Ternary effects on the gas exchange of isotopologues of carbon dioxidepce_

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1 nn Pln, Cll n Envonn (202) 35, o: 0./j x Tny on h g xhng o oopologu o on oxp_ GHM D. FQUH & LUCS. CENUSK h Shool o Bology, uln Nonl Unvy, Cn, CT 0200, ul BSTCT Th ny o npon on h o lon o on ox hough o, n on h lulon o h nllul onnon o on ox, now nlu n n g xhng u. Howv, h quon o on oop non n o h xhng o oxygn oopologu o on ox gno ny. H w nou quon o k h no oun. Th ny g whn h l-o vpou ol on n g, n p g on p v y n, uh h ophyll n o CO 2 lon,. W how h h ophyll n o CO 2 lon h n un n h p. Th p lo lg whn h lg n n oop opoon wn h CO 2 n h l n h n h. W how h h plly onl o h oxygn oop opoon o CO 2 n h hloopl n ohon n h lgh n n h k, wh vlu lo o qulu wh h oxygn oop opoon o w h o vpoon whn h l. Ky-wo: on oop non; gou uon. INTODUCTION Jn (974) n Cown (978) u h ll o npon, E, on h uv lux o on ox, h, on h o CO 2 lon,, o k (ngh) pon, n. Th w kn no oun n h ny quon o g xhng pulh y von C & Fquh (98). Howv, hy w no kn no oun n h quon vlop o on oop non, D 3 (Fquh, O Ly & By 982; Fquh 983; Fquh, Ehlng & Huk 989). Nvhl, h ny on u n lulon o p h hn ppl n oop oul. Fo xpl, n onln un o D 3, h vlu o nllul CO 2 onnon ounly lul ung h quon o von C & Fquh (98) h nlu h ny oon o wh w vpou gn. Coponn: G. D. Fquh. E-l: gh.quh@ nu.u.u In h pp w xn h o o quon o onon, non o nhn o oopologu o on ox h nlu h ny, n w how how h n o gnn n onon wh h oop opoon o h n vy n o h n h l. W phz ny on h lul ophyll n o CO 2 lon, n ny on h on o h oxygn oop opoon o CO 2 n lv, n h xn o whh qul wh w h. W u h ny o ou nw von o h quon o oopologu, u onou h h o n ou h. Th, n ny on uon w ully onng ou g, nly:, w vpou, h jo oopologu o on ox n h no on. Howv, n wh ollow w gno h o h oopologu on h oh, u h poly o CO 2 olul ollng wh oh CO 2 olul nglgl op wh h poly o ollng wh o w vpou olul. Th h on why h oopologu n npnnly n lly gl o lv unn. So h o ou n lly on onng h hl pon o wo ny y, on y ng, w vpou n h jo oopologu o CO 2, wh h on ng, w vpou n h no oopologu. THEOY Tny ung CO 2 lon von C & Fquh (98) p h ny quon o Jn (974) o Cown (978) pon o ol lux n lon o gn o ol on. Th quon lvn o CO 2 lux nvolv non o olul o, w vpou n on ox uh h: w + g g E () g w w wh n h ol on o CO 2 n n h nllul p n n h n, pvly, n h vg o h wo, n w h nlogou vg w vpou ol on. h ol on o nng g oh hn w vpou n CO 2. g h 202 Blkwll Pulhng L 22

2 222 G. D. Fquh & L.. Cnuk onun o uon o n CO 2 (n o CO 2 n ), n g w h o w vpou n CO 2 (n v v). ngng (): E g w w + g g w g g ( ) g g + w g w w E. Bu g w ª g [h ny uvy o w vpou/ CO 2 20 C whl h o /CO (Mo & Mon 972)], n + w, Eqn 2 pl ppoxly o g ( ) E. (3) o whh (2) E g 2g (3) E 2 g W on h oop o,, o h lon y vng Eqn 5 y Eqn 3. g E ( ) g ( ) E n ung /, /, ( + )/2, ( + )/2, g / g / α / ( ), wh h wgh onon o h ouny ly n o n (2.9 n 4.4, pvly, o 3 CO 2, n 5.9 n 8.8 