$@H>>lM4;R$N%[!<%`%Z!<%8!J%<%

The Physiology of Plants under Stress (E. T. Nilsen & D. M. Orcott)

5. Stable Isotopes and Plant Stress Physiology

2000$@G/(J2$@7n(J14$@F|!!C4Ev!'$O$s$P(J

I Introduction

$@!z0BDjF10LBN$rB,$k0U5A(J
$@4D6-$KBP$9$k?"J*$NH?1~$rD4$Y$k$?$a$K9T$o$l$k!"@82=3XE*!&@8M}3XE*$JB,Dj$O!"MM!9$J@)Ls$r
$@!&;~4VE*!"6u4VE*@)Ls(J
$@8w9g@.!'%/%m%m%W%i%9%H!":YK&!"%W%m%H%W%i%9%H!"MUJR!"MU!";^$GB,$k$3$H$,$G$-$k$,!"?"J*A4BN$GB,Dj$5$l$k$3$H$O>/$J$$!#(J
$@%,%98r49$d9ZAG!&@82=3XE*$JB,Dj!'C;$$;~4V$N%9%J%C%W%7%g%C%H$7$+F@$i$l$J$$!#?"J*A4BN$NH?1~$r$_$k$?$a$K$O!"?t3XE*$J7W;;$K$h$C$F%9%1!<%k%"%C%W$9$k$3$H$,I,MW(J

$@!&%5%s%W%k%5%$%:$N@)Ls(J
$@@8M}E*$JB,Dj$G$O!"%5%s%W%k%5%$%:$,@)Ls$5$l$k$3$H$,B?$$!#%,%98r49B,Dj$G$O0l;~4VEv$?$j(J40$@!A(J70$@%5%s%W%k$rB,Dj$G$-$k$,!"$=$l$G$bMU72A4BN$NHy4D6-$dNp9=B$$r7W;;$9$k$K$OIT==J,(J
$@05NO!/$7$N%5%s%W%k$7$+B,Dj$G$-$J$$!#9ZAG$NB,Dj$O!"
$@"*?"J*A4BN$N5(@aJQ2=$dG/JQ2=$rD4$Y$k$K$O!"8EE5E*$J@8M}E*B,Dj$G$OIT==J,!#(J
$@?"J*$N0BDjF10LBN$NHfN($O!"D94|4V$N@Q;;E*$JCM$J$N$G!"?"J*$N@8M}E*$J%W%m%;%9$N;XI8$H$7$FE,$7$F$$$k(J

$@!z0BDjF10LBN$K$D$$$F(J
$@F10LBN!'3K$NCf@-;R?t$,0[$J$k$?$a!"0[$J$k
$@F10LBN$O!"0BDj@-$,0[$J$k!J(JFig. 5.1$@!K!#(J
$@J|
$@J|8:(J_$@a(Ji$@J|J,$,0BDj$J7A$K$J$k$N$KI,MW$J;~4V!K$O!"(J10C$@$G?tIC!"(J14C$@$G?t@iG/!J(JFig. 5.1$@!K(J

$@!zJ|$@H>8:4|$NC;$$$b$N$O!"J,;R<1JL%^!<%+!<$H$7$F!"@82=3XE*$J%H%l!<%5!<8&5f$KMxMQ!#(J
32P$@!'%?%s%Q%/$N%i%Y%j%s%0!"3K;@9=B$$N%W%m!<%V(J
11C$@!'(Jin vivo$@$NC:?e2=J*M"Aw!#%,%$%,!<%+%&%s%?!<$G%,%s%^@~$rB,Dj$G$-$k$N$G!"HsGK2uB,Dj$,$G$-$k$,!"H>8:4|$,Hs>o$KC;$$$N$G%5%$%/%k%H%m%s$,6a$/$K$"$k14C$@!'D9$$@82=3XE*$J7PO)$NCf$K$"$kC:AG:?$dM-5!J,;R$N<1JL$dDI@W!#(J
14C$@!"(J235U$@!'CO8:4|$,Hs>o$KD9$$$?$a!#(J235U$@$O(J7$@2/G/!K!#(J14C$@$O3Km$d
$@!z0BDjF10LBN$NMxMQ(J
$@0BDjF10LBN$K$OH>8:4|$O$J$/!"%(%M%k%.!<$dN3;R$bJ|=P$7$J$$$N$G!"%7%s%A%l!<%7%g%s%+%&%s%?!<$d%,%$%,!<%+%&%s%?!<$G$OB,Dj$G$-$J$$!#(J
$@0BDjF10LBN$O!"
$@0BDjF10LBN<+A3B8:_Hf!'$"$k0BDjF10LBN$,85AGA4BN!J$"$k$$$O$"$kFCDj$N%W!<%k!K$K@j$a$kHfN(!#F10LBN$NB8:_HfN($O!"J,;R$d86;R$N7k9g$,2u$l$?$j!"J,;R$,3H;6$7$?$j$9$k$H$-$K5/$3$kF10LBN8z2L$K$h$C$FJQF0$9$k(J

