Professional Documents
Culture Documents
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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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ISBN 99940-61-67-4
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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 2Ssh^ospjhn 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)
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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
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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]
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BIOSPHEiRE
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eaS^hnbxn ihj: pH-nb baS^acm 9n6i(jo93&n 'b(«3nb 5yac3n (Ott, 1988). mhgn6alyib pjWba
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(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).
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20 -
40 -
Trifolium repens
60 -
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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 bjh^orm -
^hnp3n. J33g> b3hsom - 33KyiiiC3h ShmjJyijSn) (Kutschera, 1960; Kutschera & Lichtenegger, 1992)
- 25 -
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ARBUSCULAR ECTOMYCOHRHIZA
MYCORRHIZA
Gymnospermae
Vascular plants Angiospermae
and Bryophyta Basidiomyc&tes
Zygomycetes Ascomycetes
Zygomycetes
MYCORRHIZA
IN ORCHIDS
monotropoid type
Basidiomycetes Basidiomycetes
MYCORRHIZA IN ERICALES
Alnus incan a
C=) Betula pendula
[...I Quercus rubra
^ 3 Larix decidua
70 - 1 Pinus nigra
Pinus sylvestris
60 -
50 -
40 -
• 30 -
& 20-
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Snimhobnb (%) nfjg^b^nxn 9^33
moderate high extreme
Pollution load pa0»&n6d3lv)&CD&rib 5130361 (Kowalski, 1987)
- 26 -
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ihb36m6nbib 3dnd3 83050360b 93cn3nb36i n'bhg3&i gi 95b3o6d30o 8(5365113 33301
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Carbohydrates
Nudeotides
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 tW^oan
qQ6mcnfa)bn
9ih^n3n (336^360 b9 ohig qjncnpjbbi g i s6^o9n^»»i63c a ^oyyiiscrrrfrin,
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, >6^o9 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. i6^o6nm^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, 3C3CfT^iOiir)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)
- 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)
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
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
- 31 -
3(J365fn00to adfnOdD
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)
- 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,
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)
OH
- 35 -
90066630)6 &6 (f)09 n
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 om^i3b^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)
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
r 9"
OH
due
HO
GkioGal-O'
HO
H
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
Shm^e^n
39609363- 6606363-
3C\jhsnnbo93hn 290-360 0-4 9|jnh3 38608363pm
Cm 0013560
gm^bnbon^'b^hig
6608363- 6606363-
ij^ojho hipn^iyib 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
- 38 -
980b tfU>«SOaOlK> 3^09560
- 39 -
9(306660016 &6 ff>09P)
a b
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)
- 40 -
880U JC ? 08i &0
- 41 -
8 0 0 6 d(n 901 t) (W ft 3 8 n
0 20 40 $0 SO 100
Canopy cover [%] Canopy cover [%]
Relalive Iransmitlance
Iz = I0 *e-k LAI ( 1 .3 )
- 42 -
980 U (3 ^ 0 9 6 6 0
- 43 -
8036 ^ ^ 00 )^ to tfo an
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-bj^ihnb 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.
- 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
- 45 -
8036660016 &6 6 0 9 D
- 46 -
880b
a b
- 47 -
8 0 0 6 6 6 3 0 )6 566080
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 biiJ^ier'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
- 49 -
8 (3066630)6 &6 6 3 9 P)
1.2 .2 ic n a ^ n
- 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)
- 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^hR^enpn 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)
- 52 -
980b 66^06(306 *56 <3^09660
Precipitation
[mm]
- 53 -
30366^0016 <VMf>39n
- 54 -
980b (^061306 3CJ09660
a b
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)
- 55 -
8006663015 <V)tn38 n
Wind direction
y a/
1 / ✓
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
25
20
JL 15
cO)
'<&
I 10
£
20
IS
O 10
- 57 -
ai3066ff>30)5 &5 ff>aan
- 58 -
380l> 3^09660
&T=o.5 K
- 59 -
90066600)6 S 6 f609D
- 60 -
____________________________________________________________________________________________________ s a b a o f n & a m 0 3 ^ 6
Protocellijiar
organisrris
Phototrophic
organisms Eukaryotes
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)
- 61 -
GabBOfn&^Oli 8{ % 9M a 56 896^0 603Cn00fo0?»0b B^ff>8nd866
- 62 -
6ib . 2.2 g*9ning36xO ^>;> im^EStyi^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 ojnc^i^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
Rs6ihoi36b (3.6. 305bo,m6C<Ti63036b). bbo33ho 363(1505 jo9o3ho 66360b 363(15059
5 ih9 inj963&i 0)nCi i rnn63 ban.
<30io>mjo9o3ho 3hm^3bn n6y3&5 ;j0mhm305bo,36nb 9n3h qjih-ob 9 015601^8001.
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 93o^i3b 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-
ob bi3hcnm hiOig36m6ob 83035(19365 3058360, 700-0156 83hy3m6b 300 :1-956 600 :
1 -993 9(536ihob bib30i6obi 95 55(13601 3ohm636nb 0obx3ocn. bo6ioi0ob
<3010103680 9 3(35(19365 j0oihoiqjo0o a / fcmhoiqjoon b 95 PS I-ob 6000 3qjhm 35(?5-
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,
3ogh3 3b PS I-8oi; Jbi6o)m<3o036o 556b5330ih36x hm0b ibhx3^36 ^ih6o 363h5o-
ob 30,000^5(50580 (jbi6O)mqjo03ho 150300, ob. 01530 6.2. 1.1).
bo6ioi03 b5h3iji50Pi (5360^3680 06333b 5jO)03hn j0mhmqjo0ob 55^6365^ hbbsu
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
e^aacxn ^ “brian; PSII q)m,jmbnb,K)fo II, P680 e> 5oib^r>r)n6m6o6nl) ito^ijnhD&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^j^i’bi; CF ATP bn6cn3^i<bib ynqjij-
limhn (Hess, 1991)
8 hv
2H2CH-2NADP-H-2ADP+2P j------------- = 2NADPH2+ 2ATP+0 2 ( 2 . 1)
P 700+ P680
- 64 -
66 b30 rt66 m 03^6
- 65 -
GabaOrt&^Ob 9 n b 96 (f»0 b6 836^0 603 m03fnD60 l* ff6 ff>8 P)d966
- 66 -
66 banfn&5 ^ni> 0 3 ^ 5
- 67 -
66b90rt56m 3r)b36fn3{>6 <56 33^ 6 ^ 0 6030)0 3 6 3 &0b E 6f i3n d 366
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
qjmonvib *133659 656055363- jbmoyien 556055363- 8h33&*g 5*695536390
po 03'bmojn(vi, 558- 905 55845(10 im636nb 83lbmqj09n. bsSihifcm
45(10 3016360b fcmh36Jn839n jboi3opob 33(1393680
nh63Cn3 3*h36Jn8]- 83801813360 33(1393600 gngn 3*33019360
pn 83801801360 h5gn593h5g (56540180-
33(139360 nb " Kranz"-4o3o)
jpmhmgncn a/b . 3 . 4 . 3
CC>2-nb 30(1339*90 i J u ] 3- RuBP (b36b1jh51yi: HCO3-) 110650193^3: RuBP
(jmhn (b36b(j(u>yi: CO2) 1)06639380: PEP
- 68 -
66banff)66^nu < 3 3 ^ TWfnagreo
535Q3nbib; Sibbp 5530365b 5bg 3&b <3 oioi036 i 9 g 3 9 m(j<63 x n h50n5|jn5, 6nigi5nb
Qnjh 353(10956 C02-nb 9 a ii 6 cn;)05, <bhg5 9565011 ^3933(15^3!^ 3nhoi6369n, 9 oi] 69 n
^53hnb 956500 66335 g i oYb0163(10 b^obn.
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 isb^icniSn (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
Altitude [m]
- 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)
- 71 -
Gabaofnwem a n b a ^ a w ett a a c o ^ o enamoafta&ob ff^an da6;>
- 72 -
s a b a o fi& a m 0 3 ^ 7i?ccf>G^gn
Control
Time of day
- 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
Bib. 8.10 53ih Aeomum-dn CAM 3>ji 6mcn<b0nb gjmhajbn bjjxO^ijm&nb bihnbbot>6 iS39nh9 n
(Pilon-Smiths et al., 1991)
- 74 -
6abaofnM em (33g?a
- 75 -
G a b g o tfw m anbaartaw ^ aa^^o 603 m006060 b ^ff>anda6a
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 (goibqji^ig 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
50omjbo0i^i9, 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 (SnghoijbnSnl^i^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, ^ih6n 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.
