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III$@!%(JANALYSIS OF SENSITIVITY TO WATER LIMITATION

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$@!!KD05$O?eITB-$K$h$C$F5/$3$k@8M}3XE*5!G=$NJQ2=$K$O$?$$$F$$4XO"$7$F$$$J$$!#0[$J$C$?@8D9CJ3,$N%H%&%b%m%3%7AH?%$NKD05$O!"?eITB-$KBP$7!"0[$J$C$?H?1~$r<($7$?!J(JBoyer 1970$@!K!#M?$($i$l$J$$?e$K$h$C$FJ,2=$7$F$$$k:YK&$N?-D9B.EY$,M^@)$5$l$?$H$-!":YK&$N3HBg$OJ,2=$7$?:YK&$N&$&W(Jt$@$N8:>/$H$H$b$K8:>/$7$?!#$7$+$7!"J,2=$7$F$$$J$$!J$C$F!"?eITB-$,5/$3$C$?$H$-!":YK&%5%$%:$O8:>/$7!"KD05$OJ,2=$7$F$$$k:YK&$G$O0lDj$G$"$k!J(JBoyer 1970$@!K!#(J
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IV$@!%(JEFFECTS OF WATER POTENTIAL ON PHYSIOLOGY$@!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(J

$@!!MUFy?e%]%F%s%7%c%k$N8:>/!J05%]%F%s%7%c%k$H6&$K!"$^$?$O!"$J$7$G!K$O!"$$$/$D$+$NJ}K!$G!":YK&@8M}$K1F6A$rM?$($k$3$H$,$G$-$k!#(J

$@!!#1!%8:>/$7$??e%]%F%s%7%c%k$O!"?e$N2=3XE*3hF0$r8:>/$5$;!"$=$l$K$h$C$F:YK&Fb$N?e!!!!!!$N9=B$$r=$@5$9$k!#(J
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$@!!#4!%KD05$NB;<:$O!"M"Aw%A%c%M%k$N6u4VE*$J0LCV$dKl9ZAG$NJQ2=!"Kl$N8|$5$N8:>/$r0z!!!!!!$-5/$3$9!#(J
$@!!#5!%:YKD05$NJQ2=$d:YK&JI$N9g@.1?F0$N8:>/$O867A$@!!#6!%FCDj$N>l=j$NJ,;RG;EY$O$$$/$D$+$N(Jsubcellular$@$N>l=j$N?eB;<:$N$?$a$KJQ2=$9$k(J
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$@#A!%(JEffects on Growth
$@!!?eITB-$,AH?%$N?eITB-$rA}Bg$5$;$k$N$K==J,$G$"$k$H$-!"KD05$O8=>]$9$k$@$m$&!#:YK&$NKDD%$O:YK&&W(Jt$@$K0MB8$7$F$$$k$N$G!"$3$l$i$N>u672<$GH/C#$7$F$$$k:YK&$O9-$,$j$,$h$j>/$J$/!":YK&%5%$%:$O$h$j>.$5$/$J$k$@$m$&!#$?$H$(:YK&$N3HD%$N7c$7$$8:>/$,5/$3$C$F$b!"?e%]%F%s%7%c%k$,8:>/$7$F$$$k4V$NJ,2=$7$F$$$k:YK&$N05%]%F%s%7%c%k$O0lDj$N$^$^$G$"$k!#=>$C$F!"8:>/$7$?KD05$N1F6A$O@.=O$7$?:YK&$+$iJ,2=$7$F$$$k:YK&$KEA$($i$l$?!#(J
$@!!?"J*A4BN$N@8D9%Q%?!<%s$KBP$9$k8:>/$7$?:YK&%5%$%:$N1F6A$O?"J*$N5(@a3X$K4X$9$k?eITB-$N%?%$%_%s%0$K0MB8$7$F$$$k!#$b$7!"?eITB-$,@8D9%5%$%/%k$N=i4|$K5/$3$C$?$J$i!"MULL@Q$O=L>.$7!"$h$j>.$5$$MU$N$?$a$K@8D94|4VCf!"C:AGA}2C$,8:>/$9$k$@$m$&!#B>$NFs/$+$i5/$3$k!#$b$7!"2V=xH/C#$N4V$K5/$3$C$?$J$i!"2V$N?t$O8:>/$7!"$b$7$+$9$k$H!"A4$F$NHK?#EXNO$,;_$^$k$+$b$7$l$J$$$,!"$=$l$K$b$+$+$o$i$:!"?"J*=8CD$K$*$$$FL51F6A$J$3$H$O$[$H$s$I$J$$$@$m$&!#$b$7!"2L $@:YK&3HD%$rM^@)$9$k(Jcritical $@?e%]%F%s%7%c%k$O
$@#B!%(JEffects of Cell Ultrastructure
$@!!Kl$N9=B$$d5!G=$K$*$1$k?eITB-$N0lHLE*$J1F6A$O$^$?:YK&$ND6Hy:Y9=B$$K$*$$$F$^$?4Q;!$G$-$k!#2:$d$+$J?eITB-$O%_%/%m%\%G%#!<$N9=B$$d:YK&$@!!%/%m%m%W%i%9%H$d%_%H%3%s%I%j%"$N9=B$$O7c$7$$?eITB-$K$h$C$F1F6A$r/$7$??e%]%F%s%7%c%k$K4XO"$9$k!J(JPoljenkoff-Mayber 1981,Steinmuller and Tevini 1985$@!K!#?e%9%H%l%9$N$+$+$C$??"J*$N%/%m%m%W%i%9%H$;$(4Q;!$5$l$?05Iz$7$?Kl$+$i$N%A%i%3%$%IKl$NJ,N%$O8w2=3X7O-6J#9gBN$N=|5n$N$?$a$G$"$k!J(JPowles 1984$@!K!#$=$l$K2C$($F!"7c$7$$?eITB-$O%5%$%H%=%kFb$N;iKC>.E)$N=P8=$d3K>.BN$N<~0O$N%/%m%^%A%s$NBg$-$J:FG[Ns$r0z$-5/$3$9!#(J