$@!zF10LBN8z2L(J
$@B.EYO@E*8z2L!"G.NO3XE*8z2L$N(J2$@$@B.EYO@E*8z2L!'3H;6B.EY$d!"9ZAG$NH?1~@-$N0c$$$J$I$K$h$C$F!"HsJ?9U%W%m%;%9$G5/$3$k8z2L!#=E$$F10LBN$,J,JL$5$l$k$3$H$,B?$/!"29EY0MB8@-$,$"$k(J
$@G.NO3XE*8z2L!'Fs$D$NB.EYO@E*$JH?1~$,J?9U$K$J$C$F$$$k$H$-$K5/$3$k8z2L!#Nc$($P1UAj$H5$Aj$N4V$G3H;6$K$h$C$FF10LBN$,0\F0$7$F$$$k>l9g!#29EY$KIR46(J

$@?"J*$G!"@8J*3XE*$JF10LBN$NJ,JL8z2L$,$I$NDxEY@8$8$k$+$O!"?"J*$N@8M}E*$J%W%m%;%9$rH?1G$9$k!#$^$?!"?"J*$NJ,JL8z2L$K$h$C$F!"<~JU$N4D6-$NF10LBNHf$bJQF0$9$k"*%9%H%l%9;XI8$H$7$FF10LBN$,;H$($k(J

$@?"J*A4BN$N@8M}3X$N;XI8$H$7$F$NF10LBNHf$N8&5f$O!"(J40$@G/0J>e9T$o$l$F$$$k!J(JCraig 1945$@!K!#(JC$@!"(JH$@!"(JN$@!"(JO$@$N8&5f$,B?$$!#Hf3SE*-Mh$5$i$K1~MQ$,4|BT$G$-$k(J

$@!zF10LBNHf$NJQF0(J
$@F10LBNHf$NJQF0$O!"$"$k4p=`J*2H$@!'<+A3B8:_Hf$,>.$5$$$N$G!"(Jd$@$O>.$5$/$J$k!#$7$+$7!"(Jd$@CM$NJQF0$OBg$-$$!JB8:_Hf$,(J7.5 *10-6$@JQ$o$k$H!"(Jd$@$O(J50$@gu%^F0!"DL>o$O(J50$@!A(J100$@gu%^F0!K!#(J

II Carbon Stable Isotopes
$@#2$,(J12C$@!J(J98.9$@!s!K!#M-5!J*$r$$$C$?$s4%Ag$5$;$l$PF10LBNHf$O$+$o$i$J$$$N$G!"Hy@8J*$N8F5[$rKI$2$P!"0BDjF10LBNHf$OD94|4V0BDj(J

$@0BDjF10LBNHf$NJQF0$O$-$o$a$F>.$5$$$?$a!"C10L$H$7$F$O(J1000$@J,N(!J%Q!<%_%k!K$,MQ$$$i$l$k!#$^$?!"DL>o!"I8=`J*

$@C:AG$N0BDjF10LBN$N9q:]E*$JI8=`J*
$@?"J*$NF10LBNHf$O!"4p
$@F10LBN$NI=8=$H$7$F$O!"(Jd13C $@$,;H$o$l$k$3$H$NJ}$,B?$$!J4pm8F5[$d?M0YE*$J1F6A$,$"$k>l9g$K$OJQ2=$9$k"*F10LBNJ,JL$r7W;;$9$k$H$-$K$OBg5$$N(Jd13C$@$rB,Dj$9$kI,MW$,$"$k(J

II Fractionation of Carbon Stable Isotopes

$@3H;6$N:]$K$O!"=E$$F10LBN(J13C $@$KBP$9$kJ,JL$,5/$3$k!#C:AG$N0BDjF10LBN$NJQF0$O
C3$@?"J*Fb$G$O!"8w9g@.$K$h$C$F!"Fs;@2=C:AG$,=i4|8w9g@.;:J*$KF12=$5$l$k:]$K!"Bg$-$JF10LBNHf$NJQ2=$,5/$3$k!#(JFarquhar$@$i(J(1989)$@$N%b%G%k$K$h$k$H!"(JC3$@?"J*$K$*$$$F!"F10LBNHf$O!"6a;wE*$K$O!"8w9g@.$N:]$K5/$3$kF10LBNHfJQ2=$H!"?"J*$,
D = a + (b$@!](J a) Ci/Ca

