolium; 3 Suaeda
e u ro p aca-. 4
A ste r m a r itim a ;
5
S p a rtin a f o l i a s a ; (Baumeister & Schmidt, cit. Albeit (1982) Snb^ncn). 3m6b(j3-
P u c c in e lla p e is o n is
3j6n W>3i 3^c*no}n^3&nb nb. Rozema & VanDiggelen (1991)
5<ib pi i(n3oi6n536ob 3ihoo3&obig6n 6p5higoi6ib 5ip i06y33(5O 0611336300161

ifob. Sipoiqjn^bob nn0m6nj]6n 9 1 ib i0 5 icbhgi 8(5361(13360 3ihfl036obig3n ;jqjhoi
35hd6oi6oih3i, 3ogh3 3oi"bhgo0n 0690309360. 516^33011136001 gog bi3o3hri36ib
5160(59016 i b ^ i 'b h g i 8(5361113360: 6nsgi5ob " b ^ i 05363630, hm00363o(5 516 0 1-
5 3 6 x 1 1 0icnn qj3b336n, 8ihn036ob 3016(536^1(501, hm5mh^ 63bn, y330i'b3 3i-
Cipni. 1^801^36361 bi3j>30i3bng 3n3gn6ih3m6b } 3ihnpm 51(1380100. 3boi0o>g 8ih-
n036nbig3n 0g5hign 1301503(1010 0(5361(1, (101501(1360(511 8 15., Tamarix, Suaeda
depressa, Halocneum strobilaceum, S a lso la baryosma, ib3hb36b 30-40 mS.bS"1 8ih-
0pm630i60b 30(10163680 1(^301(53636^ (hnj 3o3bi618 36 1 gnb0m36ncn 1-2 MPa
osmosur potentials), boi0m n"0rn33636ob ^ 3 1 ( 1 0 dnhncmgip d33b 15-20 mS. b8"1
(gnb0m36non 0,5-0,7 MPa) g i 3qjhm 613036 qjih503680 [202].
3ihn036ncn 3^ (13810 3(10 91^30(10131 06333b "bhgnb 5 3 3 1 qjoih8nb 51630011-
h36ib g i qj3b336nb 'bhpob 3 3 'b p x 3 ib . yomh^&o 53016 00036016 g i 0ioio hioi-
9360161 6150(131, 0301010360 Si^ ih n , b9 nhm qj3b336ob, q3oioi03&ob, (j>3hoib 633hnb
633(101^360. 0301010360 930010936016 g i ^3636016 33h 30033 3363iSH50ob 33(10-
mgnb 5i 60i 30m6i 3 n pi 601001b b036m 6 3(536ihob 8o6nb'b3Qi 61600360b 601300
366360. bo&3hob 3hmij3bn Rjihg36i, (50^01306061^ 30161536^(11(501 303630016b g i
8 ^ x fT'6b i6b(jo'bn6ob 8313360b! g i 30100360b 30161536^1(30360.
6.2.5.3. 6iW hR D 6i 90^9D 6n^9i6flb 3nhm6D63fl
g i0 0 i0 3 6 x ^160,31^3630 513(1 (5x 36x 3(536ih336b 3ihoQ36obigdo 013691(5

3(}nh3 8353360b 51(1333 5191(16361 ih 033d0nioi. 8iho03&nbig0n h3(bnb^36^3poi-
- 395 -
6 i ihnb 9(j36ihnb j 5ihn 3hml5m3pi‘b 3i3o QihnQjbnb 933u330m6ob h 35xn h;)6nb
03933m6ncn 011309^6 bn^nprrib 3ioio 3i(j>ipn $m6i536(ihiun36nb 3ihymcgocnn 5 ^ 3 0 3-
6 i i6 M\i>i6oib 3iho036ob mbSmb^hn 013 3rna6i 0XDCn 6 * 3 0 3 ^ (6ib. 6 .7 3 ).
&hn(3&nb 039u3x * ,idnb h'ftflpn lrfo

3l536ih33&b jîfiSnbOi 3ihop36ob 8381533060b h3530oh36ob 3h i3ipn b i9 3 >-
C3&i:
ashnc3&nb 3])hi6nh;)6i. 3i 65h36nb ^01503)1010 qj3b3363o ^hiBb^mh^nb
6ihn3h3&nb 93j36 nb 33633016001 dpr^h ^>3150113636 3iho03&ob j)m6^35vjhi|3nib
I536(!)hs03h I3o0o6gh3n g i 5 i 3 (jiho bnb^Snb bb3i 6i6np33n. M imosaceae-n b 6bh-
3m 3ie53630 P rosopis farcta-dn igj,npn ij3b cgm0103680 9ihnp36ob ^hiBb^mh^nb
bij,hd6ri6 333(5oh36ib. 3iho03&nb 01^16360 (5i6bij)30ih36noi Na+) 8300i3nb36n:>6,
3 i5 h i8 i8ob 3 3 3 5 3 5 bg3&i 3icnn 9 ^ 3 3 6 ^ 0 ^ 3 3 6 6 1 1 g i (P3hm369n. 8iho0;j6nb
^hiBbSmh^nb 93fi3h3&i g i3ibiboioi36x n i i5h3oi33 bb3igibb3i i 30\53h30n 9 l3>
6ihnbioi3nb. SiQmqjn^hn 3i536ili33&n 5i6bi]>30ih36ooi 3<23Jii3hig M P33636 nm-
636b q33b3369n, p3hm36ob i 3ojp03h 6:>6np369n, qjmaip3&obi g i y3i3np3<bnb y^S-
63680, hnb 5i3m u 33hnbli)3 3 i ; 3nbihgn 0301010360 g i 30138600336300 6iym qj3dn
6 iiC 36 ig ihoi6 8iho036ooi 9.^30(101300.
3iho036nb 8nxyiQ36i. 6i536ih336o 8iho036ob5i6 aii3ob3qj0g36oi6 h ^ h ^ n n b ,
ijhmgign j0mh-83oii6ob 5i3mymo)ob g i 3^36ihob 6i6n036ob Ri3mu336ob (biQ itf
Oii3g i3nh330ig 3ihn03&ob d0O3h gi5hm33&ib yfob ig5ogo) 83633016001. 5 ih -
00360b 3ibo3ho 5i3mymcgi 3o3go6ih3ori6b b33(5oi03ho 3oht>3036ooii g i 6 3-
b;jb36noi (6ib. 6 .74 ). 3b 30(1^30360 ijô^hig 5i3oiyoiqj36 3iho036b, hitf b^pb
}6ym6b 03010103680 3iho036ob 3m6u36li)hii500b 936ihfi3636ib 5 ^ 3 3 3 3 0 qjihfc-
03680. ^ 30 10 30(1330360 ^30 1 3i65hob b b 3igib b 3i 3y36ih3b (815. A vicen nia),
Tamarix-nb bib3on63bb, Glaux maritima, Plumbaginaceae-b ^>01503(101 bibQonb^b,
Sigmqjn^h 3ih(330m3636b (Spartina, Distichlis ) g i bb3. 89hi0 h35nm 636an
6ih(i303630o Arteplex -ob ^ 5 0 3(10 10 bib3ri6ob g i Halimione-b 5i6m8ymcgn 6 30 3-
b36n gogo himg36ro6ooi i5h n 33636 j0mhog36b, 3 3 3 g 3 5 o(?3336oi6; bi6iU 30m g
5i8mo'bhg36i ibi0O &3b3b36o. 3iho03&ob5i 6 5i6oii3ob3qj036i 93od03&i i5 h 3 0 i3 3
Sa lt re sista n ce
I I
[ Salt regulation ] [ Salt tolerance
------------ 1 ------------ T — i—
Salt exclusion Elimination Dilution C om partm e n tation Tolerance Tolerance
in p ro to p la sts toward: toward:
Filtration and Leaching of Salt succulence Ion regulation Specific ion Osmotic
transport barriers shoot surface effects on effects of
Redistribution Salt accumulation
- in the rool cytoplasm salt and
Salt exudation by in vacuoles
• in the shoot oxidative
specialized glands 8tress
and hairs Disturbed
ion ratios
Shedding of
salt containing
plant parts
65b. 6.73 Q*8o*3;j&nb b^^bntagdn l^cynajn^^dnb 0Q5h>Qmdnb y T>6!jR636^^6n (Steiner, 1934; Waisel,
1972; Kreeb, 1974b; Flowers et al., 1977)
- 396 -
&363 &c*>030 I 6 r t 3 b 3 ^ 0 g 6d 6 r ) t f 3 &n
ib^m3i6n cgmcn036nb fi:>9m(5336nb 338331116001, hnb 83936*6*0 81536^3 oi*3ng*6

nun03&b ^*h6n 3*hnp3&nb bi^ 9 *mg 86n83630m3*6 himg36m6ib. *3*33 ghmb
bp3&> ib*Q6*'t>h0*. 83cn3nb36nb gngn 36ihnb 9jm63 qjmoiQ36nb ^36*.
ib3cnn dL533* gs 3*b*bnsoi363Qni SiQmqjn^hn hfrt'b3lj x n 0U36ih 33&nbai3nb,
hmj,mhni5** Plantago maritima, TRiglochin maritimum g * Aster tripolium.
9*hno3&noi 2^801633300 b3i30361j3hm 6i

6*hnQ36nb 0mj03036nb b*hnbbn 3hmlim3pib9b‘b3 g i 0 mj,ng36x n * *h* 3*cn
i6bri03^3h (^01036016^3, ihi03g $oi6i536li)hii5ni(b3. 013 jjhQgnb 0oi(530m6i 0ih-
n03&nb himg36m6iboii6 3hcm>Q 3 i (j)x < T'6b (6*b. 6 .75 ) 0 ioin ^016(536^!^^ b*i-
9 im gngn b6nb 2)i 60i 30m6i 9 n b(ji6np3hn*. 9ihn036noi £,*30163330^ b3jjX36\Mh-
m6i gi3mt,ng3630n* jomhngnb noi636‘b3. 050 g ic b b ib n ic r^ x o i 5*0mqjnli)36nb*ov
3nb, (J36n*6 80*83 ig 6(1c'J*9yrT,<33C3^ n 6*3h(JX3& X n 9l536*h336nb*oi3nb
(Salicom ia g * Chenopodiaceae- nb mj*bnb 6ih3m8igj,36036n, (b(?3nb b*6*3nhoi3-
6nb 3(536sh330m6i, 0i6j)h369n - L aguncularia) g i ^^30133 33hi0o h3j,noi636nb
Jb3hm3i0friqjoli,36ob*c7i3nb, hm330oii 3b 6n3i6ai3nb3&i i9i33 ghmb ibibi3b
63*03nbig3 n 336D3^DCn b3i3036^36ob aii3nb363h3636b.
8*hop36nb ljhi6b0fny>(jn*. Na+ g * C l’ &igiig&n036i cgom30i9n d*0n*6 *g3og-
oi. i 9o^m8 8 ihn03&nb 9* p * 0 O j)m6 (536lih 5(5n36ob cn*30g*6 *150036^, &i6bi-
2>3cnh36ncn d0o3ho (jh*6b3nhi(500b 3jm63 0301010369^ 93bid 03&30Oi 801300 8(53-
6ihob bb3303o 9 ioin 5igi6i6n036nb 93833016001.
9*hno3&ob 0 ^ 0 1 3 3 6 5 g i 33(1398 mhnbn b.h“rl»nhnhT&i

3*0mogn(j36n 6y0nb oib9mb3h 68ib 32jh3gnb 5^301 ^019336^(13636 3ihn03&nb
gi5hm336noi 33h3gnb 633690. <bp3nb 6 yi 09(536ih 336n ^h^mhb 06^6363636 9*h-
n03&nb *J^n3hn gi&hoi336nb 93933m6ooi, hob 93g 36*e*l 5 9 *cno 9 n6 *&*6n mb9 oi-
b3hn 3mvj,36(5ni0n (-2 ,6-gi6 -3 MPa-9p3) 3cghm gi6igni, 3ngh3 mj)3i6nb 6y0nb
3mli)36(5ni0n (-2,5 MPa). 9nj(533i-9mj^33nb cbm6nb 6yi09(536ih336n, hm0Q363n(5
0ihn036nb 939(5330m6i gpnb g>i69i 30m6 i 9 n 93hy3m6b, ^h^mhb ^3530^ 3 6 3 6
9iho036nb bBhicgn j>i0mymqjnb i6 03Oi3nb36nb bi93i036ncn. 9gn6ih30ii
6ib. 6.74 dihnQQdnb 3cn9i>

6ii5ni gi 9ih(jb-
36s bjhicnn:
i6n 9*hn(yib jnhi)303&nb
j>m93p3jbn Limonium
gmelinii-b qjnrcnpnb
]3ng3h9nb3n. P qjmhQin;
SC b jih j^m h x n 35^393611;
CC 9i9ij,hm336]po 35^393-
6n (Ruhland, 1915). 0jhj33-
6i b^hiCTin: 9ihncoi>nb
9x^13*^ Atriplex mollis-rfo
qjmoicnb &3b3b9n (Berger-
Landefeldt, 1959)
- 397 -
9 0 D66ff)33J)0 b6 rt3b ^ 9*?8T)&>rfiami>90
fob. 6.75 bjixO®!^6015

bihnbbn gi 0ihnp36nb
im6(j36ohiun>.
Someratia alba-b ibic-
fci'bhgi (9ihj335i
b^hicnn) 94 'bhgsbh^cn
(0ihubo6i b^hiOin)
qjmencaftnb Jbmsncnb
b^îi^tii. PP S^bhnbj-
6hn 3ih36jn0i; SM
^3iX3®ii3lln OotcignQn
(Walter & Steiner, 1936).
fchxjjnijn: Laguncularia
racemosa-b qjmoirocdn
JO^hnb ifT'6U36(jhn5nr'b
giSmjjngjbxDî
gmonpnb qjihcn'b] (•)
gi 6ypnb 3;)9u33pm6i<b3
(0) (Biebl & Kinzel,
1965) 200 (im Degree of succulence
03bihcn!33&nb 0iho0oi6o 6y03&nb tbm6i0n 0m(bihg yi3nboj3h g i 600130 6 y i 0 9 l53-

6sh3369n 5i0mnymcgi 0ihn0030u30O Dijmiio^bn.
q ^ C ^ & X 6 n !9 !j)36<b3 9m'bihgn bQ^pQonnb 01536^3360 Djh^gnb 633690 9 ih -
0036b ^111*133636 Qbmpmg 6^ 2)3030 bi'b(Ti3hi0g3. i0ob {p0m 6:>h9mnj063&:> ByQnb
3m^36un!0nb 3ih g 6 i 0 ^ 3 3 6 3 0 , hiy 0ioi bbS^bQ^feib idp33b 03oon3obmb 6n i-
gi&0o mbOmb^hig 69x n B y^n (6ib. 6 .7 6 ). 0ihn03&ob mb0mb3h 3m^36|jn!0b
30b(|)36i J,ibb6fi0n m hîBxn 0gi336nbi g i 6ib9 nh6yi06ig 36nb mbambjhn d i £ i .
(bmj)n3hCTin Sipmqjo^n, &i6b!2)30ih36rrcn 0ihi330ri3636o, 3cghm (j>ihn6ni 0ihn036r»oi,
3ngh3 mh0366ni6n 0ihn0030(5330O ©1536^3360; !0 3^6^2)630 3690 bb6ign
6ib0 nh5ys0 6 ^ 3 6 0 (0ihn03&ob gi5hm 336iboi!6 3haiig) gng hm0b au>0i0fTi636
35h3gob 633690 0ie>i0O mb0mb3hn 66330b 036^83636^00 (6ib. 6 .77). nmB^bnb
h33n9n (Na+/K+ g i CIVSO42" om636ob 0 3 0 ^ 0 3 6 ! 3 3 113 9 ^ 633600) m jib ^-
6oboi3nb g i bib3mb3&nbai3nb b33ijnq)n2>Dhn 6n0i6oi3nb3&ii. 0hi3i0n mh03b6 n i6 n
9ihn00301J330O 0i536ih3 i5hni336b 3qjhm 03,5 N a+, 309*13 K +, 3hoi03b6ni6n
3i0mq3nj)36ob 303£jbm&i 2>n ^cghm 03^ ( i6 (Jm0n himg36m6ncn) K +, 3oph3 N a~.
i6nm6nb bibncn bg3&i 'fly&jusQ C l- gi&hni336i, ib3 0ij,.t Salicornia-b g i
Suaeda -b bib3m6360o, Artiplex confertifolia-dn, 0hi3i0 0ih(33pn3i6b! gi
030ji00n. 303^)3big b30ojivjb ^ (1^ 3 3 6 3 6 Plumbaginaceae, Olantago m aritima,
- 398 -
67)6066030 3 6 d6 P)6360
■NaCI content of the soil

NaCI in substrate [mM]
6ib. 6.76a,b Ssomcgn^jbnb 35h3enb 633680 mbdmb^hn 6633011 gi3fn>pne3&x3d* Sihncjbnb $(*16(536-
eJixjn^'t'3 (Walter, 1960; Rozema et al., 1985)
Lepidium crassifolium, Tamarix-rfo bibambobn g i bb3i mho3&6n63&n. shorn gi

n9i33 53ihnb bibambjbb S^ndpjbi 3jm6g30i 0ihnco6nb ^ 6 ^ ^ 3 3 6 3 0 0 e^b ^SD -
bnb 36ihn: S also la kali ifchr^jbb K +> N a + g i C 1'> S 0 42', S a lso la turcmanica -
Na+>K+ g i C 1"> S 0 4 2’ , bmpn Salsola regina - N a+> K + g i S 0 42'> C 1'.
]jh3gnb 633660 0ihfiQ3bnb g i^ h r^ b n b 3hpi(j3bn biSnShmjbib ] j 06 nb 603-
cnn^bioii U3C^b; SiQmqjn^ban j& br^bnb 303lj3bmbi nrn63bnb ^mjbnahn 0mj-
0393bnbig0n ih ihnb ^qjhm 6ii>e3big 9&hd6<nbnih3, 3ngh] 3i(ynq)mb3b3n (j>i0m6-
iiOnbn: 030bhi63&oii6 gbi)i30nh3bx‘n i»iq)-i<b3bn). jjhQgnb 9 n{,6 ncn 930fnp633pn
Salcormo prostrata Puccinellia distans
-u Oxalate
<0
Q.
5
-3
c
<D
■4—>
O
CL
-2
o
&>b. 6.77 Salicomia prostrata-b (Na/Cl E
SiCm^ryjn) 9b Puccinella distans-rfo cn
O
(K/Cl {ijviytiSipmqjnjyi) qgrooiQ3&nb
Sjhjgnb 633611b b»3hown mb9mb3hn
3r^j]6(yu>cn (Albert & Popp, 1977;
1978)
- 399 -
3(3066fo0060 U6 ^ 3b7) ^ aânaâm aao
3*ho036ob 0016360b *801(16330010 6*g*6*6o036ob 83833016001 31101^0130*^8^

833d0o* fcyjdcmb 3*ho036ob gog ])0i6(53613hii5036b 5*1139m3 o. 3*ho036ob 38363-
boi6* 5(1013936* 3*330193680 (838133300160010-83^*60^80). ^n^miQiba* g * 5 *6 b v
3301(136001 jQmhoi30*bli)36n *8nb 5*801 b*5hd6oi60*g 5*60^30(10136* (ybh. 6.8) g i
Ijbhncn 6.8 9ihnpnb nm63&nb J>i6i6nc]6i 340 9nQd 9"3 NaCl-nb 9;A330C b^db^hi^'bj 9m'bih9
Suaeda maritima-b ojpicnpnb Lîgibbjh
ynd5m6;)6^n 3jh3Qnb dr^-jcmbnb 3hm3mhjjn:> (% ) nm&nb jmSôSiihiijris ampn S’3
Na+ CT
11 116 60
jQmhm30ib,i36n 1.4 104 98
3*i3mon 81.6 494 352
3jh3gnb 539x0 6 194 138
383*9019 361303360b bnbli)33i'b3 801^8390 8*hn036ob b^h3bo (1600936*. 8*ho036nb