- 77 -
B a b a o A M m 8nb86(f>oto aa^^o Goamoorssoiou p^Aanda66
2 .2 . b 3 b O \5 15 3 W a i5 D 6 ^ h O D 6 9 fl
- 78 -
(VttOto 03^ 91306^330
dm / dt = -D »A • (dC/dx) (2.5)
J=A C/ S r (2.6)
- 79 -
GibaOfni^^OU 9Db06(f>Oto 3 3 6 6 ^ 0 6030)03fo350b P6A 8D d8G 6
- 80 -
03^ 9(3066<n330
- 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>- cn^a^i bnb63033n
33h ^ 9 m hnbx bn3hu33o pi&hm33b30o b3&cnj3ncnn C 0 2 i 3O>ni30nbi Qi 33hnp3h-
03Qn jbm3O036nb g,:>3pncn goqj36gnh36b ^ih^amSn 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 libn^i^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
- 82 -
&68016 133^6 81)3666090
61b. 2.16 C3 gi C4 80361(1301} gmoicaiiSn C02-nb ^s^ihj&nb h3'bnbli 35>j x fT'^i (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 ^ijjhQ&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)
6bj,30i} J>}bb6}
6}j,30i} h}m036oi6}, 5^6^60036}, t>m8}, qjmh8} g i 9oidh}oi6} b}b30i6obai3ob
0}8}b}bo}oi3630o oi3ob3636n}, 013815} 3b 8^6336360360 (33}036}goi6b 3}60l>(} li)0b
- 83 -
66 b30 f n d ^ 0b 8Db86rnOM ^ 33 ^ 6 ^ 0 6 n3m00fo360 b g ^ 3 n d 866
Stomatal
width
[|im]
□ O’2
□ 2-4
□ 4-6
□ 6-8
- 84 -
<W8016 03^6 9U366fn0B0
91633636306, 6*53 0(3136*; fi*9 (,30) 33(1393600 ^h5m hnb 9*(530ob*b 6 *5 3 063(13-
6 *.
(53h5nhnb 5*90936* mb9mh353p*^iTih3co 3lim(53boi, hm030O(5 om6oii *5^13(10
^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.
- 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
o
o
w
d
04
in
in
o d
c6 06
rn
O
m
~
o
in in
aa^^o
in o g
OJ
*9J Ckj in
m oi
86 -
in
oJ — o SO
uS o
w in w oji
J Ol
(> > S>
9
w
Oi r-'
in Of
in
O* S>
in >o
>9 'O
o d d
s
in rA
- 87 -
e a b a n r t^ o u a n b a ^ a w a a ^ ^ o Boamoarno&ob pa<«i9 r)d86 d
- 88 -
cV)80D<!> 03 ^ 3(3 3 6 ^ 3 9 0
- 89 -
6ab30r t t o m 9nb&>(f>0iW> aa^^O 6030)03rt3M)b P ^ a n d 865
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\b3bx<i 5*6bb3*33b*6n *61*15
933cnb333i3n d*on*6 gngn* (ijbh. 2.6.)- nbn6n nd033O*6 8i{?*03ho>g3jon3cn
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-^itCTi (i3Ci 33hb3ilb (Populus) bt>3igibb3i y v i 63n O ioipnyijh 3nhm-
6^69n (Ceulemans et al., 1980)
- 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 biS^iQmg 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
3mprrOsh:>'bnl!)3&n 3-6 -
- 92 -
<W8015 0 3 ^ 6 9l33Gii(n0 3 n
- 93 -
6 6 b 3flff)W m 9 n b 86 A 066 896^0 G030103(f)060b ff-bff>8nd866
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
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 b^igniBn 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)
- 97 -
GabaOrt&^Ob 9nb9i)(n36i) <56 93rt6=?0 G030)03to060b B^9nd866
- 98 -
(W8016 03^ 9036^6030
Monltis Months
- 99 -
^0^0 WOto'DCnO
E
OJ
o
E
=1
o
o
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)
- 100 -
&68D16 03^6 8133666330
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
b iy i 8336bnsnm bo6ioiooai
atpSahoa):. 5*30511 &i6 >0 )363cyn6s ic &i53h3& i fs
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
- 102 -
(W8016 03^6 800666330
o
o
o
e
Trues
a
z Sciopnytes
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
- 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
(jbhncn 8.9 6i 6:>cn363QrTi6 :>bai56 93&33&X <3 'T'<n03^ 3 mhnb jp 6663:13365 (0h:>3;>Qn
i3^rTthnb 3nb3g3noi) (+ ^ojhm Sipipn, dpnjhn, 3 3^11; - jqjhm pa6s(vi, Irjbjn, 6sjoo&n; X sh ihnb
&s9m bsoXn shbj&noin bb3im6s)
- 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)
- 105 -
6^ 9066^^013 anbaartow ^ aa^^o 6030)0 0 6 0 ^ 6^6anda6;>
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) ^.h^ig 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 ish^ig 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 bnbjo^o
(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)
- 106 -
cW80 to 03 «=U 8 0 3 6 ^ 3 3 0
Time of day
- 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
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)
- 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)
- 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.
In Q j^ lO /C T rT ^ ln ^ /k !) (2 . 7 )
- 110 -
366666 L 6016360
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 SQ^i&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 ^ih6mjGngibi/mJbnj)36i'bnb 6iiih6mjbn0nh36ign
ijijn3m6nb giU30i9n. di0ni6 0i(p:>pn ^3033hi^3h3<bn i0unh3&36 6ib9nh6ignb 03-
^ibmpn'bQbi gi 6n3cnn3h3&iou> ^iBbimh^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^^Ob 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 ^ihgn-
6iQ3hn 3ijbn830n (bnjjbnb SiJbnS^Sn; bnybnb QnOn^n)
- 112 -
cW80 tt 03 ^ 903656030
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 ojo^o -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
- 113 -
G6ba 0 6 6 6 ^ 0b 9 nb 06(nD65 0 3 6 6 ^ 0 6030)00606015 ff668m 866
- 114 -
<V)8CT>5 03*^6 900666090
- 115 -
66baorfj66^ob anb 9 6 6 oaa a a ^ ^ o 6 O301OD6 D6ou P66and866
- 116 -
(W8016 133^ 803666330
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
- 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
%
10 0 - Helianthus annuus
CD
75 80 - ■31
0 _ Photo respiralion
</i 60 -
1/1 c
0 ) Dark respiration
M 2
a 61b. 2.44 &£nb eoojniyi^b 5130361
c
>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 bnsmb^Ub] (Boyer & Bowen, 1970;
Keck & Boyer, 1974 ; Ortiz-Lopez et
Water potential [MPa] al., 1991)
- 118 -
5680)6 03^6 900666330
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.