$@$3$3$G(JD $@$O8w9g@.$KH<$&F10LBNHf$NJQ2=!JF10LBNJ,JL!K$G!"?"J*MUFb$NFs;@2=C:AGJ,05!J(JCi$@!K$H?"J*<~0O$NFs;@2=C:AGJ,05!J(JCa$@!K$NHf$H@5$NAj4X$,$"$k!#(Ja$@!J(J4.4 $@ga(JA$@M}O@CM!K$O!"Fs;@2=C:AG$,5$9&$r=PF~$j$9$k:]$K5/$3$kF10LBNHf$NJQ2=!"(Jb$@$O!"?"J*MUFb$NFs;@2=C:AG$,!"9ZAG(J RuBisco$@$K$h$C$FM-5!J*$KF12=$5$l$k:]$K5/$3$kF10LBNHf$NJQ2=!#(J
$@Nc$($P!"5$9&$,JD$8$?>l9g!"MUFb$NFs;@2=C:AGJ,05$O>.$5$/$J$k!J(JCi/Ca$@$,>.$5$/$J$k!K!#$3$N>l9g!"(JD $@$O(Ja$@$K6a$$CM$K$J$k!#5U$K5$9&$,3+$$$F$$$F(JCi/Ca$@$,Bg$-$$$H$-$K$O!"(JD $@$O(Jb$@$K6a$$CM$K$J$k(J

RuBisco$@$K$h$kJ,JL!'M}O@E*$K$O(J30 $@ga(JiCO2$@$@$1$,4pl9g!K!"JD:?7O$G$N

$@!zBe$@C:AG$N4T85$K$h$kF10LBN8z2L!';i/$J$$!#(J
$@MU$OLZItAH?%$h$jF10LBNE*$K7Z$$(J

$@!z1vN`G;EY$HF10LBN8z2L(J
$@1vN`G;EY$,9b$$$H!"5$9&$,JD$8$F(JCi$@$,>.$5$/$J$k$N$G!"(Jd13C$@$O$h$j%W%i%9$K$J$k(J
$@1v@8?"J*$G$O!"1vN`G;EY$H(Jd13C$@$K@5$NAj4X$,$"$k!'1v@8?"J*$O!":GE,$J8w9g@.$r$9$k$?$a$K$O1vN`$,I,MW$J$N$G!"1vN`G;EY$,Dc2<$9$k$H8w9g@.B.EY$,%3%s%@%/%?%s%9$KHf$Y$F8:>/"*(JCi$@$,Bg$-$/$J$C$F(Jd13C$@$O$h$j%^%$%J%9$H$J$k(J

$@!z5$9&$N3+JD$HF10LBN8z2L(J
C3$@?"J*$G$O!"Bg5$1x@wJ*.$5$/$J$k$N$G!"(Jd13C$@$O$h$j%W%i%9$K$J$k(J

d13C$@$O?eMxMQ8zN($N;XI8$H$J$k!J(JFig. 5.6$@!K!#$J$<$J$i!"?eMxMQ8zN($O0BDjF10LBNHf$H0J2<$N$h$&$J4X78$K$"$k$+$i!'(J

$@29<<$d%0%m!<%9%A%c%s%P$G$Nr7o2<$G$O!"(Jd13C$@$O?eMxMQ8zN($N$h$$;XI8$H$J$k!#(J
$@Nc!':=Gy$N?"J*$N!"8w9g@.$r9T$C$F$$$k;^$O!"MU$h$j$b(Jd13C$@$,$h$j%W%i%9$NCM!"$D$^$j?eMxMQ8zN($,9b$$!J(JFig. 5.7$@!K(J
$@H>4s@8?"J*$N?eMxMQ8zN($d(Jd13C$@$O=I
$@!zNS4'$N>x;6(J
$@=c0lx;6$r?dDj$9$k$3$H$,$G$-$k!'(J

$@$7$+$7!"$^$@8!>Z$5$l$F$$$J$$(J

$@!z%9%/%j!<%K%s%0(J
$@MU$NAH?%$N(Jd13C$@$r$b$H$K!"?eMxMQ8zN($,9b$$IJ$@$7$+$7!"(Jd13C$@$H?"J*$N@.D9$dl9g$O@5$NAj4X!"$"$k>l9g$K$OIi$NAj4X!#0dEAE*$J1F6A$+!)(J