01*309*6 8(^150036* 6936* 3386h*636ob iii<33"i 't|36nb 83833016001. om636ob ^h*6b-
Soihôb* g* 3 *oin 5h*9036^nb 8 3 ^ 6 * ^ 3 6 3 6 0 ^ b*ôhoi* 36311600b g*6*b*h336n,
* 8 n(t)fn3 3*901030^360 n8ymgj36n*6 8398030 b^h^bob 30(10163680, 8*oi 3633b
bhgob, 5*6300i*h3&ob* 9 * ^m6i)3h36ôoboi3ob b*ôhoi *bo8o0*^3&ob 6*6n0ob
5*8011(3636* *8 3hoi6033ob j^ ib i^ h x ^ Q -
3*ho036ob*g8n ,yTi03hi6vjo>6*
3*ho036ob*g8o ^0103(1*6^016* *hob ( ^ b o b ^ ^ x ^ 11^ 3hoi^)oi3Q*'b83hn 3018301-
>36,511, 303(101 30163(13^39*9 ijo - 3^36*hob 36*ho 3 *ho 03&ob b^h^bob 3ohoi6363n,
3 3 ^ 0 013 6*3936*9 5**630^(1*90^ (o3 nb 3obx3non 013 hoi330 b*b3oi6*boi*6,
^0130901*6 013 b*boijm(jb0Fi qj*b*boi*6 63*j3b 6*^83) 3*oin îhymqjnoio oib3 m-
b3ho 9* ^mjbojgho 5*3036*.
*hb3&oi6b 8*ho036ob*g3o 3*ph3b*g ^0193(1*6,3390 oih6*6ob336o. 0301,301*3-
,501,3(10103390 3m0^rnb*6n Dunaliella salim a *hb3&oi6b 8*ho0cn* j)oi6i336lj)hoh3630n
bb6*h36ob 30(10163680, 3*90103003(10 6*j(j3ho* Pseudom onas salinarum 9 * b*q33*hn
Debaryomyces hansenii <336jynm6nhQ6nb 36*hb 06^^363636 8*806*13 i n> ^0193-
b*u NaCl-ob 30161536^(1*150* *(p633b 20-24 %. <b(j'3ob 6y*08(536*h336ob 3 *h-
o036ob*g3o l5Oi03h*6(i x rn^ i t>Db,i*g 833b*6*836* 3*oi *9509^81^01033080 ^ 3 6 3 -
00 8*ho0o*6oi6ob (53*036*gm6ob *330olj3g*b (6*b. 6 .78 ). *8*33 ghoib, 80^1533*-
0mj(533ob boi6nb 33(103*9063(10 9* b3&0o,joih*03ho boi6ob b ,536013*0063(1 6y*08-
1536*h33&b 8oihob 830608636* *8j>*h* 5 *6 ^ 3 *3 3 6 *.
3*001030,53(10 £3(1^0013*60 8(536*113360 1^(19360*6 2-6% (9* 83,50, 0830*01*9,
20%-3g3) 8*ho036ob ioi6(536^h*(5oob 8^0163 60*9*580 g * 833890*01 NaCl ob3oio
(1*019360161101 9*5(101336*, (1018300)5 8 3 3 ^ 6 *8 3 6 * 10% bb6*hb. 3i0mqjoi636ob 33(13-
9360 0(3133360*6 3333 1% bb6*h8o (171 880190 35''), 303(101 h3'bob^36^xn 8Di!p6-
,5360b 1,30^3(1360 0,5*636 200-300 88oi0 3016^36^(1*15036^
3(101^0130*^83(10 1^5(13^3(1361^ (1500360, 6ooi8386h*636o) 3*ho036nbg8o ^019-
3(1*6,530016* 3^(1363301301(3000* b(j(i3bnb 3hoi,j3o63&ob g* (5oljoi30*'b83ho oib3 oi-
b3h*0 *^03(10 3018301636,5360b 33833016001. 3*110036001 0*(i)3ohoi3ob ghmb 1^113-
- 400 -
636056030 li6 6 3 U7)C?0 ^dfincnO&O
Species
C lodophoro splnuloso
Bangio fusco pu rpurta
Porphyro Icucosticla
Polyliphonia pulvinala
Chailomofpha linum
C troB iu m b c rn tri
Anlithom nin cruciatum
Cafclhommon fra n u lo tw i
Clodophoro ulriculosa
Cl. proli fera
C l.ra m e losa
Callilhomnio# scopulorun
C e ra a iu n strictum
Placamium coccineum
m io p h yU w pu tclatun
6ib . 6.78 W jpDOinb 6y io 3 |j 36ih 33&nb b ^ B b n ^ h m b i Q iS Q id ^ n b bb 3ig it> b 3b bihnb bnb 9nb ]p 3ncn.
cn^cnhn bhg: i g i i ^ n i ; 3 i3 n ih ]: 91(^3331 2 4 bcn-nb 339034,. 6yic9u36ih33& n, hm 03(}oii 3:>&fvji^3-
6n piSQiSobnb gngn ojODiiD^nîmcn 5i0mnhfi33ni6, 3^301 35336016 hm&mhu 3n3m,yn63h, ^333
3n33 h(5m63p 3nhm&36b, 3 ngh3 3 mJ}33> 3 nj(j33nb 3&3 n)
3 ubm 3h36n g i b36Qnonnhi£«3hn 6yip 3 |}36ih 336n (Biebl, 193 8 )
h .g . + V-CH- o 1 i
3 h2c. ,/CH. o
H,C^ V CH, 0 X .
Quaternary ammonium Aminoacid

compound (Proline)
(Glycine betaine)
C H pH CHpH
HO
HOOH HOCH
HOCH HOCH ho/ h ho\ h
61b. 6.79 3ipmq)n(j 369n mb6mb3hig

HOOH HCOH h-sh chj4 h
HCOH HOOH
iijn^hn mh&i63on b36b(i>i6(jn3&nb CH pH CHpH OH H
3^5^0013611 (Popp & Albert, 1980;Aspinal
& Paleg, 1981; WynJones & Storey, 1981; Polyols Cyclitols
Popp, 1984; Popp et al., 1984; Jeffrey, 1987) (Sorbitol) (Mannitot\ (P/n/to/)
- 401 -
9006663050 9«?an8M 3n 6.>30
bnb 5hm^3n636nb bn6cn3'bn bgQ&i hi9g36n93 biicnnb 5i 69i 30mbi9 n 5 3 6 3 6 ^

6 ^ 33330 0 366360b iJijn3ii3nnon [36]; 9^9536000^ id (jn036nb g i 53i03n b i g i
bntfbnb 9 mj>nb ghmb bn6oi3'bnh363Qn 150 936 ^ 9b5i3b36i, h^j 9n3CTincn36b t,rnig-
iS ^ n i^ . 93orii3b36ign, 3.11. ihi^m jbn^hig 9mj03gn bbBign mhj>i630n
6i3hori36n (6ib. 6 .79) 3^(1363309010336 6m6ib6mhm6ib y n ^ m ^ g i^ ib i g i 33h3gnb
mh&i63036b 9 mhnb. 3i i 3m03&3n nm636nb ij^n3hn gi5>hm336nb bih3<b3 un^m-
30ib9nb 9n9ihcnx36ncn 6ih9mnj9636i digni6 d0ii3hn 5hign36vj36n; b6mh3g i 9
j)hign36(j36nb 5ibi63n^hi03&pig bmhun30g36i 33h3gnbicn3nb 33636300, mb9 m-
b3hn di0nb 9jm63 jjm93m636(j36nb bn6cn3<bn. 9(336ih3369n ihib33ijnq)nj)3hn b^h ^
b x n h3ij(3nnb (9ij>., 9ihn036noi gi^nhcr^nb, 63*03^ Q* yn63nb 3nhm6369n)
2,i9m30n636ig 93ndg36i fiinoi3i0mb nbsoin 6i3hcn36nb bn6ai3‘bn, hm&mhnuii
i9n6m9gi336n (3hm0n6n, i0i6n6n, j)03lj i 9n6 n, ib 3ihi{,n6n) g i i9ng36n, i 9 ni^nb
6i3hoi36n (63^06 36 0), bb3gibb3i 9ijh36n, 3m0nm036n (9ij>., bmh6n^n, SiSn^n)
g i bni0 rlii36n, hm930cni cni6iqjhgm6i g i 930ig5360m 6i 9(j36ili3cni ‘bm5n3hcnn
563gjnbioi3nb qjhnip bisynqjnjj^hni (9ij,., 9i6{>h369n 5i6bij>3aih36ncn bn6-
oi3‘bnhg36i 9i 6 nli)n g i 3n6n,jn). fcihgi i 9 nbi, bb6ign 6ib9nh6yi06i036n g i i 9 n-
6 m9gi336n 0 ^ 3 3 6 yng36bi g i 6nm9396hi636b nm636nb îh6n ^ 6 15 3 6 ( ^ 15 0 3 6 1 ^
9^363 5i3036nb5i6.
6 .2.5.4. IXJPram M hfl 9l5D6^hD06nb aihflCD&fltoean a 6 h d 6 n 6 flih n 6 i
ihi3i0mq3n(j3hn 91536^3360, 3.0. ^ 30 136 0 , hm0036o(5 6mh9i 03h 3nhm6369o

90 i93 3i6nli(i lj3&9n ih n‘bhg36oi6, ym330ot3ob ih ihni6 9iho036obig9o 9j,hd6 rn-
6nih3- bp3ob bi6i3ohm36oii6 ig&ogo ij3 b 9i536ih330m6ob <bm6i03hm6ib, b ig i(j
53b3g3&i 9ihn036nbig9n J)i6bb3i 336x^>g 96hd6m6nih3 bib3m636n g i i5h3cn33
3j)m^n336n (9i&.r P lantago maritima-b g i Juncus bufonius-nb 3^01^113360) [105].
bmqj0nb 933h63m6nb, 936i(p3m6nbi g i 930>y333m6nbcn3nb1 i3Ci!)3*13Cn 9 ^ 36i-
h33&nb 9iho03&obig9 o h3'bob(jl36(i)30m6ib gngn 96o33630m6i ij3 b d)hm3oi36nbi
g i, 5^6^^301(136001, b36^hm3nj)36nb g i90i 936x n 6nigij>36ob 5i9my363&nb
3nhm6369n. i 9 3hm6030nb 5igi^hi9n fibhoixni b303ji5Oi, £,363(10 o6go63hnb g i
bb3i 93156031136360. fcil^ m b 9x530b ib33j^noi, gngn 96o33630m6i 36o^3&i 5 ^ 3 -
6 ^ 3 ^1130133 9myhn0n 9iho036obig9o (NaCI, M gCl2, CaCl2) 9g&hig 91536^ 336 6.
bij)336n g i bibngmb3 9^36ih336nb 9ihn036nbig9n h3<bobvj 36lb X rn&n'J 9^ 33-
6360360 9 mu3930ni 6ib. 6.80. ijbh. 6.9 9m1i>i6n0ni j)'b36nb 3o(Yb3 g ib ih j,i3ig
^ 9 (^ 3 6 3 6 3 0 0 b336nbi g i 63fij36nb 9ihn036nbig9n 9g5higm6nb 3ihi93(jh36o.
9ihnp3&nbig9n 9g&higo 91536^3360, 9j)hd6m6nih336nbj,b6 5i6bb3i336ncn, g)i9m-
bi3i0b 3m30m636 ih i 9ih(jm 393i0mg 9iho03&ob îh6n 3m6i536(5hii5036obdi6
gil53i9n, ih i93g bhgnbi g i 5)i63noriih36nb 3hmi53b3&nb 3q)hm 933<b(j'3gi3ig 6ih-
9ihcn3i9n.
5'b3&nb 3nhib 9 mbihgn 9i536ih33&nb h3-30^00^0(136;) (j,i9mi5riub036i) o9nb
2>i0fn, hm9 nbn6n ym330 ,b i9oiihb 33hnm gxê n^3nhcn36ni6 9ihn036nb 5 i6 -
6633nb 93936ig, bi^9img d630ni. 6i03j36ncn 9gngih h35nm6369nij ^n 9ihn036n
ihibij)3ihnbig nh3ub36ni6 6nigi5ngi6 63n9nb 9n3h. 9ihng36n i(p633b c^)hm369n,
y0mhj)369n, qjmoi0369n g i n6333b 03bi9R633 gi'bni63636b, nj,n, j,ihgi i 9 r»bi,
ib3b(!)36b lb i9cnhnb 5i9m6hc7im6nb 3hmi53bb. i 9 9m x36>boii6 6hdmgi 93nd03&i
9ihn03&nb 3qjhm hi(inm6i03hig j>i66633nb g i g i9g6ihn 9ihngni6n 6y0nb5i6
- 402 -
536066030 U6 fnJUT)ggO gadacifnoao
Salt toleran ce Poor Moderate____________________ Good

Alfalfa Cynodon Dlsfcchlu
dactyton stricta
Fodder p la n ts Festuca
Clover
Su n- •*]re M illet Barley

flow er wi M at G round nuts
R e id crops
Oats Sesam e Sugar beet
Maize Cotton Soybean
Radish Ftotato Torrwlo Beet
Sroodbec
V e getab les Geleriac "C a rro t SW <& g S ,f.
French 100 Lettuce Asparag 19 Broccoli
bean Cucumber Melons Spinach
Apple fig D ate palm
Cherry Pom egranate
Fruit Peach Olive
Apricot Grape
C itru s
Salt content of soil CL2035 Q65% dw
Electrical conductivity i 6 S O 12 U 16 W
*1 the saturation extract m S-cm ->
Growth response LDM

Leucophyllum frutescens i ~ 1»13
Sycygium paniculatum I ~*)>13
Pinus halepensis I
Chamaerops humilis I .. ^ 9
Raphiolepis indica I Km zzzt: -7! 9
Pinus thunbergiana i '-I 9
Arbutus unedo i tas«a. Z1 8
Podocarpus macrophyllus i — w g J B 6
A b e l i a grandiflora I ESMff r; 4
Photima fraseri i kzssi. i
Mahonia aquifolium I 4
Cotoneaster congestus i vmm?™ 4
Soil salinity 0 1 2 3 4 5 67ms ■cm'1
Growth reduction □ 0- 24f .§ ^ 25- 49Y .£ ^ 50- 997<>Bl00%
6ib - 6.80 bb3igibb3i 3>>m6m8ni3hn 81536^336^ gidQb&jbnbiedn. 'bQgi 6ib:>bn:

i6h,nAXiJ3h36n. bspnb iXiSIjhabn, &>(rfi3xn iXibOhD&n (Kreeb, 1965; Cox & Atkins, 1979);
e^bi’bn: eaynh^yrjpn (*> bj 61336^3360 (Francois & Clark, 1978). p:>6g3:>oynb
ihj(V>3ij3hi9n jy>9my363&;>gn 3u36:>h33&nb Smtuu^S^bn nb. Banick (1978)
9(336ih336nb 03bid036Qig bhxymgncn 3ihi6nh36nb 63933016001. 6n:>gi&nb g >

q p h 3 i8 ^hcgnb 3nj,hmbt|m33Qn <336non 9 31^ 0 36 5 93i 9ijnhmb 6ihn036nb gij,hm33-
6 i 6nipi59n. 3b <336:1 p i ^ 3 0 1 3 3 ^33^36000 <bs8cnhnb gibibh30b 369s 0m(5n0-
93b 6nigi5b, bn0m ynaSmb^nf^nb Riîh36i ih 93nd036i. gi'bni63630n b3
9 j336^h336nb 9 ^ 3 3 6 3 6 ^ 03bid0363Qni 8bm0(ng 8ihn036ncn bi03i0m 9 9 ^ 3 ^ 0 ) 3 0
3 i£>nl!)i£)69n 8 ib 936935. hijj bnigi^n 6036nb 8i6dn0<b3 9^83653936^ 0063606
3n6m 0^303036nb bb6ihnoi (hm63cn(5 93n yi3b K +, Mg2+, Ca2+, N 0 3‘ g i SO42').
- 403 -
8006660060 b&fnOb'a^ aei& rm foom .iao
Ijbhncn 6.9 gido:>d3bntagdn ihiîcrnqjry^hn bô&nbi gi &3fr)3&nb ojihgmbncnn b]6bi>j3hm6i

(ojih<7imq)moip<n3>6n b^jbnbi gi djfrjjfcnb SihnpjinbbgSn b36bryj3hm6:> tig b x 0^ gibi6ynb3n: 0.3
- 0.5% 90hip 6m6i‘b3; b^Gbn^hn 6n63m3636nb: O.fi - 0.4%; 8ihne:j6nb:>p8n dg^hign ajbhcnrygmcrv
pm3i5n bjjftnbi gi d^fijj&nb: 0.8 - 1.6%; h3‘bnb&)6£x;n 6n63fn3636nb: giibo^dncn 0.6%) (Sucoff,
1975; Meyer, 1978; Carter, 1982; Dapier, 1991)
e*8p*33bnb:>edn bjOxxohn es9Oi33&nt»e0n 33esh36ncn h3fenbts361s3o°
(griOTpai}336fi b^^&n gi infijjbn
Acer platanoides Acer negundo
Aesculus Ailanthus
Carpinus Elaeagnus
Euonimus Fraxinus
Fagus Gleditsia
Juglans Hippophae
Ligustrum vulgare Lycium
Platanus PoterUilla fruticosa
Primus serotina Quercus-cfo bibjmb^&n
Rosa rugosa da sxva Rosa-b bib3(n636n
Siringa Robinia
Tilia Sophora
&fog963»63 qjriai(>n3s6n 93hj6ru6n 81536^3360
Hex-nb bib3m6]6n Coccoloba
Ligustrum lucidum Ficus-nb bib3n)&3&n
Mahonia M agnolia grandiflora
Trachelospermum Nerium
wiwvovnebi
Abies-nb 633^ bib3(T
i6i Juniperus chinensis
Picea-b &33hn bib^mbi Picea mariana
Pim a strobus Picea parryana
Pim a silvestris Pinus halepensis
Pseudotsuga Pinus nigra
Taxus Pinus ponderosa
6 .3 . ^6onhn3n636D hn b i!)h 0 b 3 c n q j^ m h o & n
6.3.1. ^ a n ^ n b u ^ ja n ^ n b n b 3 dq 36^ q 6}hamj06ncn

a n a e ^ a c^ n encnnDhObObnb 6 ^ a m y n ^ w aaaebnan
3 0 i i 6 0 ija ^ q jn ib n b o jD h n a n
9t>3, 3i]hn, 6yipn p i 6ni0i&n 9(536ih3b 3^363309^10336 ih ! 9ih^m 363(1-

^nncn, bij)336n 6030103(136 36001 pi b30biyh3pn 3nhmti36001 - 9u36ihnb j>ih39ro9n
ihnb bj)h30i33 3m96i 9o !3n (qjn^m^mjbn^hn) p i bs9 n3 yn6u36l|)h:>un36ooi ihb^-
63011 9^363 6i3hcr>36n. 63636(103 3nhm&369n 6ih9 mj96n0 Jn9n3h 3 0 3 9 3 6 ^ 3 6 ^ p i
6i3hcn36b Smhnb ihnb nb3cnn 3mlj36(3n3h:>Q b i9 n9 n 6i3hoi36n, hm^mhnipi: 33CJ>i-
63(10 6ih0m3m&nb SO2; ibm^nb nrjbnpn, hm93pn(5 6ih3mn;J9636i 9nihm6x n P3&-
n^hncgnîjnnb Shm^bSn pi 3030b phmb; 9 ihn03&n pi 3i 3h9n 936m6nQn 9^ 3hnb
6i6n0ii36n. ip i9 ni6nb b ij9oi6m6nb 93Q3&ip 91536^ 3^3 9mJ93P36b 0hi3i0n
3Ubm 6ih0m3 m6nb 6i3honn, hm9p36obip9o 9u36ih3 (33(1 inp33) ih ihnb 8 3 5 3 -
- 404 -
a6 m r t m n & 36'3(f>o gid&nrta&o
Ubtv^n 6.10 j^3m0qohm8n JnSn^hn >>ma}m636vj3&nl> (jninohn im5u36(jhiiyi3&n gi >hba6ni6nl» 3^3-

6n (Freedman, 1989; Lahmann, 1990; Legge & Krupa, 1990; Kuttler, 1991)
Jnanjhn jm93m63^}n (jnSnjhri (ppm) iî^mbcgjhmgn ibba&mix*
bjgjcni 3i3h3n JyjhSn bi66hdcn3m6i
CO, 340 400 2 - 6 6300

CO 0.1 40 - 70 2 - 6 0133
so2 0.0002 0.2 1 - 10 geo
HiS 0.0002 0.5 - 2 g(p3
nh3 0.01 0.1 2 - 14 gp 3
NSO 0.25 4 - 1 0 63011
NO < 0.002 1 - 2 3 - 6 geo
no2 < 0.004 0.2 5 - 10 eeo
03 0.02 0.5 ge>336 n - 0133360
CH4 1 - 1.7 3 4 - 1 0 630*1
bb3i 3nphnj)ih6r'i6i(j36n < 0.02 0.3 ~ 2 eco
36300 (ubh. 6 .10). 3i0hi3o 6iohj,3iho qgn^m^mjbn^hn Bi^hcnn j,i9 mnymcgi ^ h -