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
- 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
Time of day Time of day
122 -
<W8 (J)6 0 3 ^ 6 9 0 0 6 ^ 0 3 0
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^^o 6030100^05011 s ^ a n d a e ^
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
E 0.10 ■
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]
- 124 -
<W8(Jtt 03^6 803666330
- 125 -
66 ba 0 rt 56 ^ 0b 8nb9.iff.3to 89 rt ^ 0 G030)00 ft 060 u B 6 fo9 r)d866
- 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&v^in 3hmy3bn Mn Mg, C l, (Mo) Cl
- 127 -
6dbaort&d<?ob anb&ufti&d 9 3 r n ^ o eoamoa^a&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]
50
- 128 -
&68016 03^6 803666330
- 129 -
66b3flrt&<^ 0l> 3n b 85(*>3&6 ^ 83rt 6«=J0 6030)0063b0b P 6fn9n d 866
E
3
10 15 20 25 30 35 40 30 40
100
%
(fl
55
| 80
c
>*
l/>
I 60
I
1 40
5<D
cr
20
- 130 -
&68(J)6 0 3 ^ 6 8 (3 0 6 6 6 0 9 0
6069001 gi/i6
ph^6pn>6m6ncn
- 22 - 38 - 13 - 20 - 39
5i3ni633xn bn6iciQnb
eoqwiisn
gi&ipn 16 0bpipo
- 15 -3 - 7 - 12 - 8
M ^ahstfjhi
- 131 -
6 6 b a o rt& 6 m 8 nb&>6 0 to <$6 a a ^ ^ o 603moocnoa>ob P^^andae^
[MPa]
-10 c
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 )
- 132 -
90)^0560 90a 65<f)0b G6b30(nbi^nb ^ ^ G b O
- 133 -
66 b30 <f>aam anbaartOito aa^^o 603fl)ooff»o6 ou ^ a n d a e - i
- 134 -
80)^0660 80366<f.0U G6ba0(n66^nb
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^Sm^a39361 (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
- 135 -
eabaofn&^ob a m a ^ o ^ a a r n ^ o Goaainofia&ou E ^ a n d a e ^
- 136 -
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 , ^ijoiQb 338016333130 , 63pnb (Tranquillini,
1959). bb3igibb3i ibiyib 9hi33QO bob C02-nb 6p03hn 6i(?i6bnb 90161(539360 nb. Tselniker(1993)
- 137 -
G abaortw em 8Hb8 ^0 6 6 a a r t ^ O 603m00fo0&0b P 6ff)8n d 86i
PR = d W/dt (2 . 1 5 )
- 138 -
80)^0660 80866 <n0 b 6 ^ 30 ^ 66^01 66^66U0
- 139 -
6 ab 30 tf& «^ 0b 9 Db 8^ff)3 to 99 (n^ 0 603(J)00ff>0 &0b P 6fn9Hd 966
a>jbna>c^in 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)
- 140 -
80) ^ 0^60 8l33G6ft0b 6ab3n (n to m 66^ 66b0
- 141 -
G6b 30 fn 56«?0b anbaartO M G030103r t 3&0b P 6(f)9n d 866
- 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 Sij^oncn'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
143 -
anbaarnOto a a r t ^ o 6030)0D^ 9 &0b P 6fi 9 r)d96^
- 144 -
90)^0660 9 G0 G6 rt 0U BibaOfn&^OL 66^66li0
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)
- 145 -
6 ^ b a o ri& ^ m anb&>ri36 d a a ^ ^ o 603U)03 ft 3 &0b ? 6rtand 36 d
- 146 -
30) ^ 0^60 aUDG^fnOb 6 ^b30 A 6 ^«5 nb to«=U6bO
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
%
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
- 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
- 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; oi^anhn 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 ^idonihdn yn633&rvb
ghmb. (Gaumann, 1935). Abies vetchii (oi3m6m): 1 ai’biqjbx'^O 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 ^iging^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
2.4. ai536^h3Dcnon^6^(bn6^en3bD6nb
6^b9nhb^enb b rao w i
- 152 -
3 oa 6 ^^a~3 g;n m ^ b a s n a ^ n a & a & o b 6abao6&,^ob btroyaeo
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 nb3C3^i 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 qjoi^oiboB-
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. i8o^oi8 801300 9^361(13300 biqjihnb Shng^nnb CG R 2103630016b
(6ib. 2 .75 ).
iShn^ig, 3b ihnb qjmoioob cgihcnm6nb oi3^o8 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 ) gob^iS-
ljn3ho 5ibm03i b i33i036ib od033i j>iobm9 mb qjmcngnb qjihaioi6ob n6g3jbo 8153-
- 153 -
6 ab 3 0 6 & ^ 0li anb&HnObi ett 603m00(!>0 &0b P ^ a n d 8 6 6
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 -
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 .
1 - 2.5 (4b) -
- 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'^O^xiL) gmcnonb qphamb n6g]Jbn. nj, b igi^ Soihxn Sb^ihQ 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)
- 155 -
6 ab30 (ng»agm
6 nm33 n (Whittaker & Likens, ^ aar^o eoaomo^o&oij ff6<f>and86.b
ltj o m o in 0 o to o w O rO 04 p O
fO ol oi r<i ro O r-‘ ro d o 0 o oi -
«* K* 01 r- o «± tv
33a(}33ct»i6> bb3igibb3i
fO U*l p*. r—
ei Jcmhrngncnb
in O o ^ ^ t r - ^ p — qco O O _ 04
<D -O C> <i r - u*i d 0 -0 d <> — 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 . eoe^anenb 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 ^
6ib. 2.77 bbjigibbsb ^n3nb du^ihxJC0 biqjihnb 6jyn 3nh3x*en ihmg^Jjnnb i^30nhn 9(j36:>h33cm-
6nb eogjh^eynx n6g3jlxni6 ( i X 33n'J 9x 360^n e^ondBjbxni 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 ^an 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 Snhsx^e *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)
- 160 -
3006 ^ 0 7 )^ 0 roaeaiasngK^no&o&ob eab ao rt& am &0? )}060
PPb = PPn + ER (2 .2 0 )
- 161 -
6 ^ba 0 fn6 ^ ^ 0b 8 P)b8Art35 A 83 <f> ^ 0 6030)0 0 6 3 J>0b ff6rt8nd86i
PPn = A B + L + G ) ( 2 .21 )
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 )
- 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 bxJ^igmg 2jj$ &■%’ (0136^ biQbhjoin
ibnnb B3n0nb6 (jy^bSn, n^n S^ndp^b nymb 1,5 i&Q^a'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,
PF* -G*L> - 2
:r: ;G
B.6
S o il o rg a n ic m a ile r 35
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^ho^ob b3booi 80606:13383 93bob 93634385360; G 99hico
93bob 69038(1 93634385360 6013363^3
- 163 -
G^bBOfnii^Ob 8 P)b9^fnDM ett 3 3 6 . ^ 0 603010a(f.060 b P6(f>8nda66
- 164 -
aoa6a<na?reo m a e a b a sn a ^ rm o a o b GabaotiMqob aoaxaeo
- 165 -
66b30fn6^ 0b 8nb8ifn066 ^ 93(f>i^ 0 6030]03(f>D60b P6(f)8n d 366
- 166 -
8 0 0 6 6 6 0 ^ ^ 0 b ^ r t O b 8 0 0 6 363fn&00 b ek»n^6 d96 i
Ijbhnon ££1 363h&nnb 333u3xoi& i du36ihnb bb3igibb3i b] 6b^>6 (yii3 n (Paine, 1971; Lieth,
1975b)