$@0BDjF10LBNHf$N9-5A$N0dEAN($O$5$^$6$^$J
III Nitrogen Stable Isotopes
$@CbAG$K$O#2
$@!zCbAG$N0BDjF10LBNHf$NMxMQK!(J
$@#1!KCbAGBe$@#2!K(J15N$@$N<+A3B8:_Hf$r%5%s%W%k$HBg5$$H$GHf3S$9$k(J

A. Fractionation of Nitrogen Stable Isotopes
$@C:AG$N>l9g$HF1$8$/!"J*M}E*$J%W%m%;%9!J3H;6$J$I!K$K$h$C$F=E$$F10LBN(J15N$@$,J,JL$5$l$k(J
$@@8J*E*$J%W%m%;%9$G$O!"(JNO3-$@$+(JNH4+$@$N7A$NCbAG$,MxMQ$5$l$k!JCbAG8GDj$ONc30!K(J
$@EZ>mMO1UCf$N(JNO3-$@$d(JNH4+$@$N(J15N$@B8:_NL$O!"@8BV7O$G$ND94|4V$NB.EYO@E*!&G.NO3XE*$JF10LBN8z2L$K$h$C$F7h$^$C$F$$$k(J

$@!z@8J*E*CbAG$NCbAG0BDjF10LBNHf(J

$@EZ>m$N?<$5$K$h$C$F!"CbAG$N0BDjF10LBNHf$OJQ2=$9$k(J
$@EZ>m$NCbAG$NF10LBNHf$O!"?"J*$,MxMQ$7$F$$$kCbAG$NF10LBNHf$rH?1G$7$F$$$J$$$3$H$,$"$k(J

$@!zEZ>mCbAG$N0BDjF10LBN$NJQF0MW0x(J
$@Hy@8J*3XE*!&@82=3XE*$JF10LBN8z2L$K$h$k!JI=(J5.2)
$@C&Cb!'C&Cb$K$h$C$F7Z$$F10LBN$r$b$DCbAG(J14N2$@$,=P$F9T$/$N$G!"C&Cb$,@9$s$JEZ>m$G$O(J15N$@$,B?$/$J$k(J
$@;\Hn$5$l$?!"%"%s%b%K%"$,B?$$EZ>m$d(JpH$@$,9b$$EZ>m!'%"%s%b%K%"$N4xH/!J(JNH4++OH- $@"*(JNH3$@",(J+H2O$@!K$,@9$s$K@8$8!"(J15N$@$,B?$/$J$k(J
$@CbAG8GDj$9$k?"J*$N%j%?!<$NJ,2r!'(J15N$@$,B?$/$J$k(J

$@0J>e$N%W%m%;%9$NAH$_9g$o$;$K$h$C$F!"(J15N$@$,$I$NDxEYB?$/$J$k$+$OJQ2=$9$k"*EZ>mCbAG$N(Jd15N$@$O$=$N>l=j$KFC0[E*(J

$@!z?"J*$N(Jd15N$@$NJQF0MW0x!JI=(J5.2$@!K(J
$@>K;@$dCbAGF12=!"E>N.!":YK&
$@>K;@/!">K;@$NNL$H$H$b$KA}2C!"8w6/EY$H$H$b$K8:>/$9$k!#$7$+$7!"DL>o$N1IM\>r7o$G2:$d$+$J8w6/EY$G$"$l$P!">K;@.$5$$(J

$@CbAG8GDj$N:]$NF10LBN8z2L!'(J-9$@!A(J4.1$@ga(JA$@?"J*AH?%$d
$@E>N.!'?"J*AH?%$N(Jd15N$@$NJQF0MW0x$K$J$j$&$k$,!"$"$^$j=EMW$G$O$J$$(J

$@:,N3$r=|$-!"?"J*$N(Jd15N$@$OHf3SE*0lDj!JI=(J5.3$@!K$@$,!";^$d:,$NF10LBN$K$O(J15N$@$,>/$J$/$J$k798~(J

$@3hH/$K3hF0$7$F$$$F!"%P%/%F%j%"$,B?$/B8:_$9$k:,N3$G$O(J15N$@$,FC$KB?$/$J$k!#$=$N%a%+%K%:%`$OITL@$@$,!"=I
B. Physiological Processes
d15N$@$rMxMQ$7$?@8M}3XE*$J%W%m%;%9$N8&5f!'CbAG8GDj$K$h$C$F@8J*E*$K8GDj$5$l$?CbAG$,!"?"J*A4BN$NCbAG$N$I$NDxEY$N3d9g$r@j$a$k$+$rD4$Y$k(J