9n, Byipbi g i Bnigi^Sn nBgjb^hnnb, (jhi6b3mh|2)Ob 0oidhioi6ob, jnQrvjhn 603010-
31136360b 03lî5ob g i J,i6bi2>30ih36ooi, 6nie>nb3x n bi63i3o b 5i3oiy3636nb 8393-
jp g. y33piqj3h i 9 bb 3 6 ^ 3 6 ^ 6om(jng35nb g i bib3J36nb ihihiuom6 i 03hn &i0oiy3-
6 3 6 ^ i.'bmôb îh6o hioig36oi6ob 3 3 ^ i 6 i 6oigij>bi g i 6yo360o, N H 3-bi g i 0]oii-
6nb 5i0nymqji 3i 3h3o ^0 1330 360 6 3 ibm6ho30 3018363606 33Q362>e- o6(j36bo;)ho
bi^hiSb^rnh^m 3mdhioi6i, o6g3bl!)hnxo i3iho3&n g i b3COi363hig 6^801633300
bi6dh36o, 018360 - y33piog3ho 3b 06333b 5ih30mb gi^n h crîb 3imbobv|)3336nb,
2>m6i!)0636j)360bi g i fcprnbicoho 0 ib 3li)i636ooi.
3oij33g36i l5iCi33Cn j)Oi6u36l5hoh3630o 3 oi06i 3p i3o 6i3hoi36nbi, h018036015
06 33336 3|536ih33CO biqjihob gi‘bni636ib, dnhooiigig, bo3h(530o 0 3 'b p 3 g x n:>:
SO 2 g i 3^0153609360 5i0oioyoi<3i b i6 3i30 0oi6yoi6opoi636ob, 8 3^03^ 50 300 g i
33hi3oj)3Qo o6@3b(j)hoob bi6ih8oi36oii6; 3do03 0 3^ 0 36 0 g i 03îpoiog36o 51101303-
6 i 5i96036g 0oi6yoi6o0oi636oii6, bi6ij>3336oii6; 33ihftobo 0ig633pob 6iyih36-
o ii6; j>i6663300 0ihoc3&o, 5i0oi6i6oicj3o 5 ^ 3 6 0 g i 33(1601500360 5hoi303&i 5^ 3-
bnb 3oh36oii6; 0oi86i8pi3o jo0o*)ij)36o oii3b oyhoi6 6yp363o, 8go6ih3363o,
^636b i gi ^ 3 3 6 3 0 .
y33Qi'b3 b3 ohig ig50p0 ij3b 3hi3i0O 8i363 6i3hoiob j,oi36 o6 oh36x 801^8393-
b ib ; ib3 3 ij,., 3hoiig 60110393636 qgo(jmjo0o3ho oijbogi6^36ob 2)Oi8303;|bo S 0 2.
5^'bnb36ho 0i363 6i3hoi36ob 8oij33g36i ih ihnb 83801qjihjjODC0 8^hôi i^8oibqj3-
hmoio, boiooi booibnb bibnoi shb3&xo 6i3hoi36nbi - Snghoibqgshoioio; i8nb 8ij>i-
Onoioi 6oigij,obi g i 6yp36ob 85^336001 9^30(1013^ hiu j,i8 o i6 3 3 x o i 8gi33&ob
gîoibouoooi g i 3do03 83(i)i036nbi g i bib3jo 8iho03&ob 6igib3pncn 6oigi5n9i6
6yo3&3o. i5bCi»3X n 8 * 3^3 6i3hcnnb 8oij83g36ob 06^36^3(10 i3033^036o b fiiî-
h3bob 330036. 83(j6o3h36ob y3higc>36nb 1536^(180 hfi3&i 8i363 6i3hai3&ob 3(1010-
3han0oij83036i g i 5ih38oib bb3igibb3i bq33hoib 3(10103(10^3036^
0 i3 6 3 6i 3hc7)36ob ihb36oi6ib 6ih>)333Q iQ6O0:>b g i 5ih2)33X OôiSo 36oig3&i
n8nboi, bmom 0 i3 6 3 6i3hoi36nb 5i8oiyoiqjib - 30nbni. 0 i36 3 6i3hoi36ob 2,016536-
^hûoib 3601936b o8obo36ob >1016(536î(5ni, 050 5b8oiobi^36i 00115x0160010
6npnb (ftibbbob ou>6iq)ihgoi6i - ppm = 1:106 i6 ppb = 1:109) i 6 80115x0160b
- 405 -
9036i>ff)0060 uertDbTi^ a ^ a r m f n a n w a o
Ubhnpi 6.11 20°C-'bo ij>3mbq)3hmb {)i‘b3&nbci3nb J>igi9y3i6n jhoijxD^ 9fTH53C<T,6nan 0 PPb =

1: 109) gi &hi3n33i)hx (H 5^ 3"3) ini6u36^hi(jn36b dmhnb (Lendzian & Unsworth, 1983; Daj31er,
1991)
5 itn b qjmhSxi 5>g>y3a6i ppb-gi6 (A 9'J-9n; 5ig»(3i5> p. 9'3 es6 ppl>3n;
ppb &i9hi3Ce3U } i jh S'3 5>s8h i^ e 3b:
so 7 2.67 0.38
h 2s 1.42 0.70
nh3 0.71 0.42
NO 1.25 0.80
no2 1.91 0.52
03 2.00 0.50
c h ,c o o -o -n o 2 4.37 0.23
c 2h 4 1.16 0.86
HF 0.83 1.20
HC1 1.51 0.66
3(10133060 5(^36390 himQ36m6nb (65 9 "3) bibncn. 013 535n6vj)3h3b36b 58 603-

cnnQh^b^bnb 659m2>3C33i 6nmjn9n3h 013 qjn^m^mjbn^h 5 ^ 3 ^ 6 0 , 65010 2,1^6(336^,-
h(jfii 369b j)b9 mnb^fnb 9 mQ5h30 qjmh659o, (159556 jn9n3hn 653(101360 33(13930
9^1^9393636 9fT'C3i 3036nbi 95 nm636nb b 5bnoi. (jbh. 6. 11 6(1115383005 38608363-
0(^35631*1 85363 6531101360b 559501390b 035^(11(1360. 353(1nb 958560683(13639(1
6i3hoinb nb bimhn36liX5nm (150193(1165, (101839015 (16333b 95^056365^ 95069365
bi3hoii3mhnbm Oih556o'b5i5036ob5 (WHO - 3568hai390i6nb 0bmqj0nm 0155601351505;
IUFRO - ^ynb i3CD3nb mh5i6n‘biijn36nb bi3horu>3mhnbm iÔnhn) 95 j^Qijnb 59-
50901611030 8oi53hoi6 nb 9 n3h. mqjnijnX'lDh^ 9595360011 (b p 3h30n *)|-ri6 |536ljh 5i5036n
nd033fi56 66019019 ^01559 6}h0m9536ib, h59556 615365(13015 ^ 3 5 9 5 ^ 3 5 bib3m65
95 3060 558010(16335 556^35336390 65(18601605(1016001. o9 nbnnb 65^06593(1 ^m6-
l536(5h5ijoob b5bncn 5333906365 80121933590560 2)^6536^5(30300 3oj)36o (IMKk 30
63015693 3nj)OC7i) 95 3qjhm b565hd0O3n 901^639365 (IMKD 24 b55cn(b3 6565(18-
9030; ijbh. 6. 12). 50sbcn56 952,53801136001 3695 553001359066060101, (1018 65363
653(1013606 d0fi3hn o8nbo36o 615365(133680 0633336 b6h5C3 95 9do93 95130563-
636b, bm0m b565hd0O359, 955(156 95659 2>fT'6(j36^hii503630 80^8390 653hon36n -
jhm6o2>30 g5h(p33336b.
Ijbhncn 6.12 bb3ig:>bb3i &i(bnb 3ijbn9i03hn 1^ 16(536^ 1^360 (|1 fch S-3) (Jageretal., 1989)
6i <bn Jn^hn *,m6{536ijh;>i5n i bscfppm diS 3363630(1 js'bftb 33h9i 636(j 30<1 <y>3mymq)nb:>b
3>fi336360360 '\ gfpnbo^nb (8 - 24
(3040 BcnAxn) 33hnmgnlxn3nb
j 6 1 6pnbot3nb
HF, 3 1-2 0.2 - 0.4

S0 2 400 70 - 100 20-80
no2 200- 6000 70-100 30-60
Oj 300- 500 80-300 50-60
NHj 10000 600
- 406 -
a e m f n m n a o G ^ o U6 6 0 b7)c?0 ^d & n rta & o
6.3.2. a n 0 6 ^ a W 3 (l 6:)OhOD6nb <b D 0 n j0 D e 3 6 i 0 l3D 6 ^ h Jb 3
bibniimisbcri qj^jynjbnb Q b h ^ ^ n b bihnbbn g i 9 3 ^ 9 ^ 6 3 3 gi'bni636!0'i!

6ih0m9m6i g i9 mi)ng3630n! 633b b b 3ig ib b 3i 3nhm6i ‘b3, 5 5 ^ ^ 3 0 ^ 3 6 110 1: 9153-
6ihnb bib3m6i'b3> “bhgnb cgmh0i'b3, i b i j ) ^ , ij^njm&nb q jib ib i g i 3168(1013-
pmbi'b] (61b. 6. 0 1); j ^ n g ^ h g i 3Qi<33h 3nhm&36b ! g i 9*36 3 6i3hoinb jn-
9n‘b9t>3, y n b i^ ^ h iu n i^ , 9mj93g36nb ghmbi g i bi65hd0n3m6i‘b3. 'bm5fi3hoi
930oib33i9n g i‘bni636nb bihnbbn 933bi& i93& ! 8^363 6i3honnb n9nbnnb ^*161536-
^hi(jnib g i 9mj03g36nb bi6j|)hdpn3m&ib (6ib. 6.02). ib3cn ShcgrvjQ gi0m$ng363-
Q3&ib igftnpn ij3b 9bmpmg &ih2)33X qjih{,p369n. 3howib 9bhn, 3b qjih&03&n ^33-
3ngi6 039mbi'b(3'3h30ni jjmB^B^hiijnnb n9 t>(P3h3cn himg36m&noi, hm90nb
phmbi(5 9i363 6i3hcnnb bib&hdc^n 9m j93g36i^ ],n ih (16333b 9(536ihnb g >
bni636ib, bmpm, 03mh3 0bhn3, &ihi3330n 8 ijbn9iQ3hn 2)R6(j36^hi|jnncn, hm9gnb
mg6i3 0m0i^36i(5 06333b ih i3hm3mhijn30ig dQfi3h gihp3333&b. gngn 06n03-
630m6i 55(130133 ij3b 05363 6i3hoi36nb y33Ci>'b3 dpn3hn 9mJ93g36nb ghmb: 31*16-
li36^hiijnnb 3nj)3&n 83!9)313^3, hmg3bi(3 65533611 îbbbnpni 9 ijbn0iQ3hig, 3qjhm
bi9 n8ni, 3ngh3 (j>i9noi. i 9iboii6 3hcnig p i 9nb biioi3&9n 9u36ih3 nb33636b,
h59516 (T>i9nb 3nhm63&9n n^n 6130 3659 ihnb 3jb3m6nh3630n 9105^056363011 9mj-
9303&nbig9n (815., qjm^mmjbngiuni).
61b. 6.81 ajm^mjnSn^hn 5i0m6i6meJ3nb 1b30mj93g36nbi{(i6 6ih0mj96nen pi'bm63d36nb

(pybn^j&nb % ajmcnprib 001300 bxiSnhngib) 5161600361 cni0&ijmb bb3igibb3i ibiyibi gi
5i63ncmh36nb gi'biSn Symqj gmonpjb'bj (Glater et al., I5nii- Guderian et al. (1985) 0nb3p3noi).
80161(530360 gjmoioob qj36j(}om6iC3h gih(p333ioii 9361b36 nb. Omasa et al. (1987)
- 407 -
3(3 0 6 ^ 3 3 6 0 L 6 6 3 Lflg! 3^ s r ) 36f53r )5630
Duration of exposure (days )

Bsb. 6.86 qj^hnb 3o6jj3xn 3m36i3(}i3n jp^bnb 'bjSmjS^&ncn 5i9m6333Qn ehmj&ncnn gi
Jhm6nixn 363611 bb3igibb3i 0(536^380 (McCune, (jn^. Keller (1975) 9nb3g3nai). mbm6nb
gm'bnhjbnb ygjipi 3ihb3C3X n iDOii3h3inl>cn3nlj e* ,33 6tJ36bh30iion3nb nb. Reich (1987). bm^m
b&isjnb 3036b 33hb3nb cjfYbnmcrij^h nb. Van Hove et al. (1989)
6 . 3 . 2 . 1 . £)3han ihbD&Xfl 9^)363 6Mh0136n
SbQhnb 8^363 6:>3hoi3&b 8mhob 8u36bh336obbov3ob 5b6bbj)3cnh36ncn bb8 o9 ob

fcmfcnhgnb mh«)b650, bbm^nb mjbogo. m‘bm6n, 33hmJbrv>i53^np5nljhb^n (PAN) 9b
3b0m{,366ybc6bg36n. 8u36bh3*b3 b^QmbqohmpbB O03j3&b ^>6^30133 b90 bj)0, 6bb0oh-
6ybp6bg3&n, ybm33Q36nb b6bmhavj036o, ^bhjijcn Qb 3(J33hn.
9b<bru>636nb b3hbcno o9obn3&ob 039cnb333b3n 0hb3b0qj3hm3b6o gb bhbb33(joq)o-
i3hnb. shorn gb 05033 0b363 6b3hcno bb3bgbbb3b 8u36bh3<3o 116333b 5b6bb3b33630
Qb<bnb63636b, bnprn 0 3 ^ 3 0bhn3, 3hcno gb 05033 bod^mdo 83od036b 5 b9 m6-
3 3 X 0 nymb bb3bgbbb3b 6b3hcnnb 0O3h. gb'bob63&ob ,jo3o gb bbhnbbo 9mgoq)ouo-
hg3&b bb3bgbbb3b J>bh30m gb 6,^36390 qjbj^mhs&nb 3hcnghmxo 9mj93g36ob 8 3-
935bg, hby bd0O3h3&b dohocnbgo 0b363 6b3hcnnb ^bXD^bb. bb3 ds&.. 0606036001
gbli)3nhcnxn 0l536bh336o 6o08hb0obb gb yo63ob 33hnmp369n lbnb6g36nb6 3gjhm
dQnshbg, 3ngh3 3b68ho»xo 06903(19360.
bhb36m6b gb(bnb63&nb gb6y3&ob b g h 3 x n gob^Bmb^yib 8389350 6086360:
8b363 6b3hoi36ob gb5hm336b 9u36bli38o (6bb. 6 .83); 63033(1x0 36bhob 838ijoh36b
Jbm30p3&80; 3gb336ob 0 m j0 x 3&ocn 5b0m633x o D ^ o ^ n ^ x ^ X o 03360b 9b‘bob63-
6b 6063360b 33xn 36lbo036ob gb6bco b6 9b(?b0O bjjj^mbb (ubh.
6 .13); 03^6(^00^3606 hbrr)936m&h030 gb bbhobbm6h030 15300936360; b ^ s b jp n
- 408 -
.>6m rism r)a067)cn0 b G r t a u ^ o ^d & n rta& o
Gasconcentration (fjg S02 m-31

6ib. 6.83a-c 3 ognb J>i69i3pm6i3n S02-nb bb3i9ibb3i b^hpbxn 3oniy* 6*^3133.
a amjnhpnb 915(1(113361 60633680; b qjm^mbnEionj'bnb n6,j36bn3m6nb 91133101361; c i^hmjbnpi'bnb
iiyi3mdnb 'bhpi (Keller, 1982). S02-nb, 63-nb 91 N02-nb 3<33Jcin ln vivo Jcf’ihmqjncnb
1336(501133 nb. Schmidt et al. (1990)
(jbhnQn 6.13 i^Smbqohmb 9i8i6n6d3h36c36fKn 9n(P36x ,n b^hQbnb 6nmjn9n]hn pi qjn'bnniQmjn^hn

n6enii^nh36n (Hartel, 1972; Horsman & Wellbum, 1976; Jager, 1982; Darrell & Jager, 1984; Weigel et
al., 1989; Grill et al„ 1989; Cape & Vogt, 1991)
n6eny>^mhn gidi&n6cl;}h3&3en u>ei 938(5*361
jBbtfhjbfi
33hm^>nei'bi f2, hf, SO, X
3mpnq336mp mjbngibi S02, NO2, jneh<T|6ib9nh6i936n X
acrwuSia e^nph1^536^1 S02’ NOx’ X
RuBPiih6m#jn0i<t«> S02 X
60^0^ 11393^31^1 S02, NO* X
bj^mjbng gnbd^'bi 83133(10 bic3d36n- O3 X
bih3l>834i6mp(vj36o
ibimh6n6nb 83131 ihib^rKgnahn X
60>J;,o’™,'6n9v2 X
3mpni3n6n ihib33unqi)ni3hn X
30wp36n ihbbSjiyigni^hn X
BsaiSmcrtbQn
ig36npiyib b^j^bn ihib33(yiq}ni3hn X
qjmjmbntknj'bn ihib^nqjn^jhn X
eti^njOhn i8hD5303&n
m3^n>)3hn 18(13330360 03, S02, %i33hn 6103^ X
^^h&npn^b ^b^n djis^hn 6ip;)i)&o X
3mh3m6;)fcnb (j^Bbijjôih^bnoi Qcnnp36n) 6ih8mJ36:>; b36onj3nb n6lJ36bn3m6nb

9m 9i^36} *6 9130 36 1 (nb. 61b. 6 .5 ); qgm^mbnScn^'bnb qj36ji3n3hn 9^(^3336(1 (nb.
6ib . 6 .08); U3pnQ3636n 6ij)336nb 8mdhimbi8n 9 ! 0 ^ 3 3 6 0 ^ 3 6 :>bn8np;>^36nb
5^9iig5nc36n b 8 3 ^ 3 9 3 1 (61b. 6 .8 4 ). hibii3nh33Qni, IPCMDC bn8^ m 3 b3
Qiyhg5(n6ncn 9b33Qm6i ihnb 3ihoi3630n; b ^ h 3b xn bn^sunnb 33biqjib36QiQ
pqjhm 8 3^ 19 5^31*192,361 Qi3ibibnicn363^0 h3ijun3&nb 6ijjh36n.
409 -
3006660060 b S t f O b ^ 3 t?aD 36ffi0n 6^30
6*b. 6.84 O3 g* NOj 5^36001 (lyjClSX'M 9i

bb3bgibb3i yn96n6*uoooi) g*939*336ob 5*303-
6i (vnSnmb sbiOSi'bhg* 9uo6ihnb ibnSnQi^bnb
5*6*60036*^3. Co ime^mpn (5*^36001
9*3939*336x0 9u36*h3); R qj3b3n; ST p^hm;
L *bc*e fi*9my*on&D&Xn gmoiRQn (Okano et
al., 1984). 9m6*(}3036n b33&nb*cn3nb nb. Freer-
Smith (1984)
Q3(j*0]hn g*(bn:>63&;)&nbai3nb gi8ibib o icn 3630 oi jQmhroqjoQnb g i3 p x

<33mcnp36nb qj3hob 631530^ g^> Jbm3o036ob, 013 60130° 9l536ihnb
5 i b 3 m&*.ib^cnn 9 ^ 0 ^ 6 3 6 3 6 1 1 , hmjjmh^ 6 3 b n , 6 *h 8 m n j0 6 36 * n6 3 i 536 i h 33 &9 n,
hm6c36fH5 5i60i63&30o *hn*6 6*363 6*3hoinb 5*6m9ymqjnb 363^0^1 boib0m33b.
jhm6oi30n 0 ^ 0 *6 3 6 3 6 0 5*6nob*^36* 99h*0n 6m6ob ih m gô o b 936(5oh36ibi g i
qj3ho)o03hni6ob 93033^3660 (8*5., 8^3hn*636ob b^hnQ^hm&i). b3 8 15 36 ^ 336 0 0
5*6b*])3aih36ocn 0 ^ 5 3 6 3 6 ^ i i 96oi03hn bhgob 3hm(33bo; 03hj6ob b^h^j^hnb
(5300036360 (i5bh. 6 .14) g i 60n3ho h5Pi036ob *6*00^30 b i63i0 36ib 53*d033b ‘b^b-
ij*g g*3*g5o6mai 3hm6h3b30n n6obo3ho gi'briiSQb^&nb 9o9go6ih3m6i, g*6y36ob
ai*ho(p36o g* bi66hd0O3m6i (6ib. 6 .85). 63360b 63gm ai03i 3q3hm n930icnni,
Ijbhnco 6.14 bob 93hj*69o *^0pibq33hmb Q*9:>6n6d3h36036001 5*801633300 93b*d0Pi l33C°CO^D^n

(+ fti'bhg*; - 938^36*; x 15300036* 5*9mb*j,xn shob) (Lieseetal., 1975; Keller, 1980;
Halbwachs & Wimmer, 1987; Wimmer & Halbwachs, 1992)
9*bsboscn36x n 3n933ecn3b-efr>3*6cn* 93bj*6n gj*h30or^pm3*6cn* djhjsCn
i*96o*03hn 'bhei .
530*69x0 93tr)6ob qjmh9oh36i ±

bi03h36Jn93ho Jjmjooob 5hm5mh(jo* + .
35h3gnb i3gCob bobfo gs bo9i3ho33 - ±
6m^jmb bo{,hd] -
0)h*j3Og36o *6 vjhiJ3*03ho 3CC)9o5lj36n fboeiiohob <3*hoim6'b3) + +
(5h*i)ng36ob *6 OyMD^jbn XOÔ^^b bn^hd] -
,jh*j30g36nb *6 (jh*j3*C3ho 303936^36ob go*93^ho - -
,jh*j30g3690 9tnb*'b(T*3h3 ph]36ob Ii0(jb30 +
9mb*‘bc>3h3 ph33&ob gn*93^ho
bbo33&ob ho(jb3n (i33<Jiob qjihaim&'bj) + +
h3lbo6nb *hb3&nb hoybjo +
g*9*&hpi33&xn b36b^*6(jo* X
liXXn^ * UhmSmhuo* x X
0056060b !3hm3mtn5n* X
3jh3gob i3g0ob bb3* 039J96x 3&r' + X
- 410 -
^GuiAnanaae^tnO u & f n o u ^ o g M d n r ta 50
no
%
BA. 6 JB5 SOj-nb donjhn 6id3nb 33hJ6nb