o C 8 O 6 O), wh g E ( / α )( ) ( + ) / 2 E( + ) g ( ) 2 ( + ) ( ) ( + ) E α 2g α ny oon o oyng h pvou n lulon n whh w zo. Pog o h nx g pn on h nu o h non n h oop p nvolv. (6) (7) Eqn 3 n 3 ul o von C & Fquh (98). Th ny oon hv n nly on xpnlly hgh onnon o CO 2 (Boy & Kwu 20). Ioopologu o on ox W u h quon ov o h jo oopologu o CO 2 o n n pul pplon. I w w nly n n oxygn oop, hn Eqn 3 woul ppl o C 6 O 2. I w w xnng on oop, hn Eqn 3 woul ppl o 2 CO 2. W u h noon o h no oopologu o n, whh gh 3 CO 2 (o php 4 CO 2), o C 6 O 8 O, n l oonly C 6 O 7 O. Ju Jn (974) n von C & Fquh (98) gno h ll o [CO 2] n lon on npon, w h gno h o h oopologu o CO 2 on h oh. By nlogy wh Eqn : g w + E. (4) g g w w Thu, Eqn n 4 h unnl ny quon, wh no non wn h oopologu. Th, Eqn h no n o, n Eqn 4 h non n o, n h why w ll ou n ny on. gn, o goo ppoxon, g w ª g n o g ( ) E. (5) Con oop non, D 3, ung phooynh on lon Conng on oop, n h pl o oxylon lon (.. gnong y- n phoopon, n h pn o ny ophyll n), w n ln n pl y, wh + h onon ung oxylon, yplly.03 o hgh pln uo. W hn w α Δ / 3 3 wh D 3 h on oop non, n o lg, wh no uh ppoxon, h oluon oun ( )+ Δ 3, (8) wh h n o h nol pl quon (Fquh l. 982) h pln o n y ( ) n. (9) E No h α 0.8(w -w ), n o o ypl lo- ol on n o, y, 5 ol ol -, o- 2g pon o popoonl oon o.2% o, whh nglgl o pn pplon (.2% o 3 pl), n 2.4% o, whh gnnng o gnn (2.4% o 30 pl). 202 Blkwll Pulhng L, Pln, Cll n Envonn, 35, 22 23

3 Tny on CO 2 oopologu 223 Th oon n o gnn whn, o xpl, on h ophyll n o onln oop non un (Evn l. 986). Fo h on w gv h ull quon o non ( ppnx 2 o von): Δ α α V α Γ * α (0) wh, n h p (Fquh l. 982), h l wo n h 2n [..] w no Γ * n, pvly, k n h nluon h o ª.03, n pn on-o h n only u Eqn 0 whou ny ppoxon Eqn 5. In h quon, V kn n npnn vl. Howv, whn h quon u wh g xhng un o lon ( n h nx on), h npnn vl n uul o k h ollowng hng Γ * (von C & Fquh 98; V + Wng l. 2007; Bou l. 200; Tzo l. 20), o h Eqn 0 o Δ α α + Γ* α Γ* α Mophyll n o CO 2 lon (0) W n Δ 3 h non h woul ou h ophyll n, (wh un 2 ol - ), w zo, o h h uo woul opng n v onnon o [Coon onln pulon 9 Mh 202: In h on l o Eqn, h nuo w n o - G *.]: Δ 3 + α Γ* α Γ* α + α () Th ov non n g xhng uv, Δ 3 o, gvn y Eqn 0 whh n wn wh pl y - /P (wh P ng h oph pu) o whh ( ppnx 3 o l von) h n wn Δ 3 n h ov non Δ 3 o Δ Δ + α α + P 3 3 o (2) Typlly whn ung h onln on oop hnqu (Evn l. 986) o, on u, n y g xhng, n lo Δ 3 o. Th ov non u o vlu o, n h o ollow. Th ny uh ly n h lu n woul uul o l o o h ply, only ppoxly. To hv h, w now xn h hng n h o. Th hv y olvng Eqn 0 o, n ung h xpon o h on wh o 0, o, y. No h h vlu o. unhng, h, ll ho lul wh h ny oon n Eqn 3,. o lg ( ppnx 3), w on: o 2 o 2 3 o + Δ α α + (3) (3) Eqn 3 o no ollow ly o Eqn 3, u o nuvly xpln n h Duon. So P( )/ 2 P / (3) o o o n u o ny o vlu o whn only l vll o pulh un. ny oon un ou o vn o pon whn w nx on n xpn wh h lux ll n n unoupl. On h oxygn oop opoon o CO 2 