1. Studies of Nitrogen FIxation by Soil Enrichment
15N$@$rMxMQ$9$k%F%/%K%C%/$O!"%"%;%A%l%s4T85K!$KJQ$o$C$F(J1970$@G/Be$NH>$P$KEP>l!J%"%;%A%l%s4T85K!$G$O!"8D!9$N:,N3$+$i?"J*A4BN$K%9%1!<%k%"%C%W$9$k:]$KBg$-$J%(%i!<$,$"$C$?!K(J

$@!zCbAG8GDj$N2r@O(J
15N$@$rB?$/4^$`HnNA$r!"CbAG8GDj$9$k?"J*$HCbAG8GDj$7$J$$?"J*$KM?$(!"EZ>m$N(Jd15N$@$rB,Dj"*N>J}$N?"J*$K$D$$$FEZ>m$NCbAG=[4D$,F1$8$G!"EZ>m$N(Jd15N$@$NJ,I[$,Ey$7$$$J$i$P!"CbAG8GDj$9$k?"J*$N<~0O$NEZ>m(Jd15N$@$O$h$jDc$/$J$k!J:,N3$K(J15N$@$,C_@Q$9$k$?$a!K(J

$@CbAG8GDj$N?dDj$K!"EZ>m$N(Jd15N$@$rMxMQ$9$k>l9g!"IT6Q0l@-$,LdBj$H$J$k!'(J15N$@$NBgItJ,$OEZ>mI=LL$KJP$C$F$$$k!&EZ>mHy@8J*$N6u4VE*$JJ,I[$,JQF0$7$F$$$k!&EZ>m$N>K;@$N(Jd15N$@$O;~4V$H$H$b$K8:>/(J

$@>e5-$N7gE@$rJd$&$?$a!"EZ>m$G$O$J$/!"%j%U%!%l%s%9$H$J$k:nJ*$N(Jd15N$@$,MxMQ$5$l$k!#%j%U%!%l%s%9:nJ*$O!"D4$Y$k:nJ*$HF1$8$h$&$J:,7w$r$b$A!":,7w$KCbAG8GDjAH?%$,$J$$!"5(@aE*$J>K;@F12=$,;w$F$$$k?"J*$rCm0U?<$/A*$VI,MW$,$"$k(J

d15N$@$+$i!"CbAG8GDj$N3d9g$r7W;;$9$kJ}K!!'CbAG$N6!5k8;$H$7$F!"#3$D!JBg5$!"EZ>mMO1U!"HnNA!K$r9M$($k(J

$@?"J*$,HnNA$+$iF@$?CbAG$N3d9g(Jy

$@HnNA$NMxMQ8zN((Jw

$@0lJ}!"CbAG8GDj$K$h$C$F?"J*$K8GDj$5$l$?CbAG$NNL$O!"%j%U%!%l%s%9:nJ*$HL\E*$N:nJ*$N(Jd15N$@$+$i7W;;$5$l$k(J

$@CbAG8GDj$K$h$k8GDjNL(J=

$@$D$^$j!"CbAG8GDj$N3d9g$O!"%j%U%!%l%s%9:nJ*$HL\E*$N:nJ*$N(Jd15N$@!"$*$h$SL\E*$H$9$k:nJ*$NA4CbAGNL$N#3$D$rB,$l$P!"?d;;$G$-$k(J

$@>e5-$N?d;;$r$9$k>e$G$NCm0UE@!'EZ>m$N>K;@G;EY$,>e$,$C$FCbAG8GDj$,AK32$5$l$k$N$rKI$0$?$a!"(J15N$@$r4^$`HnNA$OM?$($9$.$F$O$$$1$J$$!#$^$?!"C&Cb$K$h$C$FBg5$$h$j$b(J15N$@$rB?$/4^$`CbAG%,%9$,H/@8$7!"?"J*$,$3$NCbAG$r8GDj$7$?>l9g!"?"J*$N(Jd15N$@$,$h$j%W%i%9$K$J$k$N$G!"CbAG8GDj$N3d9g$O2a>.I>2A$5$l$F$7$^$&!#$b$C$H$bBg$-$J8m:9$N860x$K$J$k$N$O!"ITE,Ev$J%j%U%!%l%s%9?"J*$NA*Br!J(Jd15N$@$OF1$8CM$G$J$/$F$O$J$i$J$$!K(J