Shmejjiyii'bj. gi'bni6;)&ob bihnbbo:
1 60633606 jph^nb
pi&jo&s; 2 6063360b dcnjhn y>h53i; 3 bob yjBBjhmba
p i ^m^nb 6n<(xn3&ob bdmbnb 9^6^36^; 4 bob j366]hfny
6nb b9m6i. b]]6nb bnj3pnpo (Vins, 1962). 3m6iUo33&n
fihp. Mtymyib brnhjif^ho ,jynb 8(536:^33606101306 nb.
Amundsonet al. (1990)
2,3663hm3&n 6y0oon u x ^ g 0bhb5g3&Ob6, lS^Ci)DXn lyîbD^n b0Q6> gb 6mpmb

9lS36iho
ndg36bg, hb9g36bgbu 5Pi5nhgnb mhs)b650 gb qjm^nimJbogbB^bo y330b'b3
&i 3MiXD&DCn 3^36 3 5b‘b36nb, 0bcn 0b5b0ocn'b3 j330mcn 5b6bo03go 0^636^ 653-
6bh3360n 5^0^633300 9^ 0 ^6 36 36 0 p i ou>30b(j3obb gb h3<bnb(j 36(j x |T|^n'J 93jb-
60^3360. 3b0n5366yb06bg36n *6 333 gng gb‘bob663&b 0633336 0^36bh3363o, 0b5-
hb0 3bOi bbb6obOi3&6 0^nh3 bo3h(5Xn 6^3^15X 3^ . qg(jmh6yb06bgo ftbBbbiôi-
hQboon d0o3ho 6b6b00b3Ob. 0u36bho3&0o &b533&ob 5^001 93(3*63306 338 9 35 <303-
mhogjbo 6b6n0Q36ob6 b3m30bb(i£)lin 5'bob 03033m6ocn. cnb3gb3oh330bg n6y3&b
qjmai03&ob Jgmhm'bo, bmpm 330 9 35 - qjmcngob 633h36obb gb $0933606 633b nYbo.
5n5ohgob nfopij&n
(jmjbo^h 5b'b36b Smhob y 3 X i '6o b3 ohbg 0U36bh336ob g h b n ^ b b b 06333b
S O 2 gb 3b 3hmu3bo 3o8gn6bh3m6b bhb 6m0m 4000 60nb 5b63b30m6b3 o, hmg3-
bby gbn6ym 5 ^ 0 (19 8 30 15 3x 0 0bg636nb gbpb653b, bhb03g 93gb3o6b'b3 boymybgob
Ribbb3ngb6, 3X i p 53ho bj^03m6ob hg5 o0363o. 0u36bh33&0b 33m03uoob 3hmu3b3 o
S^bdo^b 5ni5ohgob mhgb65ob 0mj03036obb00o 536mlj)033hbg 835336b.
9*363 b*‘6ob 83^6034 8ij36bh33o. 013 6b5336n 5bbb6o0ob, S 0 2 ob333 ê3n"
Qbg b(n633b 0U36bh30o, hm5mhu CO 2 (rsS02 = 0,84 rs* ° 2 ) [2 4 6 ], 0bb3h30o 64533-
&nb 938oib333b3 o S 0 2 (CO2-0b6 5b6bb3b336ooi), nb333 hr^nihu gb656bgo, bg30-
C^9 gbbb3b W i jn i x i h x 6o6bb(ji3 Q35m6bb. 5bh30m3o gb&40O im 6^36lihbô36ob
(gbbb0m36oon 45 |±g,h S0 2 .0"3) 5b9m 533hgooio ^ h sg jb o b ,33(15(^0 0uohg3&b,
hbij 06333b 6453360b 5bbb6bb. 3b3h3o S0 2-nb 0b(?b0o 330(5330fTi6b (1300
S O 2.0'3 03^0) 06333b 6b533&ob 0bb;jh3bb. 3b 0^130364 J>bh5bg bb6ob 6o6bbp9g3-
50163606 333^ 330 0 3m6bi53036ob ihb36m6bb, hmgsbby ^ 5 3 3 ( 1 bg5o0o bj3b |^hb5-
biohbyoob 0m0b^36bb, bm0rn ^ 5 3 3 1 1 30 54^014 15300b o6(j36bo3m&ob 838150113-
6bb [ 173],
S0 2-ob 015364(1380 03(^63306 330950100 5^4 390163334 C 0 2-ob 5'bbb (ob. 6bb.
2 . 14), bm0m bÔmbqyhribb gb j0mhm30bbli36b (bbtjhb6b3ohbi5ori 0b6dognb bb6m-
0m n 63h(j,o0o) 3 mhob shb3630o 3(^61536(1^4150x0 5hbg036(j0b bogog3 050334.
D^hQQnb 3393000 S 0 2 oib30b3oh330bg 6y400o obb636b, bri0m b3 hsbjôob 3hm-
- 411 -
aoaeaAoa&o a ^ a rm rta n & a a o
eDJ»SD^n ' HSO2" g* SO22- 33h3gnb 9056001 6*6009360*6 jprnhm^Cib^^b'bp, un^m-

bmpbs 3î;]mQ3&9n 83b*6*0nb*g 96 : 3 : 1 % oi*6:>qphgm6na> [ 179]. jcmhm-
Spib^&n, hm03QOib pH bo6*oi038o 8 *(j>633b, 5333006360*6 nm636nb 6*03*6559.
SmjdQeobnb *95O0*b *hb3&30O 3tirnu3b36n. 5fii5ohgob 6*31101360 (*5(130133
S 0 2 9* H2S) 9u36*hoboi3ob 3ubm *h *hob. q3o0m5363,bob *9(1330 b(i>*g036<b 3
3hmij*honi^36o 835336300 093636 03*33, 5m5ohgooi 9gng*h j,*h30mb. iQn^ma 5m-
5ohgo 03b*d03630n* fi*hoi30o oyro 6030103(136*01* U0nb ^PÔX q3rri(i03&0o. 3nh-
330 b*qj3b3hb 6*h0m*g536b bxojoôb 9*3*6 53* b x q )*6*9' hm030ou ^h m jbog*-
“b ^ n b 803(1 33h3gnb J>3903&8 n oi*3b936*, dohooi*0*g 30 8m6*6o03m6b 5m6n(i9nb
q3mljribn6oi3‘b3h 03j|)S&m0n<b08or hm00ob b*6m0mm 3 ( ^ 9 3 ^ 3 6 0 * 5Pi5ohg830i53xn
*8o6m0g*336n - 1506^3)1060 9* 830101116060 (6*b. 6. 86). 013 5Pi50h90b n6v|)36bn-
3I10 8315306360b 5*0m oiori0ob 3^16(536^11*150* 0*l530m6b, 0*8o6 ftm^nhgn cgnjbnh-
936* 503(jsoiom6ob b*booi. SH-353qj36nb 9* b30g3oôb ^*h6n h*mg36ni6ob 8 38-
oib333*3o obo6o 5*9*90*6 630030980 9* H2S-ob b*booi 2,*0moymq3o*6 8(536*hn-
9*6, h*u 06333b 6030103(136*01* 15300b 3hmi53bob 5*6^30(1013^.
^m^nhgnb mhg*65ob ^*h6n h*mg36m6ob5*6 5*6oi*3ob30036ob 93j* 6n'bSnb
3c233diiDhm6* 33^(^39300*. S02-nb 830i3ob3&ob 0m0*lj36nb 3hmi53b8 o 0g*3o*6m&*
0^(1936* 9* 3hm(i)m30*<b8ob 63033(1x11 I)6*hn *0(11063(136*, b x t3 niiD^)n*J h*m036m-
6* j0mhm30*bli)368o 0*^30m6b 9* S 0 2 60mj,nli36*b 3330136b C 0 2-nb RuBP-3*h-
6mjbo0*b*bcn*6 833hcri36ob *9500366. *9 ob 5*0m q3Pi(|)nibn6oi3'bob 8301(1*90 3hm-
]~ S° 2 ~ ]
Bab. 6.66 5^9 rnbibx 3&i n0 3hm-

U3bnbs, SOj Bognb jjhxSn, ^ 3 0 13 3
3nbn p3^mjbn>>iuni Jcmhm30iblj3b9n
{jm^nhgnb 33^*6m(Vi'b3nb bi93iC3&nat. APS,
i63 bQ36n'bn5<3nibq)(nb3c(3;>6n ATP-nbs
6* SO^" hjijiyinb 33935^9 3nc3&3(vi
3nh33Qn Shmgîyi. siyi3nh3&xn l-,X’° 3;>iin COOH COOH
(qjmbojm-APS) Ui^nhpjbs 6m&nhp939|J3X H .N -C -H H -N -C -H
Unc>b (CS), 93&36ie Un<vn3i6-{)mj)nhgm346n I 1
ifn95o3jbn (CS-S03) q}3h3emjbn63&nb H 2C - S H H2C - S - CH.
(Fdred»Fdox) 'bDQmjao&D&ncn 6ihexiJ3636s L-Cysteine L-Methionine

^X<53nese (CS-SH). *9 îCbqjrohaiur^nb
ghmb 93bidp36xn^ 0-i^>inp b3hn5nb
(AS), (jnb(Mn6nb, 930inm6n6nb gi
{,C3îcnnm6nb 6ih6mJ96i. ba&fnppimg, Bih-
9mnj963&i H2S. hv 05*11^6360 (Rennenberg,
1984; Gaised, 1985; Wellbum, 1985; Lendzian,
1987) Glutathione
- 412 -
660)ff,n3n a a 6'3(f.n b d t f o b ^ o
SO^Pollution
Lowering of Mechanisms against

toxic concentration toxic stress
» \
Reductionof Reductionof Sulfur Detoxification Highstability
SO.uptake acMfication metabolism ofen^mes
I I
Thallusstructure Buffercapacity Formationofcysteine C,-syndroine
andmethionine
Dehydration Bindingcapacity Antloxidatiue
of thalloplTytes (Lichensubstances) Accumulation systems
Slomatal closure of glutathione (Peroxidases,
mvascularplants superoxidedismutase)
fob. 6.87 S02-nbip9n h]'bnb1j;)61ix rnknb 3oii6n“b9n (Wirth & Turk, 1975; Tesche, 1989)
I53b3&n ncnh^S^bi, hijj ^ 3 0 1 3 3 nB333b 6ij>336nb C 0 2-nb bih35X >

I3nm 03ji6n'b936nb 33933016001, 5bhg:> :>8nb\ nhc>333b 36bn93bnb 93010^3300
hmQ3bb(3 bg3&i ^pogn^nb qjoiôioijbngiôi b3003^^9, 6 ih9mnJ3636i
b o g^^n b hbgnj)b036n, hoi803&oi5 (013 8boi b33i6f,-gnb331»hibb ghoi30ig ih 5:063-
n^hcD^b) 0633336 jpmhmqjnpnb g:>bo:>636:>b.
S0 2-noi gi^nhcrqnb ghmb ahba&xn h3brtb^36lj x fT)^0^ 0Oi*6nb836a bb 3>
Qibb3i 3bbn3hn gh ij^n3hn 3hm(53bob 03833016001 9|j36sh3b 5^h.!)3330 <3^503630
333d gn i 33iR3hmb S02-ncn 06,501jb o j^ o i (6bb. 6.87). 8^36ihnb boi5n3hoin 01^30-
b3&3h36>, hm03QiH5 b3Qi(j ih ihnb 9ob3boi6hn3 $:>33nh3n 3^36 3 6i3hoi3&nb 9mj-
0O936iboii6, b3gb 36yoi6b 3ib ^hdmpmb n3nbo3&ocn 5:180163330 gs^3nhoi3ib:
6iQ ib013^6(1 8u36ih336n, hm903bn(j 33hnmg3Qig 031101^3636 ib ip cgoioi036b,
ihns6 3qjhm 6ij)Q36ig bi3 n3 8g50i8ih30i6:>3n, 3ngh3 6(163(1136360, hm03Qcm> cgmav-
036 0 3hi3i060(ii6ni. b:>3oi:>hq3oia)oi00i5336o 9 536 ^ 336 11 8 38 0 ^ 5 0 18 :^ 3 03010103-
bnb Rb9mu336nb 833330161101 ooii3ob3qj03636 oi:>3b S 0 2-nb £>h6n hioig36oi6ob-
5 ^ 6 . oii0oicgnl!)36nb y330i b ^ h ^ D ^ x n , jn8o3hn 013 3^01001503110 oi:>3nb363h36:>,
(1018036015 ^ 6 3 0 3 6 3 6 S02-ob 8 3(^ 6 3 3 ^ 8536 ^38(1, b30biyhx<i. 3 i 6hi9 93801633-
3ncnn 6(13^6013(163636(1^.
^m-jbm^hn 30)3^3606 01^309^6 ibi(3n03&0ig shb3&30n 6,^360^0601306 633150-
q jn ^ h pm6nbdn3636b ^301 jo9n3hn g i 6noijn9n3hn 6i6o:>oio: Sbpign 63qj3h30n
36ih n ^363 (101636(16 3^ 0 ^ 36 30 11 83013(1636(16 93833016001; SO2-06 9o8i6h36b
6n30i03h36i0ii 153006 93mhig Shoig^j^Q^Q (00^363606 6iQhoi36o); 5Pi5nhgob
S^^bm Q nbi^ni gh 6o6gi6n6 Rslj ;,h o 6 x o oijbngiô}!™ h3iji5036n. 2)0933 3hoi, 080-
bnQ6b3 9^8013^093630, 8353360b 83ji6ob 8b 6ih3m ig536b - C4-bo6ghoi3n, hoi030-
ni3 S 0 2-ooi bsSîooig gi^ o h o ô b 3ohoi&363o 6ij)3:>oig 3<33il3ho:>: REP-ijih6m-
jb n Q ib i 50i50hgnb0ihg;)66nbbg3n 3q)hoi 6i^036bg 05hd6oi6nbh3b, 3ogh3 RuBP-iph-
6m jbO 0ibi g i CO2-06 9 ^ 0 13 3 6 0 6 83ji6ob 3o 6 33833016001 013001 RuBP-iih6ov
j b n 0 > b i (5 01^ 3063039936 ^ C 0 2-o 6 9 n b i3 h c n 36 9 ^9 . * 9 0 6 0 1 8 , € 4-81536^336 0 ih n i6
S 0 2-6:>g9 o 3ghoi 6i*)036ig 35hd6oi6o>li3, 3ogh3 €3-8(536^3360 (6ib. 6.88), hnb
5^00115 5oi5nhgnb 5i0oiyoignb ^95003601^6 ibjvnb S 0 2-big3 o 633i5nqjn]hn
3g5higoi6nb 8^0163 €3-01536^336006 (8^5., bb3igibb3i j,oi66n6366nb 3 x ^ 3 6 -
- 413 -
9036663060 9 ^5 0 9 6 6 9 0 6 6 3 0 366666 1160167)60
6ib . 6.88 6jjn-qjm^mbn6«n3'bnb

9:^33(101361 (% b^qjcm Sijh'b] Bihamjii]-
pn qjmiynbn6oi3‘bngi6) C3 gi C4
9u36>h3]69n SC^-nb bb3ipibb3a yn6o3&-
ljhi(}ni'bo Siom 8 bow ii6n 3jb3m(bnunnb
9d9G36 (Winner & Mooney, 1980) S02fumigation [jii r 1]
cn:>6 3mbihgn Poligonacaea, Mytraceae-rb 6ih9m9iQ5363pn M etrosideros collin a

3i3inb i36dxD2>t>3) 3(101^9 b9nhig 363936001 C4-0(536bh336b (M iscanthus sinen
sis, Andropogon virginicus).
bSjjjnqjniflhn 3&M6ri&nihm6i n0obn3606^960. 9jj36ih30u> b b 3i9 ib b 3i bbb^m-
dbb, jn 3b i 013 Qjjm^nib j>sifi6n:> 9*363 6 i 3hcn3&nb;>g3 n 5:^63*3363011 35 hd6 m-
6riihm&i. n3nbn3&nbi90n b^s^ngnîn 0j,hd6m&nihm6nbi g i igilBûnnb 36ihnb
I5mg6ib 9090 3hij(i)ni3cn 36 (1336300161 i ^ b gib ibo 3& x 013 n603blshn30 (1351101-
63660 3(536ih 336nb 9ihj)3nb>oi3nb (ubh. 6 . 15). ym33C 9 3601633313(1, 9*^(11636116
83116(1036* 83bid03630ni b*b3m636nb *9oihfi33nb, b*b3j3&nb 83^*6nb g i bb3*
C'm6nbdri3636nb 339330161101. i 9 5*30 36 1^ bh3Qd 5 16 3 11^ 10 3 6 1 8 3 ^ 0 3 6 * 30n-
bn36nb 36(183630013*6(1 83815111136(101.
ISbhnpi 6.15 b336nbs pi 63fr)36nb 33{,33&igm6i SOj-noi dcn3h gi&n6d3h3&3pn 3a6n,ji(j 36nb:>e3 n
(Krussmann, 1970; DaPler, 1991)
jshy> 3»:»tw* bsSxvn 335,3361

<3moi(vn3i6n bQQftn
Acerplatanoides Castanea sativa
Buxus sempervirens Ginkgo biloba
Celtis australis Magnolia h)poleuca
Gleditsia triacanthos Populus candicans
Platanus hybrida Robinia pseudoacacia
Quercus-eio Iwbjmftj&n
Sophorajaponica
6n63m3b6n b33&n
Juniperus-nb b*b3m&36n Chamaecyparis-rb bib3m&3&n
Picea omorika Picea pungens, P. glauca
Taxus baccata Pinus mugo, P. nigra, Thuju-h bib3rn636n
b-f\\jbr\
Cafluna vulgaris Berberis.nb bib3/^6360
erica cornea Forsythia intermedia
Gaultheria shallon Prunus laurocerasus
Ligusrum-rb b^m&a&n Rosa rugosa
Sambucus nigra Weigelia florida
- 414 -
Ubhnpn 6.16 SO2 Q* cnj36;)i>r>b <bhp;> b ^b i gs J33^'t>0 (Kershaw, 1985; Amdt et al., 1987 p> bbj. Shi3^ s^hn)
415
-
56m A n 3n a 36Dfto UftfF.a.'a'ô sad&nAOMt
9006663060 LftrtOliZR 9^ 3 0 9 6 6 0 0 6 ^ 3 0
S 0 2-big9 n ^5(130133 H F 95 H C 1) 5^66^^301(136001 05hd6ni6nih3 ihni6 b r ^ n -

3hoin bi3bn, 93^360 (ybh. 6 .16) g i 9y36ihnb 3icnm5363hn bm^m ( 3 i 5 „ Rhytism a
acerinum, D iplocarpon rosae, Gymrtosporangiun-rh bib3rob36o, Puccinia g ram in is).
5i6bij)3aih3&ncn 05hd6m6oih33boi 33C3b36hn 90^36360, 3q}hm 6i])C36ig ■ fohjnbi
g i b^Fijnb 9(^36360, 9 i5 h i9 ihb^brnftb i5h 30i33 S 0 2-nb i9 ^ i6n 9 0 ^ 6 36 0 ( 9 i 5 „
33Qj)i636nb 5ih9 ] 9 m 5i 3hu303&3cn 9 ^ 3 (13 ^ 1^ 3 9 0 16 ^ bib^mb^n). 30 ip o 3b 6133-
6ih33&9n gibni636ob 5i9ri963330 S 0 2-ob imi536^hi^nnb ghoin 03ib3gn CnJ3^Q'
60b 393(i3bm6i9n 06333b 6360^306 gih[p33336b> J9mhmajn0nb pi90ib g i bhpnb
93(bp 393ib.
i^Smbqfthxn m jb n e^^& n : ibnyyib rrjbneo&n, m(br>)6n, 93mhign 8 * 3 6 3 5 ^ 3 -