pfejhj/tnb
339t533pmS>a
8313693131 8.9
6C<Hjn5n 8.7
Qipi^nb 0fp3^ 10.0
5nh]3i^nb 83131 13.2
fcCDioi'bs 15.5
imon5C3im6s^n 17.6
iy>C0£>n 23.0
pn{)6n6n 26.4
^60836(1 38.9
^>3)1336360 46.9
- 167 -
6 ab9 [)(fj6 a m anba-bffiow a a r ^ ^ o 603 moa<f>a?>ob e ^ a n d a G ^
Ebb. 2.62 353h{,nnb 333(j33cm6i (mhj(i 6x n 6ib 9 nh6ignb 3ihij,n) 6063^1316(1 (C), mhe3&6ni6n (D)
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)
- 168 -
9 (3D66 (f)0 ^ 0 U ^ r tO lj 3 0 0 6 063 ti&00 b &>ctra>d8 5 d
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 &:)
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
- 170 -
eo-b^&o 6 n a n 6 i) bW 33 &o 603 mo3 <f>3 &3 &ou
3 0f)63hX?Oh 6fl30in0h3£)icni
93013nb36i
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
- 171 -
306366^36 6030)0060560)6 300130b366
- 172 -
00636^ 7)60 6O30)O36O53J>Ob 9a0)30b056
Root hair
- 173 -
8 0 6 3 6 6 ^ 3 6 6030)036066016 3301301356
OH
OH .0
2 Fe3*
+ 2 Fe^ + 2 H*
HC = HC—COOH HC = HC—COOH
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
P lasm alem m a
120 mV
PD C atio o uoiport
PD Anion uniport
cotransport
Cation-
exchan ge
C yto so l PH
A p o p la st -pH 5 .5
- 175 -
80606^36 6030)0 3 6 3 5 d0 to 330 )30 L3 to
- 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
- 178 -
8 0 6 3 6 ^ 7 ) 6 0 6030)0060606011 300)30l>0&6
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 bnR^ih]
(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 ^s^ibx^ (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
- 180 -
803666330 806066 ^ 7)6 6030)0060560)6 30 )660)d86 ^ 6 ^686606056
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
^im6ob 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
- 181 -
a06 0 fn^c5 ’3 fn 60 d0)03 (n32>60)6 30 U)30baiW>
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
- 182 -
800666390 806366^:36 60a0)0363;>60)6 90)660)d96 <S6 0569663&366
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8 ■a
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J E c E &
82 f6F ID
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bb3igibb3> ,i)nJ9 n (Rodin & Bazilevich, 1967)
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£. c c c c c c
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- 184 -
aooB^fnoan 8060 ^ ^ 3 ^ 603ai0 Drtaw(jto aowGmdaa ^ a a fo a a o w
- 185 -
Ijbhnon 3.4 3 ii)hmx33 D5 ^36nb 339 i}33Qm6i, 330130036^, &i3h(j3C3&:>, ihb3&m6nb qjmh3i ps 1336^01 (dh^sQn i^mhnb am6!(533n Finck, 1969
3nb3e3noi)
906066^7)6 6030)0060660)6 330)30U356
186 -
onnbnb 9n63h:>e3&n iebmh&nhjbxn > 3bb- K.+ &sbb6ncn 3Sh3pnb 633630 Snehi^unnb h^- a^hnb^anb sbscftith- i^&n
6ien XSHJ'1*. DCDlOjhm- e=> Jbm3nc*i. j3hjnb
jn 0n3-hn n 3 bh36jn 0i
s
SB
>,ih6m6i536n 3bb6ign > Mg2* {,ibb6ncn gi :>gbmh6nh;iaxn'Snghi^iiyinb h^j,- qjmonmen aQgih^&ncn
(gncman^n), igbpihbnh^xn: 9ynhoi mhii6x ^9 &9x n iihj,n
bncnii^n, 9fP33 6nigij,;)&9 n jcmhmqjn09n gi q)rr\jmbn6cn;)'bn
bxq^OA jQmhngn jSbnQ^Jinbi gi hnbmbm9nb
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bxajnp^&n, bx- spbrnhftnhjiix'’ ; 9e*n- Cu2+, Cir imaSpjjbjban 6630" (3o>b- ao^m en'ban. b ^ ^ D ^ X 0 (?0h|T| <*630°
iih&rn6il53-CrfbO&X n > bitor^hn ^m(}ni5nBn). 36'bn3;)&nb jma-i'bm^nb a^bm-
5„ 3m 636^n (jjn iyn jh m SR jb rv pn'bSn, B ^m hsen
Qibs, q)36mQmjbnei'bi) 33^>bmQn'b3n
amendgi^n igbmh&nhj&oen; ambrv M o042‘ a^C ^d^X S^hcnjdan; itrn^nb qjnjbi- qjmoimcn, P 3hm d6ocn
pn'bo&xn > 3gi3^ 36'bn936nb jm03m636l5n yni, hyi6nb
(6nlih ili)h3g3j^i'bi, 6(1^ 0153- sdbmhiujns pi
6:*:,) e,iei6iij3C06i
- 189 -
303666090 3 0 6 0 6 6 ^ 3 6 60301006056016 3 [J166Qld36 «?6 C563663 &3&6
9 0 6 0 6 6 ^ 3 6 6030)0063660)6 300)301366
- 190 -
8 0 6 3 6 6 ^ 3 6 6030 )0360660)6 0 ^ 0 8 0 6 6 0 0 6
bnibpm33b (bbgbij gngn hbmg36m6ocnbb Zn, Pb, Cr, Ni, Co, Cu gb bb3-).
0(536bh3Oib ijRvji bbb^mibb 33533630(1 bb3Cn 3ohm636b. 9 3533611b 3b 03^60^63-
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
calaminare, Silene-b ‘bmftr^hcno bbb3m6b (^003063(10 0(536:1(13360); b3h336oo6 x
6obgb536'b3 (Cr, Ni gb Mg-ob gogo hbmg36m6ob 930(53300 6obgb53dn) -
Asplenium cuneifolium, Alyssum bertolonii, Armeria maritima, Cupressus sargentii gb
bb3. 5b6bbi)3cnh36ocn gb9bbbbnbOi3630Ob b3o036dooib gb im6b0oooi 9 gngbhn
6obgb53&ob qjgmhb (jjb0i65obb gb hmg3'boob "b3o036dob oj0mhbn); jQShmqjoo^b
b5h3cn33 0n3j)3oi3636ob6 Silene vulgaris, G ypsophila patrini, G ladiolus- ob
bb3bgbbb3b bbb3m6b, <bm503hcno 0 bh(530ri3b6 o gb bb3bo.
3 .4 an 6 D h X 3h 6 n 3 0 in 3 h D 6 ^ D c n a ^ u rD
- 191 -
9 0 6 0 (W 0 fn 6030)00^06^0)0 300)301366
Incorporation Elimination
r
Storage Catabolism
■*• Detritus----
De Intermediary — ► Excretions —
metabolism
3.5 y b m n b aaiWbmcrvban
- 192 -
68 TO 0U 9 0 6 6 6 0 ^ 0 8 3 0
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 ^ )
^36X 1^9 ^ ibm^nb 1330001. idnomd s<bm1jrT>3!6 3 3 3 6 ^ ! g i 3hmg3jl!)n30f,’ 6 !b 9 m“
hnb ihbgbmib 9 ^nghm imhxMyiohn gi8mi)Og363036!. 3b 3636! hm&mhu |jiC i3X
3(536ih3b, ib333 g3gi8o6ob 3 u36 ih 3X biqjihb 3 oigni6 ig. 3(536ih3Xn 8ibnb 6 >
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.
363i63 3jj36ih3 093636b ihimh5 i 6x ^>6 683c i'bmiijb; nj,n ihi Sbmpmg 6ib8nh-
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 ^igibi^ibigi^ 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-
gni6 i'bm^nb g3qjn(5nob.
3.5.2 y b m n b ^fianc^rD
ibm^n ihnb mhj)i6mj)363hn 60^30383600, hiy 081b 6n86i3b, hm3 050 8(53-
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'bm^ob 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 Scru^ihni 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 ibm^ob (530180 050 ib h x 36 b 03i!>!0 86n83630Pi3i6
hmgb. Bnohivjhx^ii^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^-
& xn i b!b3fnbi'b3 gi t,m6j)h3lj30n Sibn^i^nb oi!3ob3b3h36i,b3. NR-nb 3 ijbn 3 i 03hn
!jon3fn6i i(pnhn(sb3b! O3336n03h qjibiSn, bm0m gpn3h gn6i8nj>!8n - giib-
0m3booi 83ig(nnb 838g3&.