2. Natural Abundance Technique
$@!zCbAG8GDj(J
$@<+A3$N@8BV7O$K(J15N$@$r4^$`HnNA$rM?$(!"(Jd15N$@$N<+A3B8:_Hf$+$iCbAG8GDj$r?dDj$9$kJ}K!$O!"MM!9$JLdBjE@$,$"$k!'8e$+$iJL$NCbAG$,2C$o$C$F$7$^$&!&(J15N$@$r4^$`HnNA$NJ,I[$,JQ$o$C$F$7$^$&!&HnNA$rM?$($kA0$K$b?"J*$KCbAG$,BgNL$KB8:_$7$F$$$k$N$G!"HnNA$N(J15N$@$N8z2L$,(Jd15N$@$K$"$^$jH?1G$5$l$J$$(J

$@<+A3@8BV7O$GCbAG8GDj$N3d9g$r5a$a$k$?$a$K$O!"
$@!!CbAG8GDj3d9g(JFNDfa=

$@2a5n(J10$@!A(J15$@G/4V$K!":=Gy$d:nJ*$d%A%c%Q%i%k$dG.BSNS$G!"<+A3B8:_Hf$r;H$C$FCbAG8GDj$r?dDj$9$kJ}K!$,MxMQ$5$l$F$$$k(J

$@>e5-$N?dDjJ}K!$K$bBg$-$J7gE@$,$"$k(J
$@#1!K%j%U%!%l%s%9?"J*$HL\E*$N?"J*$N(Jd15N$@$O!"$"$^$jBg$-$J0c$$$,$J$$$?$a!J(J10$@ga(Jj$@!"8DBN$dAH?%4V$N:9$d0dEAE*$J:9!"B,Dj$N%(%i!<$J$I$NJ}$,Bg$-$/$J$C$F$7$^$&$3$H$,$"$k(J
$@#2!KE,@Z$J%j%U%!%l%s%9?"J*$rA*Br$9$k$N$,Fq$7$$$3$H!#<+A3$N@8BV7O$G$O4D6-$,%X%F%m$J$N$G!"L\E*$H$9$k?"J*$H%j%U%!%l%s%9?"J*$,F1$8CbAG8;$rMxMQ$7$F$$$k$H$$$&J]>Z$,$J$$!#J#?t$N%j%U%!%l%s%9?"J*$r;H$C$F!"$=$N(Jd15N$@$K$"$^$j:9$,$J$$$3$H$r3NG'$9$k$H$h$$(J
$@#3!KCbAG8GDj$N:]$NF10LBN8z2L$O%j%U%!%l%s%9?"J*$K4^$^$l$F$$$k!#$7$+$7!"$"$i$?$a$FCbAG8GDj$NF10LBN8z2L$r9MN8$9$k$?$a$K$O!"(J100%$@$NCbAG$rCbAG8GDj$K$h$C$FF@$F$$$k>l9g$N?"J*$N(Jd15N$@$r:OG]
$@!zB?85AGJ,@O(J
$@B>$N0BDjF10LBNHf$HAH$_9g$o$;$k$H!"CbAG$N0BDjF10LBNHf$O$b$C$H6/NO$J@82=3XE*$J%W%m%;%9$N;XI8$H$7$F;H$($k!#(J
$@Nc!'(Jd15N$@$H(Jd18O$@$K$h$k3$MN$NC&Cb2aDx!"(Jd15N$@$H(Jd13C$@$K$h$k%(%9%A%e%"%j!<$NCbAG8GDj$N5(@aJQ2=(J

$@!z%H%l!<%5(J
$@CbAG$NJ|8:4|$,C;$$$N$G;H$$$K$/$$$,!"K;@$H%"%s%b%K%"$N$I$A$i$rMxMQ$7$F$$$k$+!"%"%s%b%K%&%`%$%*%s$rF12=$9$k$H$-$KMxMQ$9$k7PO)$NF1Dj!J%0%k%?%_%s%7%s%?!<%<7PO)$+!"%0%k%?%_%s%G%R%I%m%2%J!<%
IV Hydrogen and Oxygen Stable Isotopes
$@?eAG$K$O#3$N85AG$NF10LBN$NB8:_Hf$KHf$Y$FDc$$!J?^(J 5.1$@!K(J
$@;@AG$K$O$?$/$5$s$NF10LBN$,$"$k$,!"0BDjF10LBNHf$H$7$F$O(J18O/16O$@$,MxMQ$5$l$k$3$H$,B?$$(J

$@?eAG!";@AG$NF10LBN$NI8=`J*J$,Ds6!$7$F$$$k!K$,MxMQ$5$l$k(J

A. Fractionation of Hydrogen and Oxygen Stable Isotopes
$@!zHs@8J*E*$JMW0x(J
$@>x;6$H;g308w$K$h$k8w2rN%!#=E$$F10LBN(J2H$@$H(J18O$@$,J,JL$5$l$k(J