6n
ibm^nb mjbngQbn (ibm^nb gi65n N O g i ibm^nb mhgi650 N 0 2) i^9mbqj3-
hm9n 6ih9mnj9636i 9^(3^9^3333(1^311390 6311b 311^15363606 93935ig . 39 1^ 0 0 6 0 33 -
hn higniunnb (300-400 69 <3^(1593690) 9o ii6oij9 o6 93935i g N 0 2 n9g3&i i'bm-
^06 g i6 5ig g i ij^n^h gi6 56 igig, hm93gou Si^hnb gi65&igob 9mg3j)30ib a ii6 3 3 -
3hoi3bnbib 6ih9mj96ob mbm6b (O3). qjm^mbn6oi3'bnb ghmb Bih9mJ95n0 NO -b
mbm6n g i6 5i3b nb33 N 0 2-ig. i 9 bi9 mjbngi6^b Smhnb ihbg&nbb gn6i9n3hn
6m6ib6mhfn6i, b i6i 9 n6g3b^hnxf> 9^ bb 3i ij^fi3m6nb 9ggQ5ig i^m bcj^hm Sn ih
5i9mnymqji 6ib9oh6yi06ig36o. 0 3-0 ^6 3hou>g 93mhig n9nbn3h 6i3h o i36ig 5 3 3 3 -
0n6g6i ^>5(130133 33hmjbng30n (1:^0^93611, hm9Q3&ob5i6ii5 939g5m9o h3ij(jn3bnb
9 3g 35 ig 6ih9moj9636i i^ôgSn^hi^n, 3hm3nm6ri06n^hi^n, 63^0960^(1^0 g i 636-
‘bn06n^hi^n. qjmli)mnv|bogi6l536nb (9icnn 9bob bo6ioi03b3 gi9m2)Og363Q36ob 5 i-
9 m) im5j336^hiijn30n gn6i 9o*)i gi9ming3630Oi 630n6ignbi g i gpob ghmb3,
i9o6gb3 (6ib. 6 .8 9 ). 9cni9n m'bm6nb bi93i0m j)m6u36^hi(3ni 9i li 30m6b b(P3nb
gm6ngi6 bn9i(ji0nb 9i^36iboii6 ghoiig, h ig 5 i6 gi60m63n 5i9mymgjn0n 5^ ^ 36 0
lb33ncn 90063336 g i nj 5i6n(jgoi6 30^hioobcr)3ho bSgjjjhncn 9gngiho 9bob
higniunnb 6i3036b.
8*363 5 i ‘b36nb 93p 6 Q3i 9u36iho9n. y330i 3b 5ibn, 5ibb6n0o bi5D3&ob
5boai, qjmaimcfln 93gnb obg33 j>ih5ig, hr^rnhu S 0 2. N 0 2 i 3iini 30i 9 n S 0 2-b 3
&33hig bBhiqgig i(p633b (g i i 9n^m9 bb6ihnb ^n3ob 933mbo^36nb 939 33^ 16 0 0 1^
j)0 iQ3fi0ig 930cn30b36i), 03-nb 393^360 6i6ogo g3og3h9nbnb 5ih 3a ii 9h39o 5ih -
ginj963bi g i6 56 ig ig . hmg3bii5 ibmôb mjbog3&n 33h3gob ^3993680 6yigcni6
jjm6^j(j|9n 93gni6, 6ih9moj9636i HNOz g i HNO3, 110199360(5 6n^hnôbi g i
6o^hi(inb nm636ig gnbmôiijnnb 939935 ij^n3hip 93noi3nb36ni6 3hmli n 30i'b9 i 3 n.
9nj93g3&ob ig5ngib ihbg&xn 3hm|j3b36n. 6o^ho^h3g3j^>,bob 9 rn6i 6 o03n 6 noi
6nj)hn^nb 0^6360 ibn9oQohg36ni6 i 9o6m9g i3369n. i 9 939oib333i 9n|j j0 mhri30ib-
Ji3bb gngo 96o9363pm6i 36n^3&ioi 9m96i 9c i 3n 3ggjiinb 93h6og36i9o. i^ 9 mb-
gjghxn ibm^nb mjbng36o cni3nboii3ig 9y 36ihnbioi3nb 6ih9mig53636 ibm^nb
gi9i^36nai 6yihmb. 635i^03h 3 0 3 3 ^ 06333b 9g i33 ihnb 9 gj96i g i S 0 2-aii6
Jjm96n6ii30i9n 3m^36^n3ho ^mjbo^hmbob bi9n9hm36i. h ig5i6 S 0 2 06333b
6ov!)hn^h3P3j^)ibob gi'boi63bib, ibmôb mjboggbnb gi9 gnb 3hmi53bn o‘b(j>3g36i.
pVbm6n 9(336^33690 bBhitjig n9Q36i gi65& ib ig g i 6ih9mnJ963bi 3ghmjbng36o.
9mj93g3bob ig50Q36ig, 3301133936 ym30obi, 63339o63bi 30i‘b 9 i0 39 i g i bmgmb
6om939bhi63bo, hi^ 06333b 9396hi6nb 93(p633igm6nb gihp333ib g i 63*)hm'b3-
6ob 5i63naiih3bib. 63360b, 636^3606 gi 6igib36ob m'bm6ncn (n9
•jfn6i}36^hiiin369o, h(^993605 9^ 36^ 130 16001 o'bm936i 3i 3h9o - 0 ,05-0,1 ii.9-1)
gi^3ohoi3n6 9 3g 35 ig 0|j36ih3369n 039R6339Oi 93j)i6m0n‘b9nbi g i bhgob 3hrnj3-
bob Qih("333360. 33hmJ6nii53li)096n^hi^n gi65>3b 0hi3i0, 5i6bij>3Cnh36oa) SH
353qj3bob 9395339, 6i3honb, go3og36b, o6 gm03bo5ob ^ 3 3 6 6 , hm9936ogi6i(j
- 416 -
aemfnm n&aeafnO %>d6n«>9&o
O r .-- ■---1--- --- --- ---1—r-

0 50 100
Ozone concentration [ppbj
Bab. 6.09 ipljb ob aaj'by ^^dnibqQhmb J33pi ShjjbSn mVnSnb ynBujS^hi^nnb j>hign]6^n. Isjgs
b^Hbain: 0 3-nb 6(yi]hn (h^mcD^n. 01310360) g i oqn;)hn ba33aoo>36n (bVfco&n) b b ^ i b b j i
bnSsTCD’b]; 63h^ncm3a6n biVi: gn6i9nj)i 9-16 biitfi^&nb &a6di3c<n6idn bi335>ip(5nm 5;jhnmg0n.
J33ea b^hsom: 1987 6(yib i&3<*>am3n G(TnDhn U30nc3* 3 &n (Smidt et al., 1990)
6ih0mnJ9636i ‘bhpnb 3;}0'b(p3Q3DQn 3nghrnjbn93cnn0rr)bn6gf'icn.

b^xnagnuhn Sbhd&n&tohmb* n0nbn3&nl»g0n. bb3ig>bb3! 9(536:>hnb
qjfn^nmjbnei6^36nbi90n 9j>hd5fn6 nbhfn6nb 8^6^1539360 (ybh. 6 .17), hpri9p3&ni5
3yhg6m6b jpbnoi g ^ n h o g n b 8393306001 6 ^ ^ 13 6 3 0 3jb33hn036(j3&b. 63636^90
3b 55^ 36 0 9mJ93g3636 ■>m9&n6nh3& x :>P> ^ 6 ^ 3 9mb:>cfTig63cn:>, hm9 3icnn
9mj93g 36b b h x ^ g ^ 390*53335 3Jb33hn936^ ,x 9fri6:>u393&b. b6mh3g :>9nlyn9 ,
gmongnb jn9n3hn îpn'bnb s6 gs<bro63&3i>nb 6 ^ i3 3 3 C n b3h>cnnb 83933(116001
hin93 i f '6 ih36X n mjbngaJi^nb 553C36nb ng36j)nqjni>ii5n:i d:>on:>6 d63pni.
- 417 -
3 0 0 6 6 6 0 3 6 0 U S6 0 U71C ; 9 ^ 8 0 9 6 6 9 0 6 6 3 0
Ubhnpi 6.17 ^3hôm3i6 81536^3011 cgihpm&nann b36bn&)hm&> I'bi'Hyib mjbngj&nb, m‘bm6nb gi

PAN-nb 9n6ihan (+ b36br\j3hn; - 6ij£36sg b36bf\j3hn; X bidjipfne b^Sbiyjjhn) (Onnrod, 1978;
Guderian et al., 1985; Dapier, 1991; Treshow& Anderson, 1991)
3t536sh30to ityyp ! m^bn / ^,3^360 NOx; 03 PAN
WCiiO^xn fooijbjbn; 6mb(j633Qn

3ihi53pm3i6aii 393>J3bm6i + +
dhnfjjn -
b*33hi93 6iCit>3&f> + +
3ihimb636n + +/ X +
Brassicaceae - X -
Apiaceae + -
Beta -/x +
Spinati + +
Solanaceae X + +
Asteraceae -
Cichoriaceae + /x +
e3imhi^n-)Cn 63^^360
Comus - -
Cotoneaster +
Gardenia X
Hibiscus +
Ilex -
Ligustrum X +
Pyracantha -
Rhododendron + + /x x/-
Syringa + x/-
Viburnum -
ojmoi(vn3b6n 63360
Acer X - -
Betula + - -
Carpinus -
Fagus - -
Fraxinus + /- -
Populus + /-
Quercus - + /- -
Robinia - -
Tilia X -
6n63m363&n
Abies X x/- -
Juniperus - -
Larix + + /X -
Picea -
Pinus - + /- -
Pseudotsuga - -
Sequoia -
Taxus - -
Thuja - •
- 418 -
66(J)rtn3n&367)ri0 bGri3lw)c?0
6 .3 .2 .2 . a d n a o a o iw e 3 6 n - 6 n » » n w q s eycn b 0^353
6D3h013&n
8h*3*0 6i3honb 8mhob, (1^803605 3330*333001 013 3y3hagc>36m6ob 6*3m

639360* 6obgbj,3 n, 6yo36bb g * <be*33&0n, 3do03 6 3 ^ 3 6 0 ysx^ D b*3o8ob gb
bi65hd0f»3 3hm&03336b 3j36ob6 3u36bh336b, hbg&b6 nbn6n ^^60^83680
&hpi3g3&o*6, 8ri6b6o03m636 6om0mg,o3ho 6*3hoi36ob 6h36h363b3 n g i b3QO036l|)3-
6nb bbbnoibtf y i gogbb6b 333d0Obai 3ynboblj30b9o gbhR36b b*0 o9 o ^ 6 5 3 6 ^ ^ -
& 1-2
2
S 1-01
® ATPase
E 0.8
N
0 0.6
a
1is 0.4H
® Glucoee-6-phosphate
CE 0 .2 dehydrogenase
Isocitraie dehydrogenase
0
0.1 1 10 100 1000 10000
Zinc concentration (jiM) Concentration of metal Ions (}iM)
Bib. 6.90 SdndQ 3>jiC36noi &i3ro633X bôbb;) Silene cucubalus-rfo qjfncnpodnb ]6bn3]&nb hjbJ-
Uru. 3ih(ib36i b^Kionn: hjiJjni (jn6yib ^ne^^B^h^jnnb 81^36*^3; 81)133361 b;jh:>cnn: 60^1^^3935-
iîtib ijlyi3mbnb 153000361 bb3igsbb3i SdnSj Sj^cncn lb33nij8393&nb 333934, (Ernst, 1967; Ernst
& Joose-van Damme, 1983)
15036001. 8*g6ob 6 m03m33&3Q 8*p*hm36aii6 gb b b ji bb3oim-j>3mcnij>n3h 6*h-

6m3636oto6 *h b 36 xn 6o*g*&o 330(5*3b 3dn3o 03li>bO36ob (&*6b*j(30ih36ooi Zn,
Pb, Ni, Ci, Cr, Cu) nb30i hbm936m6bb, hm030fnj 8u36*h30i* 303^3bm6oboi3ob 8m8-
68*0b3Ob. 3dn33 83*5*036001 gb6o6d3h36bb *95000 bfob J>i6bi2)30ih3600a 0693b-
ljhn30 ^0163680, n6(j36bn3hn bb&'b*™ 3mdhbn6nb g i 6*fc3ob gi&hri336nb *g&o-
03680. 3 3 lji0ob gi3i333b33630n b*6*h8m36ob 2,*8m6*6m0j3368o 830^036* y3x *
9do03 33^>0nb 3m36i, b*6*h8mcn* R*6*h3ub368n 6*6^21301(136001 633hnb Cd, Zn,
F e, Pb, Cu, Cr, Hg, 5^8603690 3m6ym6n0m636nb 80*83o - Cd, Zn, Fe, Pb, Cu, Cr,
H g, &(b36nb 3ohbb gb *3(11*1^03630 - Pb, bmpm 6yc3&3 n y330*<33hn ob, h*u
6b03j36ob 013 bb3* b*33*03636ob 83833^6001 6*390636* 609*580 b^mbqfthm-
Q b6.
3dn0 a dapoo&nb 3x 633* 81536^380 9* ,yijxi£]h n 6*3036*. 83(j*0mog3ho

303336l|)36ob 330i3ob36nbboi3ob 3jh 3936b (&b6b*i30ih36oon qob3ob 33(13936^
5*bfi6oboi 83b*6*8nbo bb^hbBbSmh^m gb gb8bj,hri3363cn 83^*60^8360, h*ga*6
&33h 3dn33 83(j*C|b. nb333 hm^mhu 8 0 3(^ 30 38 36 ^ 36 ^ 0U36*hoboi3ob *J3b bbbniy
m|jb0m 36033630^16*. *8 2,^36001 81536^380 bp63336 *6(130133 8m86*80*3O XD"
- 419 -
(nb. 6.91 9dn9j 93(iiC3&nbig>0n hj'bnb-
d36(530™6nb 3obidpm 3oJs6nb9;)&n. 1
35h3pnb iOG3c3n QsdPC0^ nm6]6nl) ndro-
bnpn’biijni, 3nh330 ym3Qnbi 33^5063606
03933ndncn; 2 j>bîh36nbbg3n
3jh3gob 3j96hi6nb Bneiifnag^m&i; 3
(jn^m3Qi'b3i3n 33^>onb giS^jhn
yncjdnbi 91 3m0n333^ne36nb (cgrvjmbj-
0^063611) 6njh bxÔOinb 6ih3mj06i; 4
31^X10363° 3>ja036nb 0i30i; 5 sj^rvjhn
3jb3mh^n. CW 3jh 3gnb i3e3Cn; CYT
(jn^m30i'b 3i; VAC 3i i 3m0n (Emst. 1976;
Torasett & Thurman, 1988; G rill, 1989;
Cumming & Taylor, 1990)
836(5360. 3 m36i 3c i 3n 8(1^839365 dnhncnbgiQ 3g5m3bh3m6b bbbnym^bQrn 8608363-

mpbnb 3jm63 36'bo336o6 9501(153635811 (65b. 6 .90 ) 95 6360^3065 95 cgm^mbn6-
cn3‘bnb 3hm(j3b3n 303^(5(1(1163606 ^hiBb^mh^nb 9ih[f333363n, (15(5 06333b 363(153-
iiniDCn 6(55(53606 838(50(13656 (nb. 656. 6 .5 ), 6533360 6030103(1363606 83013063-
606 3 3 ^ 3 9 3 ^ 95 'bhgnb 3(101(53606 933(136056.
3dn03 83(35036065980 h3<bo6l536(330ro&5. 8(3365(133606 383,336(1165 h5(p5(5 3 o6 -
nSiQ^hn 3016(536(3(15(503606 '633001 3dn33 33(3503606 3 n3ihoi dpn3h 85(136(11605(13
69365. 01336511336b, (10180360(5 nlbh936n56 956(^3(13630 3560(55,53680, 833d0O5cn
3do03 33i|)5036o6 ^5(160 3016(536(3(15(503606 55630,5(150365 bb3595bb35 pm6o6do3-
6n6 03833(116001 (656. 6 .9 1 ): 33(13906 33903680 9 5 3 5 3 3 6 5 (136(1. 6 .18), hn6 5 5 0 mi3
3h m ^ m 30 5 6 ^ 36 o i56 3m 6 l55j,50 9 5 8 5 0 10 3 3 3 9 5 1* 18 0 (5h5663mh(50 5 3 m 3 0 5 6 (5 3 6 o 6 5 ^ x 16
5530001 558(110(10156365; 305<b8 n6 6565b(p3h(ii 93363606 33^63350(11606 838130(1365;
I3o^m305'b858n 5m5oh9833i3330 3oi0o333(5o03&oi56 (503,55010(1160 95 503^58061306-
(530606 93(1035^360) 95 SH-06 35303001 89095(1 1309360156 J305(in6 65311013606
83^865; 03^503606 (5(1^6033(101606556 958133300 6,5(136x0 3(101,53063606 606013'bo;
05 6010016 5 5 9 5 6 5 6 0 0 3 6 5 3533(1103630, (15(3 8110(^6335 5(15(11(155630 9 5 (11(155630
8953360156, 0336(11006 93(1035(1)360156 95 5003(11^9360156 5oi3303j63ho 653(1013606
65h3mj36o6 83833(116001.
8(5365(13366 3do83 33^50360156 3016,35^(30 59580560b 65(13306 55(1383(3 3jm 6 -
9501, 855., 8596360b 6050550b tb3053oh'b3 553mb30ob 595093680, 63h336ô636'b3
05 d0O3h 83535 6o5055'b3, 6505(5 5938060b 3(116(536(3(15(50360 (3(11^6033(105. 56301
595003680 55630015(19636 ^33013301,503360 05 63315003033I10 6563(116360 (855.,
"359606 8(5365(13360" 56 6635956635 3m6(!)0636(!)06 63(1336(1)06300 030(11(106 9585-
656050136300 6563(116360; 06. 156I1. 6 .22 ). 8do33 83(1)5036065930 h3<6o6(536(530m-
606 365(10 5363(3033(159 950^60(136x0 95 5953(35(503(159 00190030(50(1365905. 3do-
83 83(55936001 89095(1 6o505j,‘b3 Smbshgo ^38(1133(11(303360 558(110838533636 o'bm-
36‘bo036o6 633(5003033(1 653(1366 95 (530056 3hoi(5oi305'68o6 63315003033(1 03(p-
63359(11656 6b3505bb35 303036,5065930; hb(5 303(101 90905 30163(13(5x0 303836(5001
558(11633x0 9^(530(10135, 3 ooi 3q}hm obh^& s 3(5365ho6 l5m03li56l5m65 53 303836-
,3065980. 5(16360166 6563(116360, (1(11803685(5 85010 35(j'50O 5363(5n33ho 3056(50-
33(1111606 558m, 558(1108335336 65303659 633(5o50o'boh3630o 33(11(5033606 801300
h050 - 06060 h3<bo6(536^3036o 5(1056 8(153500 3do33 83,550065 95 6635 XD^D^-
- 420 -
Ubhnpo 6.18 (jffijnoi Spngih Sm'bshp 93^(jQmqjnlj369n ^nByib îSs&io^
bs pi Dhjgnb im95m636^]60n
bsbyn&s, wh5»s5m 3W ijjhognb (%> D3tv»nb
tjOtjmSos'Ws (%) C%) 43930°
Cardaminopsis halleri
ojmoincn 82 6 12
38 5 57
3np363 3x6^ nb
qjmoxnpn 64 10 26
qab3n 18 10 72
Agrostis tenuis
qjfnompi 48 11 41
93b3n 38 10 52
Minuartia vema
qjmomen 46 8 46
qj3b3n eo 8 72
^nb 3n0i 6cn; ib3 3 ij,., Agrostis tenius, A. canina, Festuca ovina, Plantago lanceola-
ta Qi Silene vulgaris 3g5hig36n ihni6 Zn, Cu, Cd, Ni g i 6i6n0oi6hn3 ^30133,
As, Al, Fe Qi NaCl 3 n3ihoi. 6536^301} 3dn33 33î036nbig3n h3'bnb^36^30m6nb
Umg6ib 36n^3di gngn 36n93630Oi6i gi6n6d3h3630n 595,003606 ( 3 ig 6 n6 gi033i-
33&nb (13500163611, 6i53n6 g33mcbn^36n) j>i363i636nbi g i (13,1)30^3^006 bij030n
g i ^5(130133 Sgbi&iQnbn 3i 363 303336^ 36nb n6gnii^mhn 013365(1336116 3mdn3bi 3 n.
6.3.2.3. nanunohn e^ n h o ran b 6nnn6enwi5n^
6536^360 3qjhm d0O3hig g i bi6j)hd0n3ig ihni6 3jb 3m6 nh3b x n ig 6 nCfTV

6(1030 g i 3 i 6 n6d3h3&30n 6i3ha>3&nbig3n, 3ngh3 13601330360 g i i g i 3 ni 6n. i 3 n^m3
nbn6n 5i3oig536ni6 3hi3i03bhn3 6nmn6gnii^mh36ig. 563013659 33od03&i 3 n-
306600101 mh5i6n'b3n i6 mhj)i6n<b336nb o ^ ib ib o ifc ig o ^ i, (1018390(3 îh6 o3 o-
6036^3 3ib3bm6b 6560(301136901 0336;)(503606 93l330nb {hjsjynnb n6 gnjpuns) i6
9i363 6i3honnb 955(10133606 (ijjS jp iy n jh n n6 gn^>u/u) 33833016001.
bibnj)6i0oi oih5i6n'b336n 55(139016 b3i0D6igoi6i'b3 (^ 5 0 (13 6 3 6 bibnuonjbgoi
001016013600363606 33b i6i3n big - nbn6n ^ 6 3 6 1^ 6 g i 0(313336056 i6 06^36603-
h ig 3hi30g36ni6 . qjgmhnb^gn 933i g j)360oi6 nb U3i036igoi6i g i 6 ih i3 3 X n 6ib-
30163606 j,ijhoi6i 3n3cnnai36b bi65hd0O3ig 5(163639 g i ^ o h o ^ i ^ . i 3nt> (56016090
9ij>i0noini 0*1^363606 553(1(5390^06 9 ^ 3 ^ 3 6 5 (6i6. 6 .92) - 3icnn lbhgn6 n6&)6-
60301606 g i cggoihobô 9 335953600160b 3nb3g3ncn 03nd03&i gi^3nhcn3nb
bihnbbnb (boi6nh36n6 gig&36i. i 3 ghoi6, S 0 2-6ig 3n bû n q y^h 3&hd6oi6nihoi-
6ibcni6 3(10159 5i6ioi3i0n66n63630ni ioi6j)h3j)30 5g5n06i3yoiC9303n ^ ^ 0 1 6 3 6 3 -
0n i0n3i^3hn 30(1016360 (55650136300165, 3i3hn6 ^360560165) g i 6366^(15^06 35(15-
6 3 ^ 13 6 0 (pH g i 1)330)360006 i6 3 65053606 33^(10103006 bihn6bn).
6 ^36 ^0 1 oih5i6n‘b336n 558010(1633056 ^016^3^300 3 i3 6 3 6i3hoi36o6ig3o
5566^30 1(136390 3j,hd6m6nihm6non. iSnôid n6n6n 33od03&i 5i3oi3oy36oioi 6nm-
- 421 -
8 0 0 6 6 6 0 0 6 0 b 6 6 0 b T )^ 8 ^ 8 0 9 6 6 0 0 5 6 3 0
<bm6p3&*p, hro9036n(5 (b^Bgih^nVihQ & x 3nhm63&9n *(^36300 6omjo9n3hn, cgn-