$@#C!%(JEffects on Photosynthesis$@!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(J
$@!!?eITB-$O!"2a5n#5#0G/$N4V!"9-HO0O$K$o$?$C$FJs9p$5$l$F$-$?8w9g@.$KB?$/$N1F6A$rM?$($k!#1F6A$OB?MM@-$G$O$"$k$,!"$=$l$K$D$$$F$NNc$d$$$/$D$+$N0lHLE*$J$3$H$,=R$Y$i$l$k!#8w9g@.$N?eITB-$N:G=i$N1F6A$OBgDq!"5$9&$NJD:?$G$"$k!#5$9&$O:,$N%7%0%J%k!"62$i$/%"%V%7%8%s;@!J(JDavies and Zhang 1991$@!K!"$^$?$O!"9&JU:YK&$NDc$$05%]%F%s%7%c%k!J(JRashke 1975,Collatz et al.1991$@!K$N$?$a$KJD$8$k$N$@$m$&!#?e%]%F%s%7%c%k$NJQ2=$K$O4X78$J$$$,!"MU$HBg5$4V$N>x5$05$N8{G[$NA}2C$KBP$9$kH?1~$G$bJD$8$k!#(J
$@!!?eITB-$K$h$C$F$b$?$i$5$l$k5$9&JD:?$O!":YK&6u4V$NFs;@2=C:AG$N>CLW$r0z$-5/$3$9!#$3$l$O!"8w9g@.$N5$9&M^@)$H8F$P$l$F$$$k!#(JCi$@$O0lEY!";@AG$KBP$7$F8:>/$7!"8w8F5[$,3h@-$5$l$k!#$b$7!"8w$N6/$5$,$+$J$j9b$$$N$J$i!"8w8F5[$OA4$F$NEE;REAC#7O$N%(%M%k%.!<$r$r4^$`;:J*$rMxMQ$G$-$J$$!#$3$N$h$&$J%7%J%j%*2<$G!"8wAK32$,5/$3$j!J(JBjorkman et al.1981)$@!"@x:_E*$K%/%m%m%W%i%9%HFb$G!"(Jfree radicals$@$NA}6/$N7k2L$K$J$k!#=88w@-%7%9%F%`!"5$9&JD:?4|4V$NEE;REAC#$N0lO"!"9b$$8w6/EY$K$*$1$k?e%9%H%l%9$N1F6A$O!"8w9g@.$K$H$C$F=EMW$G$"$k!#$3$N%5%V%8%'%/%H$O#1#0>O!JH/8w$N1F6A!K$d#1#3>O!JB?MM$J%9%H%l%9$NAj8_$N1F6A!K$K$o$?$C$F$$$k!#(J
$@!!8w9g@.$N(Jnonstomatal inhibition$@!J8wAK32$d%a%+%K%:%`$N$?$a!K$O!"?e%]%F%s%7%c%k$,Dc$/$J$k$H$-$N8w9g@.AK32$NBgItJ,$r=gEv$K@bL@$G$-$k!J(JOgren and Oquist 1985$@!K!#$7$+$7$J$,$i!"?eITB-!"5$9&JD:?!"(Jnonstomatal inhibition$@$NBh0lCJ3,$OF1;~$K5/$3$k!#Z5r$,$"$k!J(JBriggs et al.1986$@!K!#(J
$@!!?eITB-$,D>@\!"5$9&AuCV$rAK32$9$k$3$H$K$h$k%a%+%K%:%`$O!"5DO@$NM>CO$,$"$k!#$"$kJ,N`72$G$OJ}$NJ,N`72$G$O(Jrubisco$@3h@-$,(Jnonstomatal inhibition$@$r;YG[$7$F$$$k!#8:>/$7$?KD05$O!"30B&$N%/%m%m%W%i%9%H$NJ$$$$N?;F)@-$rA}2C$7!"%/%m%m%W%i%9%H$N#p#H$d%$%*%sG;EY$NJQ2=$N860x$K$J$k!#$"$k8&5f$G$O!"8w9g@.9ZAG$,?e%9%H%l%9$NM-32$J1F6A$KBP$7$FHf3SE*!"LH1V$,$"$k$3$H$,J,$+$j!"0lJ}!"B>$N8&5f$G$O8w9g@.9ZAG$K$*$1$kD>@\$N1F6A$r;X$7<($7$?!#!"?e%9%H%l%9$N4V!"%/%m%m%U%#%k$NDc2<$,A}Bg$7!"%/%m%m%U%#%kG;EY!JFC$K%?%s%Q%/J#9gBN$H7k$S$D$$$F$$$k#a!?#b!K$,8:>/$9$k!#=>$C$F!"8w2=3X7O-6$K4X$9$k=88w@-$dEE;REAC#$O?e%9%H%l%9$K$h$C$FM%@hE*$K!J8w2=3X7O-5$KHf$Y$F!K8:>/$9$k!#(J