$@2P@1$d6b@1$NBg5$$O!"CO5e$h$j$b=E?eAG$rB?$/4^$`!#8w2rN%$K$h$C$FB?$/$N?e$,<:$o$l$?$?$a7Z$$F10LBN$,<:$o$l!"=E$$F10LBN$,;D$C$?$+$i(J

$@3$MN$NC:;@1v$N;@AG$NF10LBNHf(Jd18O$@$O!"COx;6$,@9$s"*>x;6$G(J18O$@$,J,JL$5$l$k$N$G3$MNCf$K(J18O$@$,B?$/$J$k"*3$MNCf$NC:;@1v$K$O(J18O$@$,B?$/$J$k!J29EY$H(Jd18O$@$O@5$NAj4X!K(J

$@!z9_?e$H0BDjF10LBNHf(J
$@?eAG$d;@AG$NF10LBNHf$O!"9_?e$,$I$3$G$D$/$i$l$?$+$rH?1Gar(Jn$@2mI=LL$N?e$NF10LBNHf$O!">x;6$K$h(J_$@%A$F7Z$$F10LBN$,<:$o$l$k$?$a!"(J2H$@$H(J18O$@$,B?$/$J$k!J?^(J5.10$@!K(J

$@?eAG$d;@AG$NF10LBNHf$O!"G/J?6Q5$29$H9b$$Aj4X!J?^(J5.9 $@!K"*2F$N9_?e$OE_$N9_?e$h$j$b=E$$F10LBN$,B?$$!J?^(J5.9B $@!K(J

$@!z?"J*$K$h$kF10LBN$NJ,JL(J
$@?"J*$N?eAG$NF10LBNHf(JdD$@$O!"

$@%9%?!<%A$O%;%k%m!<%9$h$j(JD$@$,>/$J$/!"(JNADPH$@$+$i?eAG$r/$J$$!#;iKC;@$N?eAG$N6!5k8;$O?"J*AH?%Cf$N?e"*AH?%Cf$N?e$H;iKC;@$N(JdD$@$K$O9b$$Aj4X(J

$@%7%s%W%i%9%H$N?e$O%"%]%W%i%9%H$N?e$h$j$b(JD$@$,B?$$"*%W%l%C%7%c!<%A%c%s%P$G?"J*$N?e$rCj=P$9$k2aDx$G!"%7%s%W%i%9%H$N?e$N3d9g$,B?$/$J$k$K$D$l$F(JD$@$,B?$/$J$k(J

$@MUFy:YK&JI$N?e$G$I$NDxEY=E$$F10LBN$,B?$/$J$k$+$O!">x;6$d?e>x5$$N8{G[!"Bg5$Cf$N?e$NF10LBNAH@.$K$h$C$FJQ$o$k(J

B. Physiological Processes

$@!zF10LBNHf$K$h$C$F?e$N5/8;$rD4$Y$k(J

$@:,$+$i?e$,5[<}$5$l$k$H$-$d!"LZIt$G$N?e$NE>N.$N$H$-$K$O!"F10LBN8z2L$O5/$3$i$J$$"*LZIt$N?e$NF10LBNHf$O!"$*$*$6$C$Q$K$O!"?"J*$,5[<}$7$?EZ>m$N?e$NCM$rH?1G$7$F$$$k$H9M$($F$h$$!#(J
$@2F$N9_?e$O=E?eAG$,B?$$"*2F$N9_?e$r;HMQ$7$F$$$k$+$I$&$+$rF10LBNHf$+$iD4$Y$k!J?^(J5.12$@!K(J
$@CO2$@3$MN?e$OC8?e$h$j$b=E$$F10LBN$,B?$$"*3$4_$N?"J*$,3$MN?e$HC8?e$N$I$A$i$rMxMQ$7$F$$$k$+$rD4$Y$k!J?^(J5.13$@!K(J

$@!z?eMxMQ$HF10LBNHf(J
$@>x;6$K$h$C$F7Z$$F10LBN$,<:$o$l$k"*MU$N?e$N;@AG!&?eAG$NF10LBNHf$O!"C;;~4V$N%9%1!<%k$J$i!">x;6NL$rH?1G!#$?$@$7!"F10LBNHf$O!">x;60J30$NMW0x!JEZ>m$N?e$NF10LBNHf!"%"%]%W%i%9%H$H%7%s%W%i%9%H$X$N?e$NJ,G[!K$K$b1F6A$5$l$k$N$G!"D94|$N>x;6$N;XI8$K$O$J$i$J$$(J