- 193 -
9060 (^6 ^ 71(6 6030 )03 (*»3 &60)6 330)301366
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& n^osao
Acclimation
temperature
o 4°C
A 10°C
□ 20°C
• 303C
■ 36°C
- 195 -
806066^7)6 6030)0060660)6 300)30U0&6
196 -
;> 8 n m 83666 n ^ 08 an
Xanthium Pisum M a lu s
t ft
NO j
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 ^
- 197 -
906 a r n ^ 3 (iS 6(13(J100rt 0 &.MJto 93 m3 nbOW
anena^an / 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
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
- 200 -
aOGOfoi^Tltf 6030)0063660)6 03^01 0)630b36'3606660 bb36^6bb36 366066630
- 201 -
806066^7)6 6030)0063660)6 330)301356
- 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 gmhSnt^i^, 806311103(10 Ca and Mg
6n3om3h3636nl> 838(533crndibi pi n ix x x z c x
K
9u36ih30u>cn3nb 8o63pm8ip(’i6i'b3 pi,
br^ipip, 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-
- 203 -
806066^7)6 6030)0060560)6 800)301)056
3 .6 .2 . m sn in cD h n qs wconqjDbDhn ai536^hD36n
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)
- 204 -
3063(^^7)6 6030)0060660)6 ^ O b 016301)367)605660 bb36«?6bb36 366066630
Improved Fe Iranslocafon
to the shoot
- 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 an^ijyicn
dm'fcohgil SSCU(iqj35^in 8(336^ 33^ Sm'bihgn JXH5ni rr’C3hn
- 206 -
806066=57)6 6030)006066016 03^01 0)6301067)606660 Ub36«?6b b 36 366066630
Silene Anthyllis
1 .6 -
c
2
- 207 -
0O6Orf^;3<*> 6030)00(60660)6 300)30l>066
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)
- 208 -
8060A 6c;t)(f) 6030)00(^06^0)^ 03«=?0b 01i 30b06U(f)06660 bb3^ b b 3a 366066630
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^
- 210 -
9063(6 6 ^ ( 6 6030)00(60560)6 (603080 30366(637) ^ 1636(630
kM = (ML + M r) / M abs. (3 .3 )
- 211 -
Ubhrxyi 3.12 3n6ohiC3h 5n3oin3h]6ioii (y33Ci 3n6i|339n j^io^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> SCO^O^^&biu)
S>
u
toj
o
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r^i
r4
CN
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(O lO
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l£c> n
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£ £ <1
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On
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-£
iO +
9063 ^ 6 ^ 3 6 603010363660)6 330) 30b 356
? 11
6 <1
o £
B 2
CD /O
T*
212 -
S
©
VO
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
.1\\ , ;i
Vv;V \ y :'M '
6sb. 3.27 &x&nnb
(djbi, (p)3b3n, Bnqjxfi) 0n6yi- Leaching
sQ^n bnjrTfijnybxra ^njjjvib 0me- from crown
X<i (Duvigneaud & Denaeyer-De
Smet, 1970). ^spm&rvjhn 9mp3Qn
'bmSnjhn bsh,jypnb 6n63fT>3>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
(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)
- 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
- 215 -
B’d^Ob 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
61b. 4.1 0^363(13013 Bypnb hjgnSnb 33Pip3(jn3 Siam bd^Q301^3 5sgsb3(<sboi36 psi339nh3&noi
(Walter, 1967)
- 216 -
81)366601 7)J(A3 ^ 0b 603000
- 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) bof^oD^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)
- 218 -
3 0 0 6 6 6 0 b 3 X6 3 ^ 01) 63 3 0 3 0
- 219 -
p a ^ o b rtoaoao
’ F 3Sh3en = ( - ) T K + ( + ) T P ( 4 .2 )
a b
- 220 -
9 0 3 6 6 6 0 b T jJCA O em jj^O b 6 33090
e = A P / ( A V / V ) [MPa] (4 .4 )
- 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 ^ih^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)
- 222 -
813DG660 U f f a e m 603080
- 223 -
(*>00090
6y^cn 6nsgsj,Bn
Byipn 6oigi&8n b3g36i 6soDjo6nb biS^ic 36001 e* M T ^ b fth ^ n b 6 yoD^ 9 -
93. iih^n 5i0^)hm6nb 6 nigij, 9 n Byonb 33Cp633nb bnfijih3 63po6ig9o hi3g36n03
03^hni, oiob6ih 6nigi2,8o - 1-2 0-Oi, doo3h 3j,3hn3 6oigi2>9o - 0bmQrog hi9g36rv
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, ^imbn.
bi65hdon3n B3O0nb 33hnmg36 nbi gi ^ ib iq jb jp ^ oim3pnb g6m6nb 830936 ByiQn
6hm3g36i 6oigi£,nb <b 3g i 03363680 g i iOo^mO ig30Qi9 6i3moy3636i 0 1336 ^ 30 ^
0n3h (bibih&36poi Ridnh3oo Byion). hiy 3qjhm gngni 6nigi2,nb 0o3h Byonb Qi£,-
hm33&nb 36ihn, 0ooi 3qjhm b3obiyh3QOi 0|j36ih33&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 836m^ooo 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 3nghmb^i)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 ^inQ 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
- 225 -
b60)67)(f,n
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^B^oiOob 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 boigi^ob 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 5^ih36nb Shoi^bn nh(ji333i.
Bnigij,9 n 6yi0n 93ndo3&i o9ym<336mg36 oihoijgnb bibnoii^. (^63 6nigi5nb
- 226 -
90366 (n0l> Pa^OU 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, ib^om
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
- 228 -
8006660 b e a ^ o b ^oaooo
* * = ^6<i}ei5n + + ^ bnigi^n Jl • rl ^
- 229 -
F a ^ n u rtoiioan
— U -7 < n
Dry air \|fa » - 100MPa
Va= ap----- h / - 1-°
^stem
M b -
Humid air -10 MPa
(93% RH)
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)
- 230 -
30366 ff.ni> p a^n u rooaoan
6ib . 4.11 cnid6ijm3n cgncnQnb 6ycnb (lF]eaj-) g i pjhmb 6ycnb (’P stcm)
33h^,ny>03hn jhignQb^n. hmiy> ^hi6b3nhiijnnb n6^36bn3mdi a^iem, e^hmb 5ib63hn3 diphg^bi
dpn^hn 3nghmgn6i9nj3hn jh ig n ^ n ; hn^i gnoncabn oni3bg3&i SihyW n g i imhcnjcjfei
3nphmpn6i8ni>xn 6hign]6jnb 6^336363(30 njis^&b bha^^c™ Smtnxyiib 6nbgij,nb (T^ij) gi
ajmcnpob ('P]eaf) 3mhnb (Begg & Turner, 1970)
MPa -2.5
( - 1 .2 ) -1.2
(-0.9) -0.9
-0.38
(-0.B)
(-0-7)
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)
jH20 = v . A [ i 6 69 ' 1 ] (4 .8 )
qjmoiQnb 56 pjhrnb j>i9 ^iho 'bjgiinhn ihnb jbnco9 nb y330i 5i9^ihn 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 ijm^ii nb3cnn 00n3hn (15(^0360, hm903&bii3 933d0nioi
5y0nb bhxymcgnpig j,i^ih36i (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 ftiS^ihn <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
- 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
h3<b 3h33&b.
4.9 qjmh030nb oii6 ib 9 ig, Bypnb ij(33iC3hn 6ijpgnb bnfijihp (v) n9ymq)3<!>i
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 ):
(jbhnpfl 4.2 0i536ih3cni 6b3ig ib 63i ^n39n gi mhj>i6m9n jbnp30nb 3nghi3Qni3hn &i9^i(v)6shni6m6;>
(b926d"' 03i_1) (Berger, 1931;Huber, 1956; Zimmermann & Brown, 1974;Raven, 1977; Cichan. 1986:
Ogino et al., 1986; Losch, 1990)
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 pa^OU 603090
6sb. 4.15 Abies balsamea-Sa Bypnl) 3ih(jb;)6:> b^hsoin: jbnQjSnb b^unqjnj^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)
- 235 -
633090
Laurus - 4
-3
- 2
E,
>»
Ia> - 1
0
i Olea
I>.
X
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).
^iiQfO^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
- 237 -
sycm rioaoao
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) < > -
61b. 4.20 Stbnbse&i 9n0shoixn ojmoipnb ^ebSnhi^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(j^eTj)m6nb ^Sn^hn biS^iem 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 ffa^Ob 633090
E
0CM
1
o
E
c
o
CL
to
- 239 -
e a ^ n u rtaanao
^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 ^opob36h6o
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)
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
^ ^ ~ I r&i63 + ^ ^ (4 .11)
- 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)
- 242 -
9(30666013 ffd^Ob 603090
- 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 fii^ih3630i.
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 ^ibbnpm&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 6^e^t>333^ Tr/Ep 6hqjn3n cm6:>q)ihgm6npi6 90301001366 ,jh:>6b3nhn5nnb 3hm-
U3bnb 6ij)nb9n3h
- 244 -
8 (3D66 fn0b e a ^ o u Aodoao
- 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)
- 246 -
01306660b B ^ O L 603090
- 247 -
Edcm rtDdoao
- 248 -
3(306i>(f)0b P a^O U 6 3 3 0 3 0
- 249 -
pa^oi rtoaoao
a) b)
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 ^i39nh36ncn (Zhang & Davis, 1990; Henson & Turner, 1991)
- 250 -
90 D6 ifn 0U ffiaaoan
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
H am m ada s c o p o r i a
— Water potential
— Otmolic potential
<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 qjih^os^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 )
- 253 -
pa«=job fnoaoao
b3 0 (j35sh335n
©mthgnon 6336a ai53nbn bn3hi530n 55630015(136300 505oihoij03630n ^395-
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 i^ujni), hoi0036805 6y0nb 6i0i6bn
'601533(1 i(T^33b dpn3h 3ihyoicgnoi 6 n96x b -
- 254 -
8096 d6 0 b Eu^ot. 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
- 255 -
ffa c m rtoaoao
63'bmq}nvj3ho 6ioh6i0ib36o g i
330ng3hn h35om63&nb 6i0ibm3i6 6i336ih3363n 63b3g36i 6y0nb h3gn6nb
ainoiJ6nb y3305 J>n3n g i 3ihni(5ni: 3oghfnb^i6 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 3i6n^i(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
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)
- 257 -
f f a s m Aoaoao
r»(3 dpn^h ih Sh^j&i, 3 ioi3 o 93 bid03&30Oi 6 ih0 mnJ95 ib Bygnb 3tfoh3 9303013(1^(1
(20%-3q3), hi(3 06333b cgmoi03i>nb ^Jj6m6ib (6ib. 4 .40 ). hm^mh^ fii6b, 5i9^ihn
bnb^)30 i 33h 33j>0 i 3936 i Bypnb 5i(jili3&nb ib3ai 3'bih3 ilbih 06^366031116:^.