$@#D!%(JEffects on Dark Respiration and Carbohydrate Metabolism
$@!!?eITB-$N7P2a$K$D$l$F!"8w9g@.$O8F5[$NA0$K8:>/$7!"=>$C$F8w9g@.$r8F5[4V$N3d9g$O8:>/$9$k!#8F5[$KBP$9$k8w9g@.$N3d9g$N8:>/!"?e%9%H%l%9$N4V$N8w8F5[$r0E8F5[N>J}$N@x:_E*$JA}2C$,B?$/$NCxeIt$K$h$jBg$-$J%@%a!<%8$r?"J*$,$@!!$$$/$D$+$N?"J*AH?%$NC1=c$JE|$NG;EY$O%/%m%m%W%i%9%H$KC_$($i$l$?%G%s%W%s$,F00w$5$l$k$N$G!"?e%9%H%l%9$G$OA}2C$9$k$@$m$&!#%G%s%W%s$NB;<:$O!"?eITB-$H$"$j$U$l$?Aj8_4X78$,$"$k$,!"C1=c$JC:?e2=J*$,$$$D$b%G%s%W%s$N8:>/$K=>$C$FA}2C$9$k$H$$$&$o$1$G$O$J$$!#%G%s%W%s$NF00w$+$i0z$-=P$5$l$?C1=c$JE|$OB?$/$N@8M}3XE*$J2aDx$K$h$C$F;H$o$l$k!#=>$C$F!"C1=c$JE|$NC_$($NA}2C$,?e%9%H%l%94V$N%G%s%W%s$N8:>/$N@bL@$K$J$i$J$$$3$H$O6C$+$J$$!#(J
$@!!F|Cf$N?eITB-$N4VC:?e2=J*$NE>N.$O$^$?8:>/$9$k$,!"Lk$K==J,$K?e$rM?$($??"J*$OHf3SE*A}2C$9$k!J(JBunce 1982$@!K!#%9%/%m!<%9E>N.$N8:>/$O2aDx$r(Jloading$@$9$k;UIt$NFCJL$J1F6A$G$O$J$$!#(Jloading$@$9$k;UIt$O!"Hf3SE*?eITB-$KDq93$,$"$k!J(Jsung and krieg 1979$@!K!#8:>/$7$?8w9g@.$NE>N.$N860x$O?e%9%H%l%9$N4V$N%=!<%9$H?<9H$N4X78$NJQ2=$G$"$k!#MU$K$h$kDc$$#C#O(J2$@F12=$rMUFy:YK&$N8F5[$OMU$N%=!<%9$H8w9g@.$N%7%s%/$N4V$N%9%/%m!<%98{G[$r8:>/$5$;$k!#%=!<%9$+$i%7%s%/$X$N8{G[$N8:>/$O;UIt$NC:?e2=J*$NN.$l$N8:>/$r0z$-5/$3$9!#(J