n h l W un o Eqn 6 whh vl o ll oopologu o CO 2 (vou C n O oop). I w ng o olv o, w n xn h hoy unlyng on o h oop opoon o CO 2 n l. Suh on o h oxygn opoon o CO 2 (whou ny onon) w ou y Hwoo l. (998) n y oh (.g. Wlp l. 2008) nlung oulv (Cnuk l. 2004): α ( )+ [ ( + ) ]. (4) ( ) I w op h wh o, h onvnonl vlu wh 0, o + + [ ( + ) ]. + n wh h v oul o oh Δ 3 n 3 3 Δ Δ, h o n hng gnnly. o 202 Blkwll Pulhng L, Pln, Cll n Envonn, 35, I w onv o noon, wh x x/ -, wh ng h oop o o n, hn

4 224 G. D. Fquh & L.. Cnuk δo δ α + ( δ )+ (5) n δo + δ + δ δ. (6) Eqn 5 nl o Eqn 5 o Cnuk l. (2004), xp h n h 2004 pp, h oop opoon o p CO 2 u n o l CO 2; h o nl, no o, whh pon o phooynh ono. Th ny oon n - n. In δ δo 2 ( δo δ)+ ( δo + ) + whh n wll ppox y (7) Tl. E o ophyll n o ov on oop non o gh lv o nu oun P Vlu (ol CO ) 20.4 (6.7) E (ol H 2O -2 - ) 8.8 (3.5) g o H 2O (ol -2 - ) 5 g o H 2O (ol -2 - ) 0.58 (0.29) (ol ol - ) 363 (2) (ol ol - ) 357 (22) (ol ol - ) 285 (25) G* (ol ol - ) 42 (3) 0.06 (0.006) Δ 3 wh ny ( ) 25.0 (2.2) Δ 3 whou ny ( ) 24.3 (2.3) Δ 3 o ( ) 22.7 (2.0) wh ny ( 2 ol - ).39 (0.42) whou ny ( 2 ol - ) 0.92 (0.50) wh ny (ol ol - ) 257 (22) whou ny (ol ol - ) 265 (2) 2P o/ ( 2 ol - ) 0.47 (0.42) E δ δo ( δ o δ ) +. (7) g 2 MTEILS ND METHODS Th ny oon ppl o h ul o xpn o Cnuk l. (2004). Bly, n ho xpn w u h oxygn oop opoon ( 8 O) o CO 2 p y nu oun lv n h k. W u h un o h % o CO 2 n hloopl/ohon h h oplly xhng oxygn o wh hloopl/ohonl w ung k pon, n op h ul wh nlogou on w on whn h pln w phooynhzng. Th l un lo nvolv ng h ophyll n o CO 2 lon. Dl pon o h pln l, g xhng un n oop nly n oun n h l pp (Cnuk l. 2004). In o o o h nly pn n Tl, w u h 30, n h (Lngn l. 2008), lhough w ognz h h nly o h lng o vlu o y n o n h lgh o h pn nng. Th vlu o w ( + *) (Wng l. 2007), o oun o h n wn h 3 C o l pou ung phooynh n h l uv n h o h lkly u o pon, n vlu y lo n n. Th u o pon w u o hv n pou y phooynh n h glhou. Th 3 C non ung y pon,, w u o -2 (Gl l. 2008).Th * w lul δ C- Δ o-δ Cu (Wng l. 2007), wh 3 C h 3 CoCO 2 n h uv ung g xhng un n 3 C u h o lkly poy u. Th 3 C u w u o -27, on ovon o. Th hv n lul wh n whou h ny oon. Full l o h lulon gvn n h Ml n Mho on. P vlu gvn h n o gh lv, wh SD hown n pnh. Bouny ly onun w u onn o ll lv, n ho no SD hown. oun kn n glhou (Gl l. 2008). Th y pon,, w u o.5 ol CO 2-2 -, on un o k pon (Cnuk l. 2004). G* w n Shky l. (2007). Th 8 OoCO 2 n h ohon/hloopl,, hown n Fg. o un o phooynh n k pon. I w lul ung h ophyll n wh ny n Tl. Th w lul ong o Eqn 6 o h wh ny, wh o w lul ong o Eqn 5 o h whou ny. Th w hn lul o, ung n quon nlogou o Eqn 4 o h uon whn h no ny, uh o uon hough lqu. Thu, h w lul δ δ αw + ( δ w)+ w (8) wh w h 8 O onon ung uon o CO 2 hough lqu w (0.8 ), n w + w, kn, ollowng Fquh & Lloy (993), Th o w lul lly, xp h w pl wh o.th 8 O o vpov w n o CO 2 n qulu wh vpov w n Fg. w lul n h l pp (Cnuk l. 2004), u wh n onl lulon on n pol vlu o onon o w vpou ung uon n. 202 Blkwll Pulhng L, Pln, Cll n Envonn, 35, 22 23