'bnmcmjjn^hn g* Qmhqjmcm^n^hn ihn^^hn^QQftnb 8063931101) 363363636
00 6*3haiob ihbo6m6ib gi, ^ 5 0 3 ( 1 0 1 0300^333*00, h*mg36m6*biu j>o. b i^ b ^ m
mh5i6n'b836ig (9nj)hfTimh5i6o1b836aii6 g i ubm33036aii6 3hou>g) b9 nh*p oy363&36
6yip9u36ih336b (ubh. 6 .19), b("0m6iU3636bI jbm3n03h pi 3 5 h3930 i30v5I)h36b.
biBmBn^rnhnBam mh5i6o'b836nb 83033m6noi, 8ion bijungjnjjjh ^ 3 3 ^ 3 giyh g
6m6ncn, 33nd036* giS^nhcru^n 6*3hoi36nb himg36m&h030 g i bihnbbm6hn3n
8 *6 336 36 0 36 ^ 9*9536*. i 9 938 0 ^ 333*8 0 bg3&* 63636(103 3ohm6369o 8m’b ih g
8l536ih3‘b3 g*i3oh336* ( 1 >bnjhn dm6n^mhn6i,n) i6 5 * 8 ^ * 3 3 0 3 3 h3j)0m69n 8 3 0 ^ -
636* b(ji6 gih v5n‘boh3630n 9u36*h330n 8 ib i (ij^njhn d n 6n^mhn6^n). o9nbn3bnb
9m6nlimhoBj)obiOi3nb ny363?>36 hmfcmhu j)ho3l!)m5*036b, ^ 3 3 3 q3*63hm5*036b
(ubh. 6.20). i>i393C;>L5nDho 06903*^01(136^ hm9036nu p*bn*636ob £*(1303 *5(10133-
636 8*363 6i3haiob gog h*mp36m6*b, 5*9m g536n*6 hm^mhy Sibnphn, nb3 ij^n^hn
9m6o(j)mho6g)obicn3nb: 3b 91536*113360 dohnoiigig *5(111133636 (101830083 ^ 6 ^ ( 1 3 -
iHC OCO^D^b g i i 9o(jm9 jo9o]ho *6*0o‘bnb b*03*036oai, 03od03&* *p5O0b*9y-
ncg30nb piôhcrôb bihobbob p*p536*. 9*p*gn i6bmh6^n3po ]6ihob 5*3m
b*3b36o oc3i3ob3636 gogn hi>mp36m&ncn om636b (ubh. 6 .2 1), Cnj363&nb 3jb 30 *6-
15*^3680 (9*5-, Hypogymnia physodes 3 ^ 0 ^ 6 ^ 3 6 0 ) 8 *36 3 5*^36001 (5*6b *-
330ih36nai SO2) g*ii3nhai3nb*b jgmhmcrjncn 08036* gi, 5*hg* *9 nbi, nbo6o i5hm-
33636 9o63hi03h 303936^36b (Pb, Zn, Fe, M n - j*c*jnb 03*11503690). bmjjri36n 9*
6*^3(10360 *5(10133636 gogn h*mg36m6oai 9do93 8 3^*0 36^ *9 n^m9 6*j^3hrt3&n
5i8m oy3636i 5*9683690 9m6ym6o0ni6nb 00i9obj>*6 936*0 nb 03b*oi3ob3b0ig
("9nj)hm630o *9mj*R3*“).
9h*3*0 bm in9 o (8*5., Amanita m uscaria, Xerocomus badius, L ac tariu s
vellereus), 00^363680 (9*5., Cetraria islandica, Cladonia-b bibQm636n) g* b*3-
b3&9n (8ij)., Bryum-ob, Truidium-ob g i Decranum- ob bib3m63&n) 1986 6. 63(1601-
6o0ob i^mg^ho X D Jii^ b ig j^ h o b i3ihnob 8 39935 h igo m ij^n x^b * 933301(1*9
5*o'b*hp* [56, 9 0 ], h*p5*6 8*cn 83ncn3ob3b 137Cs, 134Cs, ^°Sr p* ,05Ru. *9
*3*hnnb 939p 35 h*gnm*Ji!)O30m6ob qjmBob g* 6o*g*j,ng*6 h*gnm6;3303ng3bnb
930i3nb36nb 0 6 9 0 3*^ (136 *9 £3(1^0^13*60 9u36*h336nu 5*9 mg536mg636. 5*g*j3*-
6nb qjyj^mhnb (TF) 938331^6001 03ndp36* 9 ^ 3 6 ^ 3 8 0 *h b 36 xn hipnm633p3ognb
^31300303(10 ij(jn3m6ob ( A g ™ ^ ^ g*o»*6n0n 63gg *6 99h*0 6m6*‘b3) 83g*h 36*
60*0*580 *hb363o ( A ^ . , ^ ) i j i j n 3 m 6 i b c n i 6 [8 4 ],
TF = ^DSihnb (Bq 36"1 ) I A g n ^ n b (Bq 35-1 ) (6.6)
hm^mhu 6*6b, 633b 83^3^1(1010^1503(1, 3p*<33h g i bb3* 5*h39m qg*j^mhcni6

3(101*9, “bhgnb q3mh9*b gi, 5*6^3301(136001, qj3b30i* bob(j39ob (jo3b, 5*63001*-
(1360b qji’bib, j30m,iijbm6m0o3h g cgo<borili)o33h 6n8i6-oi3nb3636b (Ribes nigrum
ôjhm 93(jig gi6n 6d3h 36xn i, 3ngh3 Ribes rubrum) gngo 96083630016* *j3b uD'b-
n]9nb* g* b(jhm6ij039nb obmijm336ob g*{,hm336nb b*hnbbnb*oi3nb.
* i 3 9 x i bnnb n6goi*li)rih36nb 5*6b*j)3aih3630 35303b 6*h9m*g536b 6*h])33x n
b*b3(Ti636n, hm9036ou 9do93 93,j*036nb n6pn3*vjrnh36*g ( dj^Qmqjnrfj&n; ubh.
6.22) ihni6 ubnbngo. 93^0111030^360 noi3nb3636 gogn himg36m6ooi 9do93 93^-
i03&nb nm636b g i 9icnn 3fn6uo6ljhiun* *p633b 0,5-8 5(1 99h*0 6m6*‘b 3
(0830*cn*g 25 5(1 ifc^-SgD), h*u lOO-lOOO-jQh 93^0* 8o])hm30393Bli36nb 6mh9*Q3h
im6u36l|)h*un*'b3 g* 9]3b*6*036* qjmbgjmhnb* g* 5ni5nhgnb h*mg36m636b.
^hmSn^x 9bD^*3 9u36*h3b Eichhornia crassipes 9*pi0n *i393CiL5nDhn 36*hnb
5*8m oy363636 6y*0b*u*33&ob 5*b*b3qjoi*3360*g.
- 422 -
-b60)ff>[T)3r)&067)f?>0 b&f?>0U7)c ?0 flad&DrtO&O
6sb. 6.92 cnJo6mn6eni)i(jni - 0^363606 {^3h^3Q36nb h3ji, hn3x<XS sbib;>3b n5b6h]j3n gs 9nb
îha^Qm e>6n6d]h36nb bhpnb pn6s9nipb. ibjcnn h3»,36001 djbidfjix^ bifj^hdcrijn 3m5n^mhn6j,nb
Piviph3i>5 (1947-1977-1987 65). I <bm6i: Qnj363&nb 6<nh3sQ3hn gsq}sh3Qro6s 6 $ bib]0>6hn3n
9hi3iO<3Dhrn36D&>: II 'bm6i: eôjihjvniii ib]33 6mh3nb qjsh5p363n:>, 3;>5hs9 bibjm-
6hn3fi 9hi3boqj3hnn363bi. h>(j 9030110)366 Q^Wjhj&i'bo; III t>ci6s: pmdnBnh^Si^ 6j>jhm-
aj3Cn- ^Snqjn^jhn 00^36360 pi Xantoria parietina, (^363606 psojsh3e<'n&s SOiyCiiCD6^ 6mh3nb
CTjihj,p363ni; IV (vifo636nb p>ihn6n bsb3m6hn3n 339^5360016* ps 9(jnh3 pso)sh3(yn&}; V
“times: 330608636^ pnj3636nb 9bmcmp d:>cns6 9(jnh3 himp35m6s ("cyvto536nb 3pb66m') (Beschel,
1958; Bortenschlager & Schwaizer, 1988)
- 423 -
90066(^0060 bSrtouTi^ a c ja n a M o n w a o
Ebb. 6 .96-nb j,i{,hd303&:>
Ijbhncn 6.19 Byio9n Ridnhxn aijjhnqjn^jftn hmjmh(j lj3b^-mhj>i6n'b03&n i^Sm b^hm b

Qi9i6n5d3h36^36nbcisnb (Nobel et al., 1983)
gs9ii>n6d3h36ocf) <53^-81536^3 jm6tj3615hii5ni 50% CCWXXlhm&i
(ppm) (ppm)
gj36mcn Potamogeton lucens 0.2 0.6
Potamogeton coloratus 0.6
Potamogeton crispus 0.6
cvyi^'bm^n Potamogeton lucens 0.2 0.7
Potamogeton coloratus 0.6 1.1
Potamogeton crispus 1.1
k h 2p o 4 Potamogeton alpirtus 0.2 2.0
Elodea canadensis 0.5 > 5.0
NH4C 1 Potamogeton coloratus < 5 15
Potamogeton crispus < 5 15
Ranunculusfluitam 25
H 3B 03 Elodea canadensis < 1.0 10
Myriophyllumaltemifotium < 2.0 5.0
Ranunculus penicillatus < 1.0 10
oy3n» Potamogeton crispus 2.1
Elodea canadensis 10.4
Potamogeton lucens 10.4
Elodea canadensis 0.01 0.6
Potamogeton crispus 0.01 0.6
Potamogeton lucens 0.01 0.6
b3n036dn Potamogeton crispus < 0.03 0.06
Elodea canadensis 0.006 0.3
Potamogeton lucens 0.06 > 0.3
l}n6in Elodea canadensis < 0.7 3.3
Potamogeton lucens 0.7 6.5
Potamogeton crispus 4.9 6.5
- 424 -
a6(J)ff>ror)&363(nO Stod&C)rn060
Dbhncn 6SO bd jo yn rb £)hdcm3i6n 6(536^33611 hmjmhu ^dmbg^hmU pi6n6cl3h36ftb b36bn£)hn

n6gn])ivyiih36n (Steubing, 1976; Ernst & Joose-van Damme, 1983; Arndt et al., 1987; Rabe, 1990;
Schulze & Stix, 1990; Schubert, 1991)
ejQs&n&ljbjbac0 QgaSihnb bibgm&i

S02 Pupulus tremula, Medicago saliva
H-jS Pseudotsuga menziesii, Spinacia oleracea
hf, f2 Prurms armeniaca, Gladiolus communis
HC1 Syringa vulgaris, Fragaria vesca
nh3 Taxus baccata, Brassica oleracea
NOx Apiumgraveolens, Petunia x hybrida
03 Nicotiana tabacum, Phaseolus vulgaris
PAN Petunia x hybrida, Phaseolus vulgaris, Poa annua
30*10360 Phaseolus vulgaris
(jbhnpn 6£1 bi3boi>nb ^mc3hi5^m6i 9dn93 93^>C3&nb:>e9n. 9^3^(^336300 n6gn>)i(jmh3-

6nb d:>b*cncnn (Amdtetal., 1987; Tyler, 1990)
bsbQm&s
iiU3ni Bryumpseudotriquetrum (^30133 Zn)
Dicronella varia (^30133 Zn)
Philonotisfontana
Fontinalis squamosum
Scapania undulata
b3nc36dn Calypogeia muelleriana
Merceya ligulata
Mielichhoferia elongata (ij>h30133 Fe, Cr)
Mielichhoferia nitida (i^h30133 Fe, Pb, Zn)
Nardia scalaris
Pleumclada albescens
5nj,3on Oligotrichumhercynicum (ifch30133 Cu)
Un6yi Cephalozia bicuspidata O^DotBsCu)
Pohlia nutans 0^30133 Cu)
- 425 -
9(3 0 6 ^ 0 3 6 0 OânO^fnODbâO
6.3.3. }i!>0nib<33h3Cn 30flbn36nb 6^3036^ 3J>mbnbi!>303h

6Cn62)C3h 9n6336<b3
59016593(10 bibrujmijbcm îhâmb i O X ^ D 9&hd6oi6o3h3 <3363 3hnb i^ m b -

<33(101. 3939036013 DÔiiD^rnftnb 65h9oig53659o ^Qmbqj^hmb 035(159360 55^ 30 593-
6 3 9 5 9 g0g05. li^hnb <3365, hmSQnb cg3h503&9 o mhjp>6nb9b 033d0O3 5hb36m65
63901363(10 g39b95(i3 pm6obdn3636ob 53(1393, 3(n633b SbmQmg 6000 9 bo35Qv
03693. 013 ( 1 3 8 9 3 6 3 9 8(30(133 3b 03363, 393090 63(18016595360 oj63&5 83b 939g 35,
hb(3 6 2)3 bn5hd3b 901360863301 5mhn‘bm6ly>03h 606(1^3^13. 3^3hnb 50 306(101 033-
6i 9n j,i9 mnymqjb 55(130133 5g58o56ob b iJ9 n^6m6 nb 93g365g 6ih9mj96n0n 5 ^ 3 -
60. 013 553001359066063601 5(15 9ih^m 3^9016033(10^ bo(5oi(5b0ob3oi3ob biyhp36
03363b, 3(13939 90130 ^hm3oi6033(1016, hm930O(3 3(1(5309363 15 39 bo33(j>933g3,
801390 59 qj36nb 9d3h 55^3606, mhcnjQnb, 63301360b p i 9lj33hnb 65609533606
83013063606 363(10 303(101 9 3 6 3 9 0 3 , 3nph3 39563^06 6635 bqj3hm369n. 06936^039
j33y6369n 9(13633901606 d0O3h95 b h gi9 (y 330 i 5j3g56 5590189065(13 30 33^ 0 6
53013390660636001) 9 3 9mbib03m6nb 3^301636(505931185 ' 6 ( 1 9 5 8 93958060b 53630-
0 1 5 (1 3 6 5 9 93 (*> 3 (1 0 6 (13500163690 5390106303 3^9016033(1390 090603606 d0ri3hn
0 m 0 ^ 36 i (93, ij3Q>6 5 3 8 0 1 6 9 0 6 3 ( 1 3 , 8(5363(13390 63035(106 g ib n j^ & i, 5^8016033(1-
0163 g i 6y036nb g36o6d3(i363) 0bmqj0nm 859,356001.
633(10 85363 653(1010 63fi0nb, 010130065 g 5 6nb0nb 3333016001 ^53(10956 550mo-
ymqji 93 003^363 6o3g35ob 013 6y03&o6 '63g53oh36‘63. 83533 6593^360 63936036
8(5365(1339 bbcgbhb3 95 30163336 6115955011, 6yo369o. 563 65h8oioj865 " ^ 3 3 6 0 6
56590 djfiSnb 95^05636360" 33hm3nb ^hn^mhnnb 303^3b 656n00n, (13(5 53801 nb-
5^365 035(10101 553(1(53936590, jh 0160339016390, d0n3h (53393639 6083(30185,303580
g5 633606 bb35g5bb35 65630160b 95^0636590. ^ 3 3 6 0 6 3b 6565(199030 95^11563-
6 5 65(1801595366 jjm9303jb3h 9 5 5 3 5 9 3 6 3 b , (1019390(5 5601(101301536390 qjij^mhnb
55(1383 3(1 65h9mnj9636mg5.
5(j9mbqj3hm0n 65600(101(13565065 g5 8301360b 038153390160b 90195,3360b dnhn-
cri5gn 90^13^3605 6o5(pnb3X n b363330 95b5gob 539903(136390 b5h33s, ,jy336ob
^15, b5bmq30fn-b5033h63fn 9n636nb 53^(195 95 (56013303606 901936365. b6mh3g 59
015306015359 3 3 6 3 6 x 0 65“b36ob 8903(195 80185^3653 5580106305 650163(106 3033^,30.
6.3.3.1. ii>y33d>nb i>n6C)n63i!)3hn 0ib3W 6nb 9^0^636360 - 3iPl-

bflbi!)303hn eîD3flh013nb bn69hn0fl
93959065^3 3oh33059 60-n36 603680 939^63390 oj65 Ij36l»)h503ho 33hm5ob

bm^65h36nb 33b 30933 5535365110, 35bm6h030 95(bo563636n, bm0m 5010 60nb
938935 5b3cno 95^105636360 3333 g55qjojbnh3b Rh9O0m3cn 93 U36l!)h503hn 33hn3ob
65d365h363n. 1980 600956 5b3cnb533 9g5m95h3m659n 5(p8 mfi6g5 cru>3g53nh3305g
6063013560 ^ 3 3 6 0 , bm0m 81^15305636001, 55h3cn33 qjmai0fTi336n C»y336n (65b.
6 .93). oi53g53oti330o 95bm6ho3o g5lbo563636ob 936935 955600136001 1985-1988
693690 633(1 ^yo36 h35om69o 9306096360193 39 30130360b b^56o0o“b5(305, boi0m
to i5 595095b - 93b33636ob (5 3 6 9 3 6 (3 0 3 [2 1 2 ],
^ 3 3 6 0 6 9m3393b 93^03636366 y(1133901306 3jm693 595090. 935h39 59 phmb
- 426 -
6G m ^ n 3P)&3G7)(nn b<?>(f>3b3 ^ o sM&nfna&o
ig3fi(vi nym 93bi6i0nbn Srfb ^b n b 3mdn3&i: 0(536ih336nb j)ip i6 3h 36i1 3^3636-

CjSn pi 3ihi'bnvj)06n, j^ n S i^ h n 3jbu3b3&n, (jynb ihibSmhn 5i3m y3636i, 3n63h>
Cjhn 6noin3h3&36nb ajmbnb^^S^hn 6h^53nb pih(r>333i J33<336n0nb 0m(jn03&nb
338535, 6nipi£,nb j ^ m c g n ^ 0m6mi30j)3h36nb 93036^0 0^> np36(yiq)nunh36:>pn
3C33o6(!)36non 0m6i03Q:>. "liyoo^nb ibipn ^n3nb 9 ^ 11^ 6 36 ^bnb" 930016333^80 hin-
63 i(,i6ih3(i3Cn 3n'b3'b36nb 9^95363 33b b3hbg36mp:>, 3 i 5 h i3 6icn3pn nym, hm0
i0 3h(THJ3b3n 9m6i6np3m636 3i3h9n ih b 3 6 x n 0*363 6i3hcn36n. ObmQmg 3ib 83d-
936, hiu id 9^ 11^ 6 36 3&nb j>i6bn03i pinBym 3^mbnb,i363(1 p i b^P^bnb pn6i 3ny»b
ib33i i 3n, i 3 im33Q3jb3hn pii3ip3&nb ‘bm<!)n3hcnn 3n<b3<bn 6i6n0m6hn3 J)i3m3cn5-
e>-
(jynb pi‘brii536nb bn33lynd3&n