$@!!;q8;G[CV%Q%?!<%s$O?eITB-$N4V!"JQ2=$9$k!#B?$/$N$C$F!":,!?CO>eIt$NHfN($O8:>/$9$k!#?"J*$NH/C#>uBV$KHf$Y$F?eITB-$N%?%$%_%s%0$OG[CV%Q%?!<%s$K1F6A$9$k!#=i4|@8D9CJ3,$G5/$3$k?e%9%H%l%9$O:,!?CO>eIt$NHfN($NBg$-$JJQ2=$N860x$H$J$k!#Hf3S$9$k$H!"@8?#4|!J@8D98e4|!K$G$O?e%9%H%l%9$O:,!?CO>eIt$NHfN($K$[$H$s$IL58z$G$O$J$$$,!"2V$H/$7!"2L
$@#E!%(JEffects on Nitrogen metabolism$@!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(J
$@!!>K;@1v$H%"%s%b%K%"$NC_@Q$O!"?eITB-$N4V$K8:>/$9$k!#:,$+$iMU$X$NCbAG$NN.$l$OCY$/!">K;@1v$H%"%s%b%K%"$O==J,$K?e$rM?$($??"J*$N:,$h$j$b!"?e%9%H%l%9$N:,$NJ}$,9bG;EY$G$"$k!J(JNilsen and Muller 1981a$@!K!#?e%9%H%l%9$N?"J*$N:,$N9bG;EYCbAG%$%*%s$O!"EZ>m$+$i$NCbAG$NC_@Q$rM^@)$9$k!#=>$C$F!"CbAGC_@Q$N8:>/$O!"M"Aw%?%s%Q%/$dC_@Q$N%a%+%K%:%`$K$*$1$k?e%9%H%l%9$NFCDj$N1F6A$G$O$J$$!#$`$7$m!"CbAG$NJQ2=$rMQ$$$F!"N.$l$,CbAGC_@Q$NF0NO$rM^@)$9$k?"J*$N>uBV$K$J$k!#(J
$@!!?eITB-$O!"%?%s%Q%/J,2r$NA}2C!"%?%s%Q%/9g@.$N8:>/$K4XO"$9$k!#2C$($F!"B??t$N%]%j%j%\%=!<%`$N8:>/$O%?%s%Q%/9g@.$N8:>/$K4XO"$9$k!#M7N%%"%_%N;@$NA}2C$O!"Am%?%s%Q%//$HF1;~$K5/$3$C$?!#B?$/$N%"%_%N;@C_@Q$O%?%s%Q%//$N$?$a$G$"$k$,!"$"$k%1!<%9$G$OFCDj$NHs%"%_%N;@$N@89g@.$,3h@-$5$l$?!JNc!"%Y%?%$%s!"%W%m%j%s!K!#!!$$$/$D$+$N8&5f$G$O!"?eITB-$N4V$K%j%\%L%/%l%"!<%<$NA}2C$,5/$3$k$3$H$r<($7$F$$$k!##m#R#N#AK]Lu$,1F6A$rM?$($i$l$k$3$H$K$O$[$H$s$I>Z5r$,$J$$!J(JShah and loomis 1965$@!K!#=>$C$F%5%$%H%=%k$N#m#R#N#A$NC_$($O62$i$/%5%$%H%=%k$N%j%\%L%/%l%"!<%<3h@-$,A}2C$9$k$?$a$K8:>/$9$k!J(JTodd 1972$@!K!#(J