Sndhmajn^^bb, hm003doi3 0^9360^6 ^ 33^380 g i ,i)36ni6, Rhgopoi6 3i 6 o,ji-
^3680 “bn6icn0ob 03J>3&ob“ (^ 3 8 0 3i5nh3 bmSob 5i6iOi 36300 i05o036o, hn3p3-
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^b^o 5i63noiih36ob 5i3m, Bypnoi bi^Sihnbig 3<bh36330ymq3i ih
bg3&i. iSn^mS dpn3ho imhoij036nb 3nhm6369n nbo6n j^n^unnbigSo ihoi6 3 og-
h3j>nc6o (5i6bii3oih 36001 I33i^036b-0b g^ihnb 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(^i6 o03hn ^o3nbi36 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-
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- 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-
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6y0ob h3jo3b 33C'4h 43016^10103636: 6049450 543oi33li40o4, ljh46b3nli4Uonb 3hoi-
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@ib3330b sgoijn (jhs6b3nhbons'b3 jjshm
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gn^ahi3nb j33gs dbihjb 648336360:1
gmcncnb gsfc>33311b bb34g4bb34 Smfenijni
(O’Toole et al., 1979; Hsiao et al., 1984) V 0
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pa^n b (noaoao
3gi66rob 0jjQ6ah336n
9l536ih30ii bibn^nijbcm cgmhdgbnb 61153500^0136365 &56bij)30ih36ncn 90905
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63^360 , hm03pcn5i5 555660501 b^jjXD^iSDhn 95 5bn0 D05(jnnb 36ihnb 9 jm63 j^hjob
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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
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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
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b ^ s n b &i0m) ih 03n6n9636i (6ib. 43). 6i&336nb 90033^3(10 5it^ihni6m 6nb 9 i-
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'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
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b b . 4.43 lutuhnb
3(jj 6ih ;)o ii iih i5b 3nh i(jn i 98hi(>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
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0 ...........I1" ....... 1 q ay* ihnb h m 'b jjx n bibn(jmyb(vn
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Time of day [h] 3nhm&3&3n (Stocker, 1974b)
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9 i336^ o u fnoaoan
libhnfxi 4.5 bb35p5bb35 ^impm^n^hn ja^ignb 9^36ihnb ibnQnciyn^h mh5j6mcni 0n6n9john Byonb
31^ 536(30100 (MPa) (Richter, 1976; Doley, 1981; Nobel, 1988)
o^o>d-Aim 39566m
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Ceraloides lanata (C3) 154 2.2 1.4
y>ij3bnb 39566m
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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^ By^ob (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)
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80365 ^ 07)^0 m .b e ^ s n a ^ r m o & o u p a c m rtoaoan
hoanan
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rioaoao
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
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bob 3m 3i^36ib (Schnock, 1971). ghi>ftn£Cn 63060160 iiy336obiort3nb nb. Brunig(1987)
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9036 * 6 3 ^ 0 cn ^ u sn a^ rm o & o u A 33090
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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)
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ooij0nb boi8o3ho bih^yQnb 65(159863563 i!>9336'63, 1101839015 601(18 606336b
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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.
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(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;
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U992)
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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.
- 277 -
a^fnoanb ^ 3 *^ 06 ^ auoG^fnOb 8 foe;6-&b6300 tt(o3 to 83
- 278 -
I
- 279 -
a^rtoanb a^ 3 ^ 3 G^ 9 ua 6 ir t 0 b 8(f^d-&> 63 om,Xna&<>8 a
- 280 -
8rt«;oM a^ 63om^fnaini>
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
- 281 -
&*6 0 9 [% &*3 C?36 * 91)06 * 6 % 8 6 e;*-&*63 na)*63& *83
a b
- 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-
„ 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
I IP T T P
\ ; u
- 284 -
8G36^ o b w u o o r m ^ n &d6300 tt 6 0 &0ii ^ 583i 0
- 285 -
a^A 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
Growing W inter
Deriod d o rm a n c y
r f (\ l Y
®
D e sc h a m p sia c e s p ito s a
4 I
- 286 -
3 i)06 *(n0i> u ^ n u n o b ^ n &*63 om*6 a&ob 3*8350
- 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*.
- 288 -
aooG ^ob w u o u n u b ^ n a^Baom^fnaanu ^aa& o
a b
- 289 -
a*tf>aanb 8 0 3 6 6 6 0 b 8(f)t?*-a*630 m*(fi35*80
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)
- 290 -
3l)36^ 0b l ^ u o o n u b ^ n &>6300to A 350b 968360
$>
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)
- 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 bj^a^^hn {>i9mbib3C36i
(Cleland, 1986)
- 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)
- 293 -
aartoanb cv>3 ^ 0 5 6 aoo66fssou
- 294
8U0 6 6 6 0 U l^ u o o n o b ^ n &6630 m6 6 0 5 0 b 368060
- 295 -
8 l3366fn0 l> 8( ^ 5-&6630 m6 ffiaM 8 a
'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
9i3ifijsh3bxo 3^6x3^
9bnb h a ^ n y ii
<3bh-nt> ejiynyxyi
jhdx° ojr'yy’ iS^inmgn 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
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)
- 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^ j^ifjh 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 .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
- 299 -
a ^ o a n b &63 c?aG6 8 Q066 fn0 b 8 6 t? 6-&663 nai6 ff>05680
- 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
Bib. 5.16 Bpnb 5>i69b3(xn6b3n pjhmj&nbs g i <3 Db336nb 'bhgi uofj^hic^hn ^hmSnb bbsigibb^
b] 9(536sh33n (Hoffmann, 1972)
- 301 -
{Wrtoanb <y>3 ^ 36 .> aiiae^fnou 3 rto^-&563 om6ri 3 to 83
- 302 -
86«?0b6 a66300)66060b b08063(f>0 908^06660066
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)
- 303 -
(WfoOanb <V>3 c ?366 9 (3066601) 8 6 ^ 6-a 663001663 &68 a
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
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)
- 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)
- 306 -
86^016 ^6 &66300)66060b 10806360 909^06660066
Predor
mancy True dormancy Imposed dormancy
K /
Correlative Exogenous
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)
- 307 -
(Wfoaanb 663*^066 90 3 6 6 6 0 b stfe ^ a ^ o o to fiia to so
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)
- 308 -
8 fn m a ^ a^ 63 nm^(na60 b uosnsDtnO 809 ^ 06 atf 0 m.b
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
- 309 -
cW6 3 8 C% a^3^ 36^ 90366601 8 6 c?6-&6630a)6635683
Meadow Forest
Beflis peremts Carpinus betulus
Primula elatior
Anemone nemorosa
jf||
a I
:t T :T :i t - rn r
F M A^M J ' J A'S 'O N
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
- 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
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 qjoi^oOohnoign (Roller, 1963)
- 312 -
8( ^ 0b5 & i 6300)6rt050b b08C%D<f>0 809^06560066
Climatic factors
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)
- 314 -
86m a ^ &d6300)660&0L 1)08067)60 808^ 060630^
- 315 -
aoosafnob 8 rf>g;a-ga630 Qi;>fo3 ii>a83
- 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 biS^iOfn 6pn;)hn d^ijhs^^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
6 .1 .1 h i b 6 f l0 6 ^ 3 b U b h O b fl
- 319 -
9 006 6630 60 ISfoObTJC? 9 CJ& 0 9 6 6 0 0 6 6 3 0
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.