5 Tny on CO 2 oopologu 225 o (whou ny) (wh ny) Full qulu wh 8 O o CO2 n h hoopl o ohon ( ) 8 O o CO2 n h hoopl o ohon ( ) () Phooynh () pon, low [CO 80 2 ] () pon, n [CO 2 ] k 32 k 32 k () Phooynh () pon, low [CO 2 ] () pon, n [CO 2 ] k 28 k 28 k Evpov w 8 O ( ) Evpov w 8 O ( ) Evpov w 8 O ( ) Fgu. Th 8 OoCO 2 n h hloopl ung phooynh (, ) o n h ohon ung k pon (,,, ) plo gn h p 8 O o vpov w ( ) o nu oun lv. Opn l how h 8 OoCO 2 lul whou h ny oon ( o), n lo ngl how h 8 OoCO 2 lul wh h ny oon ( ). Pon o w oh wh Cpp l. (2003) kn onon o 32 (,, ), n wh Mlv (978) kn onon o 28. Th vg CO 2 onnon ou h lv w 363 ol ol - ung phooynh (, ), 47 ol ol - ung k pon low [CO 2] (, ) n 347 ol ol - ung k pon n [CO 2] (, ). gon quon lng h ny o o o h pnl ollow: () ( , P < 0.00, n 8); () ( , P < 0.00, n ); () ( , P < 0.00, n 0); () ( , P < 0.00, n 8); () ( , P < 0.00, n ); n () ( , P < 0.00, n 0). ESULTS Mophyll n o CO 2 lon In Tl, h ul o h ny oon o hown o n xpl kn o h o Cnuk l. (2004). Th vlu o n o 0.92 o.39, n n o ol - o 5%. No h h n nl wh h ppoxon 2P o/. On h oxygn oop opoon o CO 2 n h l Th ny on o h % o CO 2 n hloopl h h oplly xhng oxygn o wh 202 Blkwll Pulhng L, Pln, Cll n Envonn, 35, hloopl w ung phooynh w ll, wh h % nng n 00%, n h 8 O opoon o CO 2 n h hloopl ng nly n qulu wh h opoon o l w h vpong (Fg., l pnl). Th w vn l ung k pon low CO 2 onnon (Fg., l pnl). Howv, h on h un n h k n [CO 2] k, hown n Fg., gh pnl. Th o vlu uh h h 8 O opoon o CO 2 n h hloopl/ohon n h k now lo o nly n qulu wh h opoon o l w h vpong. Th % o hloopl/ ohonl CO 2 n qulu wh vpov

6 226 G. D. Fquh & L.. Cnuk w lul o h lop o gon wn ny o n p w 0, 78 n 30% o phooynh, pon low [CO 2], n pon n [CO 2], pvly, ung h gu o uonl onon (Cpp l. 2003) h w u pvouly. By opon, h Mlv (978) kn onon vlu gv oponng ny o png o 7, 89 n 49%. DISCUSSION Tny ung CO 2 lon Tny now wll lh n nopo n h ow u n ol g xhng y (DC Bon 2004; Wlz 2005; PP Sy 2006; L-Co Bon 2008). Th o u u h on ox lvl. Boy & Kwu (20) hv on h h quon nlung ny ny, n n ly. Th quon o h o h xnon o oop, ng u ly h quon o h jo oopologu, n hn, wh oon o onon, quon o no oopologu. Con oop non, D 3, ung phooynh on lon Th quon o on oop non whn o o ny o w vpou gn how n n n h ovll non o gvn o CO 2 onnon nng o oph, o h l u, own o h n- n nlly nllul. Th puly l h uo lv hn h hvy on oopologu, ung h ul-up o hvy oopologu n h oluon n p whn h l, whh hn u y h wp (lo onvv) o CO 2 ou o h l y h npon lux. W y lo onvv onn h g w ª g, kng Eqn 3 hv h o o pl onvon o po y Cown (978) puly no hvng n h gnl ny n pulh y Jn (974). W no h ny on on oop non y C 4 p, Δ 3 4, y nlu n n quon nlogou o Eqn 0 n p o Eqn 9 o Fquh (983) Δ + p p p φ p p p +, p φ p + p p p (9) wh 4 h v onon y PEP oxyl, h popoon o CO 2 x y PEP oxyl h uqunly lk ou o h unl hh, 3 h onon y