$@#F!%(JGeneralized Response to Water stress
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$@!!B?$/$NCxe>:$K4XO"$7$F$$$k$N$O!"8F5[$,8:>/$7!"uBV$N?"J*$K;w$?MM;R$N7c$7$$MU$NF0:n$r5/$3$9!#$J$<$J$i!"?eITB-$rm!"$^$?$O:,$NG^2p?e%]%F%s%7%c%k$r5/$3$7!"?eITB-$N1F6A$+$iFCDj$N%$%*%s$N1F6A$rJ,N%$9$k$3$H$OFq$7$$!#=8Cf$9$k0lHLE*$J%9%H%l%9$KBP$9$kH?1~$O=R$Y$i$l!"$=$l$i$O;q8;ITB-$N:YK&J,;R!"2rK6>e$NH?1~$N4V$GB?$/$NN`;w$,$"$k$,!"3F;q8;!J8w!"?e!"G.!K$O$^$?!"$h$/9M$($M$P$J$i$J$$@8D9!"H/C#$NFHFC$N1F6A$r;}$D!#(J


$@!!!!!!!!!!!!!!!!!!!!!!!!!!(J $@!!(J1999,6/7 $@%+%s%@a(JBR>-$@#S#T#U#D#Y(J-$@#R#E#V#I#E#W!!#O#U#T#L#I#N#E(J-

Soil-Induced Water Limitation

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$@#3!%M-5!J*$OEZ>m4V7d$K$*$1$kM-5!J*$N1F6A$K$h$jEZ>m?eJ,J];}MFNL$rA}2C$5$;$k!#(J

$@#4!%A}2C$7$?EZ>m1vJ,$OEZ>m?;F)%]%F%s%7%c%k$rDc$/$9$k$N$G!"?"J*$KBP$9$k?eM-8z@-$r!!!!8:>/$5$;$k!#(J

$@#5!%EZ>m?eJ,$N$NMW0x$O!"EZ>m$NAX!"(Jcompaction$@!"L5@TDGF0J*$N!!!!3hF0$G$"$k!#(J

$@#6!%EZ>m$N?eNO3XE*%3%s%@%/%?%s%9$O!"EZ>m9&$rDL$k:,$NI=LL$rN.$l$k?e$NB.EY$rD4@a$9!!!!$k!#(J

$@#7!%?eNO3XE*%3%s%@%/%?%s%9$rD4@a$9$kMW0x$O!"EZ>m$N@-m$N9&!"EZ>m$N@-mCDN3$N@-
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$@#9!%2?$i$+$N72Mn$r9=@.$9$km$N0LCV$+$i?e$N!!!!Cj=P$r@8$8$k!#(J