^ynb pii3ip36nb 3hi3i 0 n b b 3ip ib b 3i bn93,jri3 nb 6ih3 mj35i p i3minp363Qni:
63 3u36ihnb bib3m6i'b3> bhpnb qjmh0i'b3 (0i&., 6id336nb 6inh£,3ihn gi^m^î),
ib iib 3, 5 i3h i3X 3^ n^ i 06nC<'33 (pi&0m6n 013 o^hpmbn, bn3i (?03 <b[?3nb pm6npi6,
3jb3mbn(3ni), 6nipi5nb ji63&nb jorr^m g^h b ^ h ^ ^ h ib i pi bb3ipibb3i
b^bm hnb 3hoin3hcnj)ipiq}ih3i'b3 (nB^mjbnîijni p i i0 i 3 3 phmb, 3ihi'bnlj36nb
0m0hi3Q36i i6 i 0 n3i£)hn 3nhm63&nb 0i33Oihn 5i3ih 3b 36 i). j,i3 mi5pn0 ^ynb 3v
cnmQrri536bii3 b3 nhip 3^nhcn ihb3&30n pi‘bni63636nb 03^6^3 "(ijynb picvp^nb"
P3iQihnh36i. dnhnoiipip, pi(bni636nb hin03 im6ih3^30n 0n(b3‘bnbip0n 3n-
i)D°’3636nb 3hm^3b3n bi^nhmi pnpn bnqjhonbn03, 3ij,„ 3 n h 3 b x n n6cg3junnb, bm-
]>rri36ncn pii3i036nb, [5601330361101 g,i0m6333pn pi‘bni6363dnb Qi 363i 33 n6^pi-
jbnii^nnb (3i&., 3ihn036nb pnpn i,m6u36^hiuni fctnb 3nhib, pibn5d3h36nb ib-
0mb Symqjn 6yihm) phmb J,ibicn3i0nb5n6363pni ij,h3cri33 0nihm(ji0c>36ni6n 33-
100-1 100-
90-
o> C
0 O
Abies alba
80 - /
1
O
70 - 70-
60 - Picea abies 60- Fagus
50- ^ --------------- 50- sylvatica^rf**"*"^
40- 40-
30- 30-
20- / Quercus
CM
O
1
J robur
10 - 10-
0 — 1------- 1------- 1------- 1-----r 0- — 1------- 1------- 1------- 1 1
1983 1984 1985 1986 1987 1983 1984 1985 1986 1987
6bb. 6.93 5,]h0i6ni3n 1983-1987 66 *>3 9l536ih30ii 9^^636361101 J>b0m6333cun3&n &i6bne3cns 336535
9hm9369n: Kandler (1992), Huettl & Mueller-Dombois (1993)
- 427 -
0360b 5^3036^. hmgjbiu 00033(136(50503110 0o556oi"bo 60301001366 30163(13(1)30 6 ^
9m96333'b3, 006^03^° (3(13006301*60(305) 0)003(10365 imdSQQjb^hn
055350360b 339cnb333i 3n.
9 3 ^ 0 3 6 9 ° 5(1636300 ?P3hii3330iTi2>nb 8036305350, “^ 3 3 6 0 6 56500 ^nSnb 95-
'605636360650130^' 05656560501363005 55(1333300 bnSS^nOs^^n 63(15010 p i (jbo-
33300 6n0636n: bhgnb 561^18500360 (055. 0jb0on9rihnb36nb 0590130365 - "yshys-
(jnb 6393"), 606336065 05 03010103606 5^ 30 0 10 36 5 (Mg-nb 6si03&m&nbsb y30cnQ-
036s 303(101 5653013560 6063360, Fe-nbs 05 Mn-nb 65303&m6ob5b - 303(101 sb sc-
5 5 ^ 0 3 6 0 ), 6n6336nb q]mcn036nb5 05 (jm(i)36nb lJ50i3D0n 366360b 55600165, 6n6-
3360b 6580153365 (35h33&ob 5583663(1365, 6sd33&ob 6590130036x^1 (301^3606 55 -
80930365 - "33hub0ob 63090b bo60hm9o"), 33663(1016 05 ,501^3606 633h36ob bn0 9 -
h503 05 qj3b30i5 bo b ^So b 559366(1365. 6063013560 ,^33606 35hsob 6950195(13(^1-
60b b5qj3b3h36ho30 05b5bo5oi36oboi3ob 559moymcg5 35^3501(10360, (10190360(5 050-
y5h3&30O5 553033(13036565 (0 - 553qj3h30365b 505000 5h5 sj3b, 3 - d0o3ho 5 5 3 -
<33^30365) 05 6063360b ^591^1(33365^3 (65b. 6 .94 ). ^y336nb 05bo563&ob 930356365
y3305 J33y5650o 5h 5hob 6(35605(1^0^(136300. 53h956o50o 6063013560 ^ 3 3 6 1 1
00130365 m0653 05^05636303650 (05^0563606 I 6503363(10), (1010365(5 6o6336nb
35I1535 5(^6336 11-25% , 658350010 0 5 ^ 5 6 3 6 3 0 3 6 5 0 (II 6503363(10) - 26-6 0 %
939016333590, 6oi0oi d0O3h50 0 5 ^ 5 6 36 30 36 5 0 (III - 65(3363(10) - 61-9 9 % ghmb;
IV 6503363(150 00130365 ,jynb 05^3335. 536^(10580 09363636 5-6 bsq^b^hb.
(1565330(133005 3ob3503ho 930356365 05560013600105, 556653301(136001, (150936003
b5b3m6ooi 93j96 o0 ^ 3 3 6 8 0 . ^ 3 3 6 0 6 05bo5&3&o6 930356365 93b5d03630O5 53hm-
<3301^015(1503006 93933016001 - 0603(156001300 0300,4(13606 55601936360656 05^0 5636 -
300 63360 gojboh036n56 bb35 033(13680.
^y33&ob 05^563606501306 059sb5bo501363005 3o6hm 60311(3390 05bo5636nb
bshobbob 0000 9115350033110136365. 563 9555000150, 96oh, J3056 6 05055^3 9m‘b5h-
00 b3 9(5365h3 303(101 d0osh50 0 5 ^ 5 6 36 3 0 0 6566, 3O0h3 801(10560016 9o63h503hn
6o30i03h3636ob 00O05h cpsh^scgSn 55605536300 93bri 630o. 3hoib5 05 09533 ^P&n-
0^13 05^05636360 600056 605003 93nd03&5 55ob5h0oib 56 939(joh03b. 560,3010,
(jyob 051bo5636ob 93b6530ob5b 55b5oi350ob6o6630o5 30163(13(1)300 ^560^5^06
y 3 3 0 5 35 h5 9 3 (jho.
,jyob 05,bo5636nb 0nb3<b3&n, 65(160190165 05 9060065(130165

(jy336ob 30(13300 95^56363606 65h9oi9oi6o6 0hmb (36m6o0o oym, hoi9 390-
05(53h 95(530(1013360156 3(10150 59 3(101(53680 8016560030166 353(106 0606036(1.
5b30io 9mb5lbh365 dohooi505g 059yh36x n nym 6503^3606 63533 (135:1(505^3.
5(j9mbq33h(Ti9o 93533 (135^006 9jm63 3019301636,1)360 5hob 93530b 65h9m9J9630n
55^360 (S 0 2, N 0 /N0 2), 015306303500 9353360 (H2S 0 2, NHO3, NCI, HF) 05 H .
03-20 65333606 935660056 33(1013580 (656. 6 .95) 05 6635 06036(3(1039
(13500163690 (R(i0o0m3oi 503hoj)5, 05301605) 5569o(i05 83533 6306360.
6503J3611b 63530560165 05801300363005 50500016(1030 o9obo36ob 63(150^33 95
b5^h56b3mhljm 655ob(jh5036ob 5hb36oi65b 3. 1970 600056 33(1013560 55boi9nc
oj6s pH-ob 90165(539360, hm9036o(5 03hy3m605 4-bi 05 5-b 9mhob, '6015 505005b
jjo - 3 05 4 . ob30i 506003690, b505(3 6ob0o 05 <33h0m636ob 6ob0o56m65 b9nhn5,
03530560165 95(p50O5. d0o3ho 93533 6306360b ghoib 5hb3&30O R50mh3(3b3nb
3ohm6369o bob doh9o pH-ob 60165(330360 mho 31101330001 303(101 05&50O5, 309(13
35(1300.
63533 93301^0(50360, o9 ob 006303001, 013 (101501(105 9 5 Cno i)ri 9 3 oi6 3 6 (i)3 6 nb
- 428 -
660)f!in3n a 067Jfn0 l»6 <f>3b'3 ^ 0 ^d&nrtObO
6ib. 6.94 Picea abies-ab jihjnb 9;)3unh;)&s. 3hm(j36^36n 5i8rnl)bbi3b ^hj'bj BnBjjdnb >^>h&3^b
(Muller & Stierlin, 1990)
- 429 -
80065(^.0060 a ^ a n a a fn a n w a o
6ib. 6.95 33hm3nb iyhrymhni'b] 3?p3ni6m6nb ihn^njxn gi^n hoî. 8gs3n:>6m&i 5i6nbi‘b(p3h36i
1 Ji-'b] BjcnBigan (Hettelingh et al., 1991). 8016^36360 W33S jjm6^n636^3&nbsCT>3nb nb. Kuttler
(1984), (Coziol & Whatley (1984)
^rnjbn^hm6i, 0633336 bibbnSnob^nm mh5i6m3bnb 30350m g 5l6056365b, 6 55.,

cgmcnQnb 2)ng33&nb 63^1*16360 g 5 60630b i3iin^30nb 95^111^636^. 5 3 m 3 0 5 b l|)36 nb
9 5 0 3 5 3 0 5 6 3 6 5 0 9 30 0)036 5 550 0 10 6 30 0 16 <30^01101^180163606 5 5 6 5 6 0 0 3 6 0 b 13 3 5 9 3 6 5 9 0 1-
6 5. 63ho0 o j 3 b 3 3 6 9 n 35^ 139 36 0 6 g 5 ym q jo b 5 b 9306006365 jh m 9 m bm 0 3 &nb 56m9 -
5 0 0 36 0 9 5 33(139360 '6 0 5 6 9 3 6 5 .
6 0 5 9 5 5 0 b g 50353O56365b (jyob g 5t)056360b 65h0mj3650o 360^365 9090 86033-
630m65 (65b. 6 .96 ). 01539530(133059, pH 7 5ohm63&0o, bb6590 C a(H C 03)2 556m-
h3ijb3ob 8393559 69365 (53,40560160b g 5j 3300i365. ^3^3 605955,00 50ob 80363-
9535g pH b55hd6m605g 5h 01530365, 855(158 5056 031^0365 55801063001b >5003-
0ob5 95 hio6ob 65^036^65. bo0oi5lj3h 60595500 8080115300b h35ji503&0b 0 3 9 3 5 5 9
69365 556b5»)30ih36ocn Ca2+ 95 Mg2+ 55801(13^35 95 obo6o 560016360156 3(10159
R5doh30o 6yi0ob 83033016001 6500936056 605955b. pH 4 95 6 qj5h50360o oiob5-
3303bob ijm0303jb36o i)5cnooi636ob 0080115300b 03833016001 5^303636 63033(130
qj36j(505b. pH 4 j333ncn oiob5-3o63h5036o 000365 95 83,550360b 309(101^09360
55959056 bb65h9o, b595(5 5030o6nb5 95 0do03 83^503606 oi53ob3q)50o 00163606
(15019360165 85,530016b. 6059550b 9g53056m6ob 80185,5365 55h3<n33 06333b 01^1556-
300 b36bljh5ljob 95000b 06(53660301606 9515385b 95 Sn^hncgoûoob 3(101153606 33-
630365b. 6059550b 950353056360b 06003630^1356 85633636306 65h 0 rri59 536 b 6 5 -
tb503ho <>5010016360015 95 60,5(15,536001 bop5ho&3.
- 430 -
^GtJ)tf>n3 naaG 3 ffio b ^ a u ^ n g6d6nA0&0
Soil acidification
Direct damage Indirect damage
Increased ratio Nitrification Metal toxicity ion- Binding 0) mineral

H*/[Ca**, Mg1*] inhibited cora petition nutrients
inthe rhizospiiere highNHJ/NOj--ratio (absorption, adsorp
tion, precipitation)
a) High permeability a) Proton efflux: a) AIJ* a) General: a) P(boundasFe-
ol cell membrane additional H* Cat
— andAl-phosphate
(release of bases) acidification b) Heavy metals ions
b) Destruction of cell of itiizosphere Fe*' Mn2*
> b) Mo
b) Particulary:
membrane
b) Ion competition
(plasmalemma: NHX*.
Al^-barrter) c) NH* toxicity
Mg** — H*
c) Dissolution of (carbohydrate NH*\ AP*.
cell wall deficiency) Mna*
(Ca/Mg-peetinate:
tissuedecay)
6bb. 6.96 Bnigi^nb 9gi3ni6m&nb 5*30361 3 nb ai3nb3&3&ti3 g i d)i36ih3cni 0 ^ 3 3 6 ^ 3 (Isermann,

1983)
dy33&nb cgihmm0*3li>*6o*6 o g*<bo*636ob 133*036*90 bo03(j)ni036ob 3ho»m6pnm6i

6ih9iTinj3636b 8 3*33 6*p3j3&nb, j>on0*lj3hn, 3Q*cg3hn g* bnm^^hn cg*J^mh36ob 3h-
01^60030 0mJ03932>nb 330 35*9 (Bib. 6 .9 7 ).
9*'bns53&nb ghnb 0n09n6*h3 qjrfbnmcmftfvjhn 3hmu3b36o 33h33hm6oo» 83b-
5*30000* 0bm0m9 6*6o0n6ho3. y 3 X * 838016333*80 6o*g*j,ob 9*83*30*636*
860836301113*6 hmgb *bh303&b. boooiûh b36bl^h*(j‘b3 0n<b*hgo 6*d3ob 63ho0o
03363360 6 *3^ 30 36 30 0 * 6o*0*2>ob "b39* 03363630, (10180360(5 j33<336n0ob 9*80nb
bihj'b3 303(101 6:^*03(1360*, 3ogh3 303(101 ph8* 3mho<bm6lJ36o. *0ob 5*801 b33&b
6o*g*5ob ’b39*3o hxn a*8m3hfn6ob 303(101 9090 b*q3hoib3 383^136*01. qj3b336nb
9*6*00 bo3hi530n 5*63oot*h36ob* 9* <33b33&ob bm*|fn3601*6 bo06om<bob 838-
iyih3&ob 5*801, b3 8(536^3360 ooi3ob3&36 6*5036 b*^3p6 6030103113636b. (10193-
b*(j 6*h|»3330o ^ 3 3 3 6 0 om636ob (5*6^3301(136001 Mg^+, K+ 9* Ca2+, 'b<T'633h
Mn2+) 6 *i0 36 ,Ti6ib 38*^36* 8 3*33 6*03J36o, (10180360(5 *8 001636b q}oioi03&o9:>6
5*801(13(5^336 (6*b. 6.98 ), 6*h8oirvJ863&* 93030(50,50 9* 0016360b 6m6*b6mhm6ob
9*hp3333&o.
013 il58mbq33h30o 6*03^60 830(5*336 ^*h6o 11*01936016001 *^01,5013*6
6*3hovj6b (NOj-ob yn06o6*(5o*b NHg-b* 9* NH4+-oi*6), 8(536*hob bhg* 01*39*-
3nh330*9 ob3 8*0iXfT'6b, (1018 b*6oi0»no»9 000(31633* 5*6(b*336ob 30 33^ (ob.
01*30 3.3 .2 ). (1*95*6 Mg2+-b* g* K+-ob 8380*1333*80 b*J83 53*feb *9300*9 5*9*-
*950036*9 XD936^36oi*6, obo6o 5*8010,5*6360*6 8(536*hob 90033(136(50(136x0
6*6003609*6. 9n0(Ti8oôb ^,3^nb, 30^3(10^ (MgS0 4) 9* 3*oo38ob 8*563<boob
5309*b8ooi 83hR33ob 83833016001 83od036* 6063013*60 b3 8(536^336ob jomhm'bj-
60b 8363(136*. ^y336ob b3*601136* *hob 0boi0oig 30(13300 9* b6h*qjn 0*b8*h36*,
hm030O(5 3oh3x * 9n 80^53^360(3 (i3hdm0 30 3*3hob p*6o6d3h36ob) 5*Q*0*b3ob
- 431 -
Pollution Biotic Direct impact
Sulfur dioxide, nitric oxides, factors through humans
heavy metals, fluorine, ozone
Attack of parasites
and opportunistic Inappropriate
Direct damage infections forest management
to l<
Litter removal,
Shedding of needles forest as
Dysfunctions of and leaves, disturbed pasture use
stomata regulation, growth, decreasing Insects
photosynthesis, resistance to
growth, development frost, pests etc.
1 Game damage | Inappropriate
choice of tree
species and
provenances
Leaching of 1
mineral substar ces Deticien cy ot
/ mineral n jtrients
Viruses
bacteria Impact of timber
fungi technologies
...............
Leaching of Nema odes Recreational
mineral substances Disturbances in Disturbances of activities
nutrient uptake water uptake trampling damage
Soil acidification
Release of toxic Damage to Impa rment cf

metal ions fine roots mvconrhiza
’
m
Alterations in Drawing of
composition of soil ground water
organisms
6*b. 6.97 ^ynb Q^Qjjnh^&nb QnVfao&n (Eisner, cit. Hock & Eisner, 1988)
6ib. 6.98a,b 9^30 63n3 o&nb <b39mjd3P3&> (Cronan & Reiners, 1983; Glatzel et al., 1987)
- 432 -
a60 )ff>mr)&a6 3 (n0 u & ^ a u ^ o ^danfna&o
^ 3 8 3 bi55hdcn3 oqftj^b 33(1 90153153636.

ijy33&3n bb|)336n 6fl3cnn3h3?)36nb 6iQi6bn qjhnig Qibnp^hni g i g:>9my>63&3-
Qfii 633(1 bb3ig>bb3i 1530^3 (6ib. 6.99). ibm^nbs g i 9n635Q3hn 6fi3cnn3h32>3&nb
6niQ i5n g i6 33oi3nb3&nb 306n3363Qm i535C36gm6i3ii5 311 83^0365 5i3mn63nmb
9^11^6363&nb 5iph3i336i, 56 Snhnjnai - 3u36ih33&nb 9:^336365, bBmh3g } 9 n^-
m9 ym33C ibip 63pn6sgb 333dpro 3m3cmg63pn l53Cr'C3^32>nb 013 bn^h^hrVbjbnb
9m^>6 i.
Atmospheric input
i- L , Canopy
ICanopyf
exchange
Growth immobilization
Throughlall
in biomass
Litter
I Soil w aterlPlant uPlake*
^Decomposition* Organic
Soil
Soil layer
leaching
exchange*
Exchangeable
soil pool
Soil water (mmol m*2)
Soil
weathering Element pool (mmol m'2)
Element - flus
Total (mmol m*a)
soil pool •calculated values
Groundwater
Forest with Mgbalance Declining

no visible d a m a ge forest
r
42 13 »l 275 29 7 J 146
*l ’1
29 iI 32 ►1 22
113 63
6* . 6.99 6x * 36:>hnb
U3On0D&* 3*6&i639nb
‘bodmj93936nb 93006*6
(Horn et al., 1989) | | P o o ls (m m ol • m "8] • Flu xe s (mmol • rrf2 • a '1]
- 433 -
90066 fo0 a &0 LStfouac? aân&w nam aao
6 3 .3 .2 . n6qjh^6n01DCn bSDjiihflb 03O1^6Cnj0DCn d ^ D b H b dCP16-

ic a h n a n a ^ 6 3 6 ^ bm ahnb o < so m
03-18 b i 32)35 nb 9 j i 603609*6 C02-nb 3 D9 b 3DCm6 i 3 *^ 3 oibqj3tim3 n gbnBym

0ii6gicni6m6ncn 8 ^ 3 6 ^ , bmpm 83-20 b*3$36nb 33 *b 6 ng:>6 3b Shm ^bn b:>j)hd6m6-
C*g 6*dcn3hg:. (b*93*Qmg 1,3 jj.q-1 6 x n 6 *g 9 n; 6*b. 6 . 100 ). ghmnb *9 n S ^ f^ s Q -
9 n fihgn0n3cn j^hnjjnb ^hn^mhnnb gngn 6*6nQn R*9my*0n6g* b*b6*3-bicn3-
b3&*g, P3g^8n6nb 633hn g*3b*b03630n h^nmBn :>oi3nb3630n nJ6* *g * 9 n*6 nb
9n3h, gsn6yn n6g3bljhn3Q 3h*3n 5*g*b3g:>, h*b*i5 01*6 bg33g:> n6^36bri3hn 9 *^ 3-
hniQ^hn 6h363nb 9jmE>3 m936n. y33C* *3 8013036:18 5^81^0630^ 6 nm9 :>b*3 n g i
6 nigi 59n 6 :>b9 nh6 sgnb h3'b3h336nb 939 ynh36i gs C02-nb 3 Qbi6 >9 nbn h^mg36m-
6nb jp g ib 30 i i^9mbq33hm3n. >3 nb 3}hip3p3h:>g 3:>3h9n S m n S i^ 03cni6nb
933 l5330m6 i 3 .
6sb. 6.100 6ib9nhmhgi6jnbi 91 33crb6nb 5^16(336(ihiyn^bnb 3^361 s,50nbq)3hm3n 83-18 bi3i36ngi5

etT]6e]. 1958 6pngi5 9v>3nh33&3&n ^9361 8136^ ptnib rn6b3h3i^mhni3n (3i3inb i36d303&n); 9i6-
1693 ihb3&3pn 0m6iU3d3&n iSîh^ngnb ynexD^Q^ (Keeling et al., (50,4. IGBP, 1990 &
WMO, 1990)
6*b3 nhmhg*6 j,n 6ygnb mhoijcoi:>6, 8301*601*6, m'bm6oii6 g* N 20-cni6 3(101*9

533390636* i0n3i^nb h3530*(j)rrih*g. 3b 5*^360 8bnb 301^03,5*0(31013*6 h*gniun*b
anncnj3 nb 3333(3nh360*g *^>*113636 g39*8n6nb ^39*3011*893, bmpm 0603(1*60013011
hbgn^nnb 90 %-b 50 - 901*601^*336. * 3 ob 5*0m, 3*3hnb 0336360 01636*. 6 36 36 ^ -
30 "b*oi63hnb 3qj3j^nb" 553m 939806*^3 5*6*^01636300* bouon5b0ob*oi3ob b30-
b*yh30n ^3933h*^3h36n. Hb*oi63hnb 3<33jiiob" 5*90196333 63636hn3 5*“b36b 36*^3-
6* n6g3bvj)hn30n *j^n3m6nb 33g35*g 6 *h9 mj96 oQn 5*^360, 5*6b*i30ih3&n0> qj^mh-
J0mh-6*b9nh6y*c6*g36n. y330*qj3hn 3b g* 63636(1030 5*b36ob 5016(536 ,ih*t5onb
3m0sj)36i i>ng33 3qjhm 'bhgnb 3*3hob <33636116 ,53933^,53(1:^. *hnb 3016*1538360
hoi3 g3gi0n6nb b*93*001 ^3033h*^3h*9 6n6* *b60330ob 33*b6ng*6 8 oio3 *^>* g**b-
00136001 0 ,7 °C-noi. h*b*i3nh33pn* *b30i 9m6*iJ3936b qjhoibno*g 36g* 9n33g530i,
- 434 -
6 Gmrtn3 n aa 6 T)(nO u & r ta u ^ o sfodsnfoo&o
(159556 ii3a3Qhi^3)hnb 8009065(130165 9390113 535036590165005 (nb. 65b. 1.4 4 ).