uo, p n p h pl pu o CO 2 n h unl hh n n h ophyll yool, pvly. In h o lg pco 2 n h unl hh, p, h uul plon y n h l. Slly, loon o k no oun y pon, phoopon, o nopl qulon o CO 2 n HCO 3- n h ophyll y lo nlu y Fquh (983). Mophyll n o CO 2 lon To ough ppoxon, h ol vlu o n o n y ª2P /, wh n h o h lul whou h ny oon n h quon o Δ 3 (Eqn ) n o D 3 (Eqn 0). Th y unoo nuvly o npon o Eqn 0: whn h oul o ull o ov non hng o h wh 0, o h wh ull ny oon, h on y oughly 2 popoonl n n h on k, h ppox y /. Th n h o kp h vlu o non onv ( h ov vlu), h nw oul qu n oppong popoonl o 2 n, h, n olu op o 2. Wh x vlu o, h qu 2 popoonl op n / n hn n olu n n ( - )P/ o 2P /, h, n n n o 2P /. Th olu n o ol - n (2P /) (Tl ), whn ny oon nou o h on oop quon n ppl o h o Cnuk l. (2004), wll no unuul whn op wh oh o o, h vlu o opon o.8 kp l-o- vpou pu n, lhough h vlu o w h lg o 2004 vlu o. Th oon wll g lg lo- vpou pu n (hgh ) n hgh CO 2 onnon, n n pln wh low phooynh p (low / ). Th oon popoon o wll g n pln wh low.thpng n n h pn (5%) wll unuul, ol - o n unuully ll (uno) n (. von C & Evn 99), lhough poly h o. oun. Th vl ho o ng ophyll n, o nv, ophyll onun, g (Loo l. 992). Th pn uy ugg h h oop ho h un h n, h, ov g. In h onx, nng h h oop o g (/ ) y Vál l. (2009) gv 26% hgh vlu hn h ngl pon vl J ho. Vál l. u vlu o 7.5 ol ol - o w -w, o h w Th oon o h Cnuk l. (2004) vlu (h uno vlu o g ng 5% lg hn h o on) o o 0.06 (Tl ), hppn, poly onnlly, o oughly opl wh h o Vál l. n h n h (0.006/0.06)5% 9%. Th nng y Vál l. n on o h l ovon y Loo l. (992), u n gn wh ho o Flx l. (2007). Th l xn g n 202 Blkwll Pulhng L, Pln, Cll n Envonn, 35, 22 23

7 Tny on CO 2 oopologu 227 op hln n n ol pln w 22% hgh whn n y h oop ho (Evn l. 986) hn whn n y hloophyll luon ho (Hly l. 992). Nvhl, h ny ou o o n ll uh hnqu nvolvng g xhng, n o n h un o l pu ll poly h o oon n gnn (Moow & Sly 97; Mo & Pk 20). Bu h ny oon o pn on, n, oul ponlly p upon ppn pon o o l-o- vpou pu n, n, n l pu, o xpl, whn n y on oop non. To xn on h ppn pon o o, w ppl h ny oon o h pov n h upplny l o Tzo l. (20). H, h n n wh h ny oon ng o 0.05 o ol -. Ovll, h w only ll popoonl n n (3 o 8%), u h un w lvly low l-o- vpou pu n (5 o 9 ) n h whou ny oon w lvly hgh (.66 o ol - ). Thu, h popoonl p o h ny oon on on wll vy ong xpn, pnng on h xpnl onon n pln l un nvgon. P onl pon o h oon uz n Eqn 3. On h oxygn oop opoon o CO 2 n h l Th puzzl ou h l ul (Cnuk l. 2004) w h h popoon o CO 2 olul h h qul wh w n h ohon/hloopl pp o ll ung k pon n [CO 2] hn ung phooynh o k pon low [CO 2] ( Fg. ul wh no ny oon). Th ul pn h ugg h h popoon qulu w only lo o uny n oh lgh n k. Th lul ovll lonhp wn h oxygn oop opoon o CO 2 n h hloopl/ohon n h o h w n h ognll nv o h vlu o h kn onon o u n ng h oop opoon o w n h l. W u h vlu o Cpp l. (2003) n h upp pnl o Fg. ( Cnuk l. 2004) u h o onovl (Luz l. 2009). Wh h low vlu o Mlv (978) o Bkn & Luz (2007) (low