Non-Soil-Induced Water Limitation

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$@#2!%CO7AE*$J0LCV$HCO0hE*$J5$8u$O$I$N>l=j$G$G$b9_1+$rD4@a$9$k!#(J
$@!!!!9_1+$O#1G/$4$H$K(J2000_$@$+$i>/NL$^$GJQ2=$G$-$k!#(J

$@#3!%0lHL$K!">/$J$$9_1+$O!"6/8w!"G.!">xH/$N<{MW$H6&$K5/$3$k!#(J

$@#4!%B?$/$N;X?t$O!"AjBPE*$J?e%P%i%s%9$K4p$E$$$F>l=j$rJ,N`$9$k$?$a$KH/C#$7$F$-$?!#!!!!$7$+$7$J$,$i!"?"J*$KBP$9$k?eM-8z@-$N:GNI$N<\EY$OLkL@$1A0$N?e%]%F%s%7%c%k$G$"!!!!$k!#(J
$@#5!%EZ>mCf$N?eJ,$,==J,$G$b!"MU$N?eJ,MW5a$K6!5k$9$k$K$O0]4IB+$N9=@.$,IT==J,$G$"$k!!!!$N$G!"?"J*$O6/8w$dF|Cf$K$7$*$l$k!"(J

Sensitivity Analysis

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$@#2!%AH?%?e%]%F%s%7%c%k$NJQ2=$KBP$7$F:G$bIR46$J@8M}3XE*$J2aDx$O!":YK&$N3HD%!":YK&!!JI$N@8D9!"%?%s%Q%/9g@.!">K;@1v$N8:>/$G$"$k!#(J

$@#3!%JQ2=$9$kAH?%$N?e4^M-NL$K:G$bB.$/JQ2=$5$;$k?e%]%F%s%7%c%k$N9=@.ItJ,$O!"05%]%F!!%s%7%c%k$G$"$k!#(J

$@#4!%05%]%F%s%7%c%k$O!":G$bNI$$?"J*?e%9%H%l%9$N;X?K$H$_$J$5$l$F$$$k!#(J

Effects of Reduced Water Potential on Physiology

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$@#5!%:YK&$N05NO$NJQ2=$H7k2L$H$7$F@8$8$k:YK&JI$N<}=L$O!"867A
$@#6!%FCDj$NIt0L$G$NJ,;R$NG;EY$O!"$"$k(Jsubcellular$@0LCV$NB;<:$N$?$a$KJQ2=$9$k$+$b$7!!!!$l$J$$!#(J

Effects on Growth

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$@#3!%:YK&JI$,$b$?$i$98B3&$O!"effects on Cell Ultrastructure

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Effects on Photosynthesis

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$@#2!%?e%9%H%l%9$K$h$k8w9g@.$N5$9&$G$J$$AK32$O(Jrubisco$@3h@-$K$*$1$k1F6A$HF1MM$K8wH?!!!!1~$N9=@.ItJ,$K$*$1$kD>@\$N1F6A$r4^$s$G$$$k!#(J

Effects on Dark Respiration

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$@#4!%8w9g@.$NE>N.$O?e%9%H%l%9$N4V!"8:>/$5$l$?!"$J$<$J$iMU$O!":GE,$N?e%3%s%G%#%7%g!!!!%s2<$G8w9g@.$KBP$7$F6/$$%=!<%9$G$O$J$$!#(J

Effects on Nitrogen Metabolism

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generalized Response on Water Stress

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Mechanisms by Which Plants compensate for Water Limitation

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Drought tolerance with Low Water potential

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osmotic Potential Adjustment

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tissue Elasticity

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Symplastic Water Fraction

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Cavitation Susceptibility

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Desccation Tolerance

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Drought tolerance with High Water potential

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reduction of Water Loss

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$@!!!!&K$O>x;6$r8w9g@.$NHfN($N(Jsecond differential$@$rI=$9!#(J

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Enhancement of Water Accumulation

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