$@?eMxMQ8zN($N9b$$(JC4$@?"J*$O!"(JC3$@?"J*$h$j$b=E$$?eAG$NF10LBN$,B?$$!#?eMxMQ8zN($,Bg$-$$1v@8?"J*$O!"Hs1v@8?"J*$h$j$b=E$$?eAG$NF10LBN$,B?$$"+?e%9%H%l%9$,$+$+$k$H!">x;6$K$h$kF10LBN8z2L$N1F6A$,6/$/$J$C$F!"=E$$F10LBN$,?"J*$KC_@Q$9$k(J
$@$?$@$7!"(JCAM $@?"J*$OB?Fy$GFHFC$N:,7O$r;}$A!"Lk$K5$9&$r3+$1$k$?$a!"?eMxMQ8zN($,9b$/$F$b=E$$F10LBN$OB?$/$J$i$J$$$G$"$m$&(J

$@!z?eAG$N0BDjF10LBNHf$H5$8u(J
$@G/NX$N?eAG$N0BDjF10LBNHf$O!"D94|4V$N5$>]>r7o$N;XI8$H$7$F;H$o$l$F$$$k(J
$@l9g!"G/NX$N(JdD$@$OJ?6Q5$29$H9_?e$N(JdD$@$K1F6A$5$l$k"*%K%H%m%;%k%m!<%9$N(JdD$@$H@80i4|4VCf$NJ?6Q5$29$H$N4V$K$OAj4X!"%K%H%m%;%k%m!<%9$N(JdD$@$H9_?eCf$N(JdD$@$H$O(J1:1$@$NAj4X(J
$@$NCO0h$N?e$rMxMQ$7$F$$$k$H$-$K$O!"G/NX$N(JdD$@$O5$8u$N;XI8$K$O$J$i$J$$(J

$@!z(JNMR
NMR$@$O!"J,;RCf$NF10LBN$N0LCV$rHsGK2uE*$K8!=P$G$-$k$,!"?eAG$K4X$7$F$O46EY$,Dc$$$?$a!"B,Dj$,Fq$7$$(J
$@>l=jFC0[E*F10LBNJ,JL(JSNIF-NMR$@$rMxMQ$9$k$H!"E|$N5/8;!J%5%H%&%@%$%3%s$+!"%o%$%s$+!"2L=A$+!K$,FCDj$G$-$k!#$^$?!"%o%$%s$d%S!<%k$N%3%s%?%_$rFCDj$9$k$?$a$K!"%(%?%N!<%k$N?eAG$d;@AG$NF10LBNHf$,MxMQ$5$l$k!#$I$NG/Be$N%o%$%s$+$i$G$-$?E|$+$b$o$+$k$?$a!"8w9g@.@8;:$KBP$9$k5$>]MW0x$N1F6A$rD4$Y$i$l$k2DG=@-$b$"$k(J

V. Summary
$@%9%H%l%9$KBP$9$k?"J*$N@8M}3XE*$JH?1~$dE,1~$N8EE5E*$JB,Dj$O!"%9%J%C%W%7%g%C%H$K2a$.$:!"?"J*A4BN$K$D$$$FCN$k$?$a$K$O!"O+NO$,$+$+$k7+$jJV$7B,Dj$,I,MW$G$"$C$?!#0BDjF10LBNHf$O!"?"J*$N@8M}E*$JH?1~$N@Q;;CM$G$"$j!"$5$^$6$^$J%9%H%l%9$KBP$9$k1~Ez$N;XI8$H$7$FM-8z$G$"$k!#(J
$@@8J*E*!&Hs@8J*E*%W%m%;%9$G!"=E$$F10LBN$,J,JL$5$l$k!'Fs;@2=C:AG$N3H;6!J(J13CO2$@!K!"?e$N>xH/!J(J2H2O$@!K!"(JRuBisco$@!J(J13CO2$@!K!#F10LBN$NJ,JL!JB.EYO@E*!&G.NO3XE*!K$H!"F10LBNAH@.$H$N4X78$,J,$+$l$P!"?"J*$NAH?%$NF10LBNAH@.$rD4$Y$k$@$1$G!"?"J*$G@8$8$F$$$k@Q;;E*$J%W%m%;%9$rCN$k$3$H$,$G$-$k(J
$@C:AG$NF10LBNHf!'8w9g@.7PO)!J(JC3$@!"(JC4$@!"(JCAM$@!K!"?eMxMQ8zN(!"MUFb(JCO2
$@CbAG$N$NF10LBNHf!'CbAG8GDj$GF@$i$l$?CbAG$N3d9g!"CbAGMxMQ8zN((J
$@?eAG!&?e$NF10LBNHf!'?"J*$N?e$N5/8;!">x;6(J