- 320 -
b&fssouo: ^ 0 6 ^ 3 0 3 0 do b06 ^A nan
- 321 -
9 (3 3 6 ^ 3 0 6 0 LGrtObTF? 9 ^ a n 96fn3 n 5630
Time [ d ]
fob. 43 <3n^™i63n6n(jmb c^'bpdn fShgncnb sd&>& Oxyria digina-dn 9nbn d(xi3h bnE^oico^
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 (iC^O 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 ‘bm^ign,
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.
- 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 Smcn^iSnpijdnbs 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'bx^i 0ii3db3&3h3d:>6n. (C) "jm^hmen; (Pb) 200ppmPb-ncn 5-gcni6n ^>30^1^3393-
6ngi6 10 ecrnb 338^35 (Maier, 1979)
- 324 -
b&fnObO: b06«?rtn30
- 325 -
9006660050 g^anaaftan M ao
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 bOB^OSt)
20 30
Temperature [°C]
- 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
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
- 328 -
U663U0: ^66^3036 UOG^fnOaO
(jbhnpi 6.1 3u j6ih30tt S^iibmp^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
6 .1 .4 .4 6 ^ h R 0 6 i b ib h O b X 3nhP1606an
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 )
- 329 -
8006660360 bftrtoiyac? a^anaoAonboao
6 .1 .4 .5 b ib h D b fl Q) D 3 H C 3 l3 n i
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
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 3hcnn3hcn9i0rTiing36xD^i (Kreeb, 1974a; Levitt,
1980a,b)
- 331 -
9 0 0 6 ^ 0 0 6 0 bfirnObac? a ^ a n a ^ a m a a o
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> cnQfHjnh^i b^h^bx^
ojiiynhjdncn (Bell & Bliss, 1980)
332
{>'36066030 UftrtOb^O 3 i>jJ6 P)6 060
0.2
a
0 I- 1 r ■----- r ---- 1----- *----- 1----
0 60 120 180 240 300 360 420
ia.
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 (<I>a) 9mhnb {Jujfjihjou, Vj9mtn9my3i6nc mh bib3m&}9n
(Demining & Bjorkman, 1987)
Ulva Gracilaria
a e
ii
CL
- 333 -
3006660060 u s t f O b ^ a«;an 0 6 6 o n 66 ao
- 334 -
6sbi 6.13 (yipijjmnQob S3di)ha6j9n JbiEi^mojncnb ^yi^yib {>s9sfHy>336](yi b£)9s. JcmhmScib^nb
j^ioij&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
Ebb. 6.14 3nmC4jbi6tjn6nb (VX) {>ihg>j96i i6crohiJbi6rtn6ip (AX) pi 1b3iJbi5ijn6ie (ZX) CUorella
pyrenoidosa-Sn Qi^icn &*5ioi36nb Snhm&^Mn (Hager, 1967)
- 335 -
8006660360 u s f n O i ^ a^sp m fn o m aao
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 ^iQ 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 mjbngiS^o-
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 b^i^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
2.0
1.8
1.6
1.4
1.2
1.0
- 337 -
0036660360 b6 6 0 L 3 ^ 3 s?&n8 6 6 3 n 6630
- 338 -
Ubhncn 6.2 :jfnnyicm^°Qh3Cn 9o:)6:>h336nb Q
* 0nyiromhjp6rfb83&nb aijbn3^C3hr» iW^D^iWhZJC0 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 bS^ObTI^O 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
^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? 9^ar)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
- 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
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 Ii6n^i,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’bX^O 9b<^»e
UiCi33Cn 0^1x0(136300 35(139360(^1 013
5191(16361 (Larcher & Wagner, 1983)
- 343 -
8006AAD360 U&rnObT)^ 8^50966306630
,•16 660
6bb. 6.20 (jaQSjh^hnb 'bhpi pi ^jQSjhi^^hnb fchipn^B^ bi3i6i9ri bi6dhnb ghmb (Vareschi,
1962)
- 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)
- 345 -
3036663060 8^ & n 8660 n 6630
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 ^ihdmjbnQibi 49 56
PEP ishbrrjbnci'bi 46 54
3-PGA yi6ib i 51 51
ip ^ n c ^ ^n6ib i 47 49
cgmtxrfm33;)brn<b n'bp^hi'bi 52 55
Ru5P ,yi6:>‘b:> 44 52
6 .2 .2 .2 b fllib O
- 346 -
67)6066030 U66flli'as=:0 36d6r)6060
- 347 -
8006669950 I 6 fn0 b'3c5 8 eJ&0 9 6 fn90&690
Time of day
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)
- 348 -
6-36066030 b&6 0 bT)c?0 36:1606060
3 49 -
9036663050 UStfOUTJ^ 3 ^ 2 0 3 6 6 3 0 6 6 3 0
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
■ '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 pn6i3n^i (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 b^O g 6 dSH(fi050
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)3^ot,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 hlo ro plast s t r u c t u r e
Plasmolysis
Vital staining
0 60 120 180mm
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
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 qjfvn^aiSn
6n3<nn3h36icni ^i&jSmh^nb bnfijih3 (Giaquinta & Geiger, 1973)
- 353 -
90066^0060
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)
- 354 -
57)6356030 1>&6 3 U^ 0 96d6 f )6 0 5 0
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)
- 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.
- 356 -
5360&6030 b 6 (f> a u ^ 0 ^ddnftOftO
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
- 357 -
0036663360 8^80866306630
- 358 -
67)6066030 US60b7)E50 36d6D6060
9
©
2
course
(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-^aa&o usrtauac? a^an aartam aao
® 40
■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
- 360 -
67)606(f )030 ^ d 6 D ( f .a b O
Months
Months
Bsb. 639a,b 93hj6ns6 9u]5 ih30u> yn63i^iddp3fn6nb 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
- 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)
- 363 -
8006660060 bS 6 0 bc)cN 8 ^ a n 8 6 6 0 n 5690
- 364 -
57)6056030 l»6 6 3 b 7ic ?0 3 6 d6 P)6050
- 365 -
8006660060 a e ja n a ^ a m a a o
-4-6
-6 - 8
-5 -8
6sb. 6.45 9]b5ihn ^ynb yn63ij,i9dp3m6i. Rihfirn9n 9mou>3b36x<i bjhicn^bnb (jnqjh^dn h(p6n36:>3b
n8 ^393^ ^ ^ 366, hm930cni ^333001 ps'bni63&36n 31,33 {>s0mnb:nj3&:> (Larcher & Mair,
1969)
- 366 -
67J636 (n030 g^d 6 Hff)360
- 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
(CD^ahn iKJ83ahieohi) ° C
- 368 -
67)6066030 liS fn O U S ^ O “J t t d & n r t S b O
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
id0n3h3636 8(5365(106 b ^ h ^ b x 8 9 50 185(130 165^ g53^ 3o h o i5336 o b 3b 3(101016900165
“bidcnhnb 30(10163680 0 6 333b 3 ij3 6 ih n b g 5“b o 56 36 56 , hnb 6560655089350195(5 35b
555^605 bb3^Q>bb3i 958(53390 33;j;>6 o'6 3 n (nb. 656. 6 .3 5 ). 60(50306 3301(1590 g y j-
($01113606 6 1 3 0 0 ^ 8(5365(106 bibnum ybpm 0336^(5036^3, 33 0 (133 0 36 yot30o65, 55801-
n b i(j36i 3(10193^030016065 g5 “bhgnb 33(1001906 838(50(136530, 556b 5j)30i(i36nan
bi66hdpn3n 01013906 65035(106 8 3 6 0 16 33 35 8 0 .
yo63nb 6o33h503
(1019365(5 6059550 55yo6x n^>. 6(5365(13360 3305(1 0 ^ 3 ^ 0 10 3 0 0 3 6 3 6 (956500
bmhCTijp36nb 30)10163680(5) 6ygo6 3010 160 1360 0 3656. 6 0 59 550 6 5 ^ 0 6 3 ^ 650101306
6086536 60595506 6088(15936. 6yg o b (1330606 95^30(101358 cbb3oihnb 30(10163680
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3>"-
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655336b “658015(130(5 66(110319 503636. 56301 30(10163680 ,5(156630(15(5006 83936^9
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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 ^oSb,
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’b^ih'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'bc^ih^ 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 UftrtOUTI^O 3 6 d&r)6050
Incomplete development
of cuticle
J
Desiccation damage
- 371 -
91306^0360 bGtfObTJC? 9£? a n 9 6 6 0 n 6630
- 372 -
&360& 6030 V&MTMa&O
GA+CK levels
normal in xylem GA+CK levels
reduced In xylem
ACC absent
in xylem ACC accumulates
- 373 -
9006660060 bS^OUT)^ 9^ S n 9660 D 6630
- 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
6.2.4. 5 3 X 3 :)
- 375 -
0 0 0 6 ^ 0 0 6 0 U&tfOUT)^ 9^ a n 9^ A 0n 6^30
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 Jj0o9i^aii6, 330ng3hn jjgnSi^nbcr^nb ^n, 3nhnjncn,
g i 9ibibnicn3630Oi 6i 03j3&nb bn^ih63, 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 ^igig 903010-
0136b 3i 6n^i^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-
^ob 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 <b 3g i3nh xn Ri9 rn-
h3bb3i 6i03J36nb 5i ‘bm93nbib ih 5inoi3i0nb6n636i.