9or)po p39i9n6nbai3nb (53033li5(43hob b503500)9 550010130000 0m05(5365 b^Q^li ih
ibit>53b n9 ShmyQbjftb, (101003605 0o00o65h3oi6b bb3595bb35 (135001600. (135001-
6 x ^ oiS3nb363h3&36n 013 55956(1360 95001^19363005 (15905500b 65056b'b3 95
2,:>6bMi3cnh36ooi Sb^hnb 0ibQ6nb 9n6i0n^bi 95 b ^ o b 9063635^3.
ilj)0mbq33hm9n C 0 2-ob 95511013365b 5556605 0h5350O 5b33j^n: ( 1) 65b9nhmh-
35650 5hnb 0301,40153(501(4(10109300 mh556 ob036ob b5],336 n 603010311365 95 05365-
h33611 50 930oib33350o 53330063605 0ob 065(133303659; (2) 65600(101(135650,
n6gh 56ooi30o bl33jljhnb 0 ou>6 oij0 nb 550m, 5hob 50o6 gob (135305501^ 0600363-
0013560 303036^0, (i5055 03000365 55001063001b 93950060b 05365(13300 b5qj5hob
335305; 013 505opo 31)6365 05365h330m6ob 0600363001356 535036590165b, 3b
0153^015359 5530365b 9 m5b936 b 65b0oh65gob 50m6503h 65p56bb3-
^53(130 C02-ob 0305330m6nb 0oi05v53&5b5 95 05365b3360b On^h COj-nb

Sou&njEfcb Smhnb 5hb3&3pn 3hoin3^*014530oho
05365I13360O 6 nm05bob 3hmp3j5n5 dohooi5g59 0 3 ^ 3 9 3 0 0 5 C 0 2-ob 63636(1030
65115536001. 550mi3033&05 363363b, hm0 q3rn^mbn6cn3‘bob 36ihob 5 5 ^ 9 5 03od-
0360195 5li|0mbqj3hm0o C 0 2-ob 9CP3b9P3m6ocn 5565^,01636300 4016536(5(1550361^
(95560. 350 |i0-l, h55 033G5650365 35 Pa $5(15030 66335b) b5033h 0 oi05(53&ob
0300163335905 io. b50i&3h9 o C 0 2-nb $016536(5(155006 TOO Pa-093 6^(19* 06333b
65365(133606 06,536603(1 bhg5b C 0 2-ob 501536,5(155006 55mh05536ob 0393559 053-
6ih33&o 0^936056 303(101 66(150359 (95 303I101 "6(150)59 56013(101636") 95 05Oin
6nm05bnb 3h0i93j505 05(j>50O5 (65b. 6 .101). C 0 2-ob 05^500 2>m6536,5h5503&0b 3o-
(10163600 6553360 65403659 5(1056 55bb6o0o 95 03b5650nb59 6y0ob 501(101^03655
05&50O5, 500(4^10 535930b ^0(10163600 ©oibshg 0536511336b 933d0O5Oi ©3,50
1990 1991 1990 1991
5bb. 6.101 a,b Citrus aurantium-rh b h g i COj-nb 63 636^ 3 (• ) p i S m S i& j& x ( ° )

(Id s o & Kim ball, 1992)
- 435 -
8006660 D60 U&fnOUTlc? 3 ^& n 06 AOn 663 O
h*mp36pi6noi CC^-nb 930131^36* (tfbh. 6 .23 ). 3 *5 h*9, 01536*11300 bb3*Q*bb3}

bib^mbiSn 5*3mj)303336n 363363636 5*6663*33639 9* b m j^ h b*3nhnb3 nhni 3 pv
6*1533366 [ 161 ], CC>2-nb ^*h6n im6(j36(jh*un36nb *h*b30b*yh30n 5*3036* 9 ond-
036* 5*3fnnb*vi)fnb 8*5., j0mhm3p*b^36nb 5*9*3b3&*3n b*b*036onb 3*hi53Q36ncn
(5*^393*) *6 3n33h^hmqj30n cgmoic^bnb 5*63001*1136*00 (3qjhm qj*9 *hn
Ijbhncn 6.23 8(3361113011 g i 3(336^3011 oii6:>bi'bm6igm3&3&nb S^bidpm h3 1 ^ 0 1 ^QmbqjQhman CO2-

nb 510 ^ 9 16 36 1^ 3 (Strain & Cure, 1985; Farrar & Williams, 1991; Hunt et al., 1991; N obel , 1 991c;
Woodward et al., 1991; Kom er & Amone, 1992; Weigel et al., 1992; Coleman et al., 1993; O verdieck,
1993; Thomas et al., 1993) (+ îbhgi; - 91:133001361; 0 31330300; X 6^0^3333300)
SmtKgmb^Gjbn g i 5,1630011*1361
33<bfnqjnpnb bnbj3 0 /+
qjmcnonb qjihcno / ajmoipob 39hi0n 60161 0 / -
qjmaipnb ojihoinb 06 93^ 0 (L A I) + / X
p>3hmb ‘bhpi +
Qi^m(j3nb bnb0nh3 + /-
<33b3nb ‘bhgi +
011610)1(190161: qj3b3n / ["3(101 + / -
y3i3n0ob qjmh3nh36s - / +
oi3bpnb ‘bmSs +
qjmaipnb ^0181 91 9163b361 + / X
8>p&(ncn<b8n. 09hi(yi dibnb 3hrng3ftyii gi 8(536^3011 Bypob (133080

63ljni-gjmljnbn&cn3'bo + (C 3 > C 4)
RuBP-j)ih6m jbnoi'bi 0 /-
C A M <336^501 +
9(VjmJm6ghn3(yi b36oij3i + /-
3 ij(i36nb ^hi6b3mh^n + / X
33hicn 9ibnb 3hmg3jijni +
^h i6b3nhnjm - (C 4 > C 3)
Byonb 5160^36 360b 3q)3^j3hm 6i +
I'bm^nb 6i 3my3636nb 3q)3j^3hm6i 0
93 hic 3ib i 3 o 8063(110360b 3016^36^11301
33hsen 3ibnb 3(10193^1301 boyb33n, 631031b! g i gi9oi936n b 30(10163680 +
8(5361113011 ou6ibi'boi6igo>3&36o
bn6ioionb 830131^361 + /-
6yonb 330>3nb36i + /0
3n63hio36nb 330i3nb36i + /0
301633(136(301 + / 0 (C 3 > C 4)
qjmoiQ^bn, hm0C3&b*i5 qjmoionb qj*hoinb oi*6*cg*h0Pi6* 00h*0 6m6*boi*6 - cgmov

Cnb b33i5nqjn*)3hn g*hcnn - 9*6*90 *^301). 6*b3nh6y*06*036nb ^*h6n i 016(336-
ji)h*i5nnb 83935*9 *bn3n0*lj)36nb lt)h*6 b!3mhli)n qj3b33&3n 33(^036* b*5hd6m60*p
5*n‘b*h0mb. 3b, oi*3nboi*3*g, 5*0mn6333b 0 n6 nb'b3g*/0 n6 nbj339* 6*6n036nb 93-
cg*hg36nb 03i530*b, qj3b336nb b36cnj3nb n6^36bn30i6nb 3m3*(j36*b 9*
bn6oi3‘bnb Shmg^j^&nb 5*3mymg*b 0 ^ 336 0 9 *6 . 013 C02-nb ^*h6 2^6(536^*-
15036b *h 033b*6 * 036* 9* bb3* b*i336n 6 0 3 0 1 ^ 3 6 3 6 ^ 0*h*5n, 0(536sh30n y3* 3-
r»036nb* 9* 6*ymq}36nb 5*63noi*h3&nb 06,1)36^3016* i03630m6b (6*b. 6.102).
- 436 -
a60 )<f>n3n&a 6'360 g6d&ncf>a&o
6nm]03936(!)36nb shybh9 m6 n xn cn:>6:>qph0m&i &:>9mn6333b bhgnb Shmuobnb

33lbc13e3ib g i (^ b n b ^ B iix ™ ^ ^ gijsncn^bib. (bmj)igbg 9 in 6 (j nbQ Ri6b, hm9
C02-nb 9 m9 i^ 36 s 3qjhm b^pbiyh^pn o;j636mgi 9(536:>h33&nb-
01306. ym33p 9360)6333^90, ]6 g i 6S3ncn3ipnb6n6mcn, hm9 y33Qi ^39^101 9my3>-
6npn 6^0012(3033^ fiîh 3J>xn;> iDOiKJ^X 9i53(»h33&'b3, ob&mh^mhnx 3nhm&3&9n
gi ghmnb 9 mj)03 n663h3*c 3 n.
jibpmpb 6ihb3p9n ihb^b^pn 9j536:>h336nb bhgnb 06(1)366031*1606 930 ^3656
gP3i6g3gou>6 93ndp36i 5bh^333pn 5ih9mgj,36i 9363 nj96 ib n9 nb 93bib3&, 013
hmj,nih 93 nu3036 mg636 9(336^3360 C02-nb îh6n jjm6i536(jh:>(5n36nb bi6j)hdpn3n
5P3C36nb 3nhm6369o. 6 i 9 ihbn g i ib6g33p36nb 6n6soi gijjm6b3h3363pn qjmcn-
C3&nb iSipn'bn 9n3onnon3bb C 0 2-nb 9 b(piQ ^0156(536,^(3056^ 95 6:>533&nb g:>6ip
jpbb5 npm6 bb 9 mhnb :>hb3&30 3hcnn3hoij>:>39nh'b3. bd 9 n9 bhoi3P36 ncn 6pn3hn hj,m-
0360b i6ipn“bnb 93g3&ig 9 np36x n 9016x530360 6n6i^9g35m6hn3n>; C 0 2-ob
îh&o j)m6i536ljhi(3nnb 6^3036:1 9hi3ipn bb3^gibb3i cgyj^mhnb
0mj03936ncn. C 0 2-nb 9 ipipn j)m6i536(ih:>(5036nb:>g9n 030^1001001503110 (RuBP-^ph-
bmjbnpi'bnb ^0301606 030unh36i) g i Omhqjmgmj^hn (9 ij,., qjncnpnb qjihcnnb
n6g3jbnb 9 m9 i^ 36b) 03&33&i 5^001063339:) cn:>39i 3nh33p:>g 9 ipipn 3hmg3jt5n3pm-
6nb 36ihnb 930(5oh36:>b. 3b 0013036:) *51130133 5 * 3 0 3 6 ^ 9mibg36gi 0(536ih3Otb
bb3^gibb3i 3530336b 9 mhnb :>hb3&30 501653113615030 oii6>qjshgm6i'b3, 9*5-, C3-
9(336ih336bi gh C4-0i536ih33&b Smhnb, bBhiqjig 9 m'bbhg 6inh6ipib36bi gi
9hi3ip6pni6 6ipib3&b 9 mhnb, :>5h30133 b3 0(536:>h336b 3 mhnb. 5*hg:> :>0 obi,
b(j>3nb 30i6jlyn6nb 3hmg3j^n3pm6i (013 j,i3nou>pnb6n636cn C 0 2-nb 5016(536(1^113-
6nb 03ji6n‘b9b) bbj,hd6m6pbg 33h 9mn9^36gi.
55bMJi3ipnbBn6363pni nb 5*h30m36:>i5, hm9 6 3 6 3 6 !^ 3501606(5303600, 9 hi3^p-
6^nhn 3ham3hoi5:>30nli36nb ^63601606 5*0m, C 0 2-nb 5oi6(536(ih:>1500b 9m9i(j)36i9
93ndp36b bx^lS sh j,i9mn63nmb 6 nm9 ibnb 93bi6i9nbn 0m0:>(536\ 5^66:15301-
h36ncn bi6{jhdpn3n ^hnmgnb J,i69i3pm6i9n. 3qjhm 930>^g 9fnbipmg63pni y330i
hoi]p bnb(i39i 9 n ^636300 h3j,3piijnnb 03j:>6n‘b036ob 5iau>6i6h36i. ym33p 030-
aib333b9n, 6 :>b9 oh6 :>go6 j,pm6ip3hn 6h3635 bnmbqyhmbb g i S(50mbq33hmb 9 mhnb,
i 9 3p3036li)nb pncrimbq^hmbb g i 3oghmbqj3hmb gngn bn3h(333&0n 5(163601606
5i9m, bij)9 img dpn3hbg ihnb 5i6m6ib6mh363pn (6ib. 6 .103 ).
iCo9 ivyib U3pnp36nb 93bidptn non^w

ipn05v5nb (3300036369^, “bboi63hnb 3qj3j^9n" 9m6i6np3 5^‘b36nb g^5hm33-
6iboii6 3hoibg, 9(536ih330m6i‘b3 jBgb 9 mibgn6 m6 3cghm gngn 5^3036^, 3ngh3
C 0 2-nb 393>pm 3033^9^ 6n30in3h36iau> (33pibi gi <bhgi‘b3. i 9 9n9ihoi36noi, $pn-
9^n b (33ip36im6nb bihnbbn y33p^'b3 6i$p36ig ihnb (36m6npn. gp3bgcp3m6noi
gi5hm3npn (3mg6nb biqj3d33p(b3 giyhg6m6nai, 3<j|hm sg3npni ipn9i(jnb (33pnp3-
&36nb (hm9 p36 n(5 6 ihb3p3 n ihb3& x l53CnCDÔ^,<':iO 3cghm b6hiqjig bg3&i) 93 g 3-
6360b 9303^36^.
93(53mhmpm536nb, mi3i6mpmj,36nb g i 5pi(3nmpmj,36nb 9n3h ipn9iiinb (53pn-
036360b g3g^9n6nb 9ib 9 (i)>6 nai Rijjih36x n 6i3rT'i3C3336nb biq33d33p‘b3 93nj96 b
3ha)9i63cnnb5i«6 dpn3h j,i6bb3i3363pn 3hmj,6mb3pn 9mg3P36n. General
Calculatuion 9mg3pb (GCMS) bnb^30icb3 gi 9 yih36x n j>i9moi3P36nb 9nb3g3noi,
2050 6pnboi3nb g30i 9 nBnb bi93ipm lt)3933hilj)3hi j)in‘bhg36i 2 ,5 °C-noi gi C 0 2-
nb t,m6(336lihii3ni ^9mbq]3hm3n 5imh9ij,036b [101]. 9mg3p36nb 393i53bm6i
53nR3363&b 3qjhm 9ipbpn 531*15^030300 5i53g36nb (un06 nhn, ii6ig^) 93^bg
5ioi6m6ib, 6iip3& (33pn0363&b ^hm3nj,369n gi bn99 hipnb 'bm63&nb 93hngni6 x
- 437 -
81306^0060 b660b7)^ 3^sn0^0n^90
Ia. 7
® O
J3 c
<Q 4
u
I 11
2
%
n> 10
E
A
m
o 9
300 I 600 900 pi co2 r'
Ksh 6.108 C0 2 -nb ^bhpnQn yn6u36k>h:>(jn35>nb ^136130361 Abutilon theophrasti-rh y^ n p m b ib i gi

6iymj)fn6i'b3. y3i3neo&nb hnubsn, 6iymq)nb 9 * o>3ljC36n- b:>3haim 6mr>8ibi
5ieii65ihn336xni 8oiX 91336^3^3 (Bazzaz et al., 1985). 8^6x5383^ bbji 8^361(13011 8361636
nb. Garhutt & Bazzaz (1984)
61b. 6.103
6ib3nh4ignb ac*’* -
ip3hn bnionb SmeSC1
(King, De Angelis &
Post cit. Jarvis (1989)
9nb3e3noi)
- 438 -
66 mf5 n 3n a 06 D(n0 b&rtabTô 36 d&rirt060
5i 3h(3XD^^- JicnQ^hn QmB^QQp&n 32)i39 ohg36i 9 hi3i 0 3jjm-q}obom0m5o3h 9 m-

930b, hiy 6ih9 m9 m6b 9^130363606 5i63noiih36nb bb3igibb3i bi536ihb. 6ib.
6.104 ibibi3b (j^ ^ h i^ h n b 2 °C-nai 5ibhgob 3ohm6369o 9(536.^3300 biqjihnb
3ri3h 6ib9 nh6 ignb qjojbiôob h35nm6i03h 5301103636b. 5i9moi3036ob bicg3d30ig
i(P36xn 9m6i(33936n ^ 3 0 1 3 3 5i6i3nhm63636 5:^(5303606 :>6;>0obob 3nhm6366:
5i 3hu303&nb 6ibp3h36nb ^^36(50311 io r ^ i^ h (ihi bnm^h) 3ohm636b 3 g i 9 yih 3-
6xn J)mh30i(3n3hn go;>5hi036o 2^(15 6 ih 9 mg536ib 53 oj96 oi6 9 i 0 o9 oôh3630 50m-
6i 03h i 0 n9 i^ 3h cgi;j,jnh3&b3 (6ib. 6 .105 ).
9(j35ih336nb h3iji50i ,539 33(1^ 3^ 6 9 m9i^ 36i'b 3, 6 n99 hion6 33(1011103606
jpbhgibi gi gi^)6mh3i'b3 (y330i<33hn 3b 369.1 9m3y336 j)0o9iôb (5300036366)
193000 5b6bb^3h3^n^: biqjb 30nb ^qjhm 9 i("i 0 n ^ 3933hb^3h3bn ,5369(10^ g i 6 m-
hs^hn ^ 3 3 6 1 ^ bm6369n, b igiy i 9 gi 9 :>g m3ljo9 :>03 hb3 0i6i0O (53033(1^3(1360:),
j)i9 mn6333b 6n30iii3h36i0ii (530nb^ g i bhgnb 3hm(53bnb 5id0D3h36ib. ibo9 o0 i(5ob
fcibhgng 36ihb, b36cnj3nb 06,1)3660 3m6o6 9m9i(i)36nb 5i9m, gi33ohob3ohg36i
C02-nb îh6n 5i0mymq}i. i 9 h35nm6369n giai6m&ib 9m3y336:> J33<336n0nb
0nj)hm630n g i 9 gnb 06^36603(116116 9m9i^3bi gi i 9 nb 93 g 35ig C 0 2-nb i^ 9 mb-
q}3hm9n 5i9mymgjnb 5ibhgi. bi93iQm 53015(1:103030 5i63g3b‘b3 3353,5:11500b 33(10-
mgob bi65hd0n3m6i 5iobhg36i; n9 939onb333i9n, 013 q)moim09133360 9^36^33-
6n 9 30 6 ^ 36 3 6 3 6 9303^1010306 6ig(j'3o6mg ^636300 ^homgob bi65hd0O3m&ib
(hig5i6 9 icnn q3mcnri0(5336i ^qjhm cgm^m33hnmgn'b9cni6 ihnb 0^390(136300,
3ngh3 (j3933hilj3hnb 9il*i36iboii6), 9ihig963i63 9(5361(13360 9 m53 &30n gihfi36oi6.
h6ri0n 'bi9 aiihn 5 >9 mn6333b lb i 9 cnhnb 9 mb33636ob 33homgob 53ni6 gig 5 m9 ib,
hiu 33b giii9iymqjn036b y0mh^36nb 6o(5n3obbg9o 9maibm36n03&ib 113500163690,
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