9)536ih336nb 53i 03noi gi^3nhcn3ob h3i 03hig ^ibi'bmSig, 5i6bij>3cnh36ooi 339-
I536ih3m6ibi g i 93iiy333'T>6i3n, 5i9rriny3636i b b 3igib b3i jho^3ho39n, hm9036ou
- 376 -
536366030 U&63U'3C^0 ^d6D 6360
90
Arctic tundra
80 Cold desert
- 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. <b 3 p i bjhsom : Shmgnpo >(»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
6 *. 6.57
bbjigibb3i
h3jnm6nb
5Qn3ipn>
5hs93&n
RDI = W S D ^ / WSDihn0) ( 6 .3 )
- 379 -
8oa6 6 6 3 3 an a ^ a r m tfa m .b a o
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
- 380 -
b'36a&A030 bSffjOUD^O 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
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)
- 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
► 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
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)
- 383 -
90a6^ 03&n b&6 0 li7)C? 8^ & n 9^(f.0n 6^3 l)
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
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 b3iX3^iyi. hmSx^HJ 6yi0b i^hr^^db p3hm9n (y>j^3bn/33q)mh6nib
^n3n); c b3j)3036(jn, hn9xo|J 6yi0b :>5hm336b cpmoi0369n (i&^i/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 (h3^i9ib ^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
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)
- 387 -
80066(^0060 usrtObT)1^ a^ana^onw 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)
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
- 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)
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 , ^Bbii^oih^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
- 390 -
bDBObrtOaO Ii6 rt0l>7)c?0 g 6d6 r)(f)060
- 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
6 .2 .5 .1. e ^ 0 C ^ 9 35 X n ^dfliWiWbfl
- 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)
- 393 -
9 0 0 6 ^ 3 0 6 0 b6 (f)3U D ^ g ^ a r m r t o m ^ a o
- 394 -
6716066030 UftrnOliTl^O 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
- 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 ).
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
- 397 -
9 0 D66ff)33J)0 b6 rt3b ^ 9*?8T)&>rfiami>90
- 398 -
67)6066030 3 6 d6 P)6360
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)
-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) ^ a^ana^am aao
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 L^igibbjh
ynd5m6;)6^n 3jh3Qnb dr^-jcmbnb 3hm3mhjjn:> (% ) nm&nb jmS^oSiihiijris 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
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 ^ihymqjnoio 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
Porphyro Icucosticla
Polyliphonia pulvinala
Chailomofpha linum
C troB iu m b c rn tri
Cafclhommon fra n u lo tw i
Clodophoro ulriculosa
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^n^imcn 5i0mnhfi33ni6, 3^301 35336016 hm&mhu 3n3m,yn63h, ^333
3n33 h(5m63p 3nhm&36b, 3 ngh3 3 mJ}33> 3 nj(j33nb <b m 6 i3 n (163 yri6b ^ i 6(j 3h ig 6m^nm 3 nhmi>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 .
C H pH CHpH
HO
HOOH HOCH
HOCH HOCH ho/ h ho\ h
- 401 -
9006663050 9«?an8M 3n 6.>30
- 402 -
536066030 U6 fnJUT)ggO gadacifnoao
Electrical conductivity i 6 S O 12 U 16 W
*1 the saturation extract m S-cm ->
- 403 -
8006660060 b&fnOb'a^ aei& rm foom .iao
- 404 -
a6 m r t m n & 36'3(f>o gid&nrta&o
- 405 -
9036i>ff)0060 uertDbTi^ a ^ a r m f n a n w a o
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
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
- 406 -
a e m f n m n a o G ^ o U6 6 0 b7)c?0 ^d & n rta & o
- 407 -
3(3 0 6 ^ 3 3 6 0 L 6 6 3 Lflg! 3^ s r ) 36f53r )5630
- 408 -
.>6m rism r)a067)cn0 b G r t a u ^ o ^d & n rta& o
409 -
3006660060 b S t f O b ^ 3 t?aD 36ffi0n 6^30
i*96o*03hn 'bhei .
- 410 -
^GuiAnanaae^tnO u & f n o u ^ o g M d n r ta 50
no
%
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 ^e3n"
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 6^36lihb^o36ob
(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 b^Ombqyhribb 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 hsbj^oob 3hm-
- 411 -
aoaeaAoa&o a ^ a rm rta n & a a o
]~ S° 2 ~ ]
- 412 -
660)ff,n3n a a 6'3(f.n b d t f o b ^ o
SO^Pollution
» \
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)
- 413 -
9036663060 9 ^5 0 9 6 6 9 0 6 6 3 0 366666 1160167)60
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)
- 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'^o sad&nAOMt
9006663060 LftrtOliZR 9^ 3 0 9 6 6 0 0 6 ^ 3 0
- 416 -
aemfnm n&aeafnO %>d6n«>9&o
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)
- 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
- 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
& 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 b^obb;) 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-
i^itib ijlyi3mbnb 153000361 bb3igsbb3i SdnSj Sj^cncn lb33nij8393&nb 333934, (Ernst, 1967; Ernst
& Joose-van Damme, 1983)
- 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>b^ih36nbbg3n
3jh3gob 3j96hi6nb Bneiifnag^m&i; 3
(jn^m3Qi'b3i3n 33^>onb giS^jhn
yncjdnbi 91 3m0n333^ne36nb (cgrvjmbj-
0^063611) 6njh bx^OOinb 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)
- 420 -
Ubhnpo 6.18 (jffijnoi Spngih Sm'bshp 93^(jQmqjnlj369n ^nByib ^iSs&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 33^i036nbig3n 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.
- 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
- 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
^iha^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: e^ojihjvniii 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
- 424 -
a6(J)ff>ror)&363(nO Stod&C)rn060
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 O^anO^fnODb^aO
- 426 -
6G m ^ n 3P)&3G7)(nn b<?>(f>3b3 ^ o sM&nfna&o
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>b0m6333c<i 333(yih36i.
3hrTH536^)36noi i^ihQtSDC0 jhoin bnoi gigbh3on qjihavn&n (Schmidt-Vogt, 1989).
S(r6n3630n biyicnbnb bbjipibbai ^3^5360 Qi djyxicm n6,53h3h3^>un3&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.
- 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 ho^i. 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)
- 430 -
^GtJ)tf>n3 naaG 3 ffio b ^ a u ^ n g6d6nA0&0
Soil acidification
- 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
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
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
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? a^an&w nam aao
- 434 -
6 Gmrtn3 n aa 6 T)(nO u & r ta u ^ o sfodsnfoo&o
- 435 -
8006660 D60 U&fnOUTlc? 3 ^& n 06 AOn 663 O
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(5361113011 ou6ibi'boi6igo>3&36o
bn6ioionb 830131^361 + /-
6yonb 330>3nb36i + /0
3n63hio36nb 330i3nb36i + /0
301633(136(301 + / 0 (C 3 > C 4)
- 436 -
a60 )<f>n3n&a 6'360 g6d&ncf>a&o
- 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
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&rtabT^o 36 d&rirt060
- 439 -
9006^0060 a ^ a r m fn o n ^ a o
M ean te m p e ra tu re fo r
th e w a rm e s t m onth
10 15 20 °C
- 440 -
_________________________ eadbaao ooaornaaagw g?n6a< ^ 6?)60U 6~aub5
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