pnl o Fg. ), h lul w oop nhn ll. Th k h CO 2 pp lo o ng 00% qul wh h w ung phooynh (pnl D), ung k pon low [CO 2] (pnl E) n ung k pon n [CO 2] o l w nhn o 8 o l (pnl F). No h h l pnl (B n E) how uh l ny oon hn h oh h oop opoon n h lv n n h n w vy l o pnl B n E, h ng o h nng h h o h ul w o CO Blkwll Pulhng L, Pln, Cll n Envonn, 35, Tny oon n o ou? I nowohy h o o pupo, h oon o ny on h vou quon o lux o oopologu,. o no n o png uh p, pov h h g xhng quon o E n lo o ny oon. Fo xpl, lulon o wh ny xlu o oh h g xhng quon n h quon o Δ 3 y only ol - o h lul wh ny nlu n oh o quon o h hown n Tl. Howv, whn on h h oon ( yplly now h o o g xhng un), wll ny o uy (o l o onny) o u h ny oon o h CO 2 oopologu pvouly. W o no wh o gv h pon h on houl lv ou h ny n nol g xhng lulon, uh oon vlul n ng h pl pu n y h oxyl, o xpl. W hv no u h ny on lux o w vpou oopologu, hy o opl n wll l wh n p uy. CONCLUSION Th xhng o on ox oopologu o w vpou n nvolv ou p g (, w vpou, lgh CO 2 n hvy CO 2). G xhng quon hv o o nlu h ny o npon on CO 2 uon hough o. Howv, h n quon o on oop non hv gno uh. W nlu h n how h hy g whn l-o- w vpou ol on lg. Thy n hv lg on p y ul n wn wo po. On xpl w hv xplo h ophyll n, lul ung oop hnqu h n wn hol (n h n o ophyll n) n ul on oop non, kng u o vl n y g xhng. Howv, w ugg h oh p h hv n n y ng ovon o hoy, uh h onon o wh phoopon,, n y pon,, wll lo n o. Lg o ny oon lo ou whn h lg n n oop opoon wn h CO 2 n h l n h n h. Th onl n o h oxygn oop opoon o CO 2 n hloopl u o l o h n ohon n h k n lvl o CO 2, n ough oughly n ln wh un n h lgh, n n h k low [CO 2], h, n n qulu wh w h o vpoon n h l. Th ul pn owh on h vlu u o h kn onon o w oopologu.

8 228 G. D. Fquh & L.. Cnuk CKNOWLEDGMENTS W hnk h uln h Counl o uppo (DP o GDF n FT o LC). W lo hnk h nonyou vw. EFEENCES DC Bon (2004) LC Pol Phooynh Sy: Inuon Mnul. DC BoSn L., Hoon, UK. Bou M.M., Wn C.., Fquh G.D., Fo G. & Bown H. (200) Vly n ophyll onun wn ly gnoyp, n on npon ny n on oop non. Pln, Cll & Envonn 33, Bkn E. & Luz B. (2007) Duvy onon o H 6 2 O/ H 7 2 O n H 6 2 O/H 8 2 O n n h plon o oop hyology. p Counon n M Spoy 2, Boy J.S. & Kwu Y. (20) Phooynh g xhng y wh nnl CO 2 ly u. Envonn Conol Bology 49, von C S. & Evn J.. (99) Dnon o h vg pl pu o CO 2 n hloopl o lv o vl C 3 pln. uln Jounl o Pln Phyology 8, von C S. & Fquh G.D. (98) So lonhp wn h ohy o phooynh n h g xhng o lv. Pln 53, Cpp C.D., Hnk M.B., DPulo D.J. & Cohn.C. (2003) Ioop onon o w ung vpoon. Jounl o Gophyl h 08, Cnuk L.., Fquh G.D., Wong S.C. & Su-Wll H. (2004) Mun n npon o h oxygn oop opoon o on ox p y lv n h k. Pln Phyology 36, Cown I.. (978) Sol hvou n nvonn. vn n Bonl h 4, Evn J.., Shky T.D., By J.. & Fquh G.D. (986) Con oop non u onunly wh g xhng o nvg CO 2 uon n lv o hgh pln. uln Jounl o Pln Phyology 3, Fquh G.D. (983) On h nu o on oop non n C 4 p. uln Jounl o Pln Phyology 0, Fquh G.D. & Lloy J. (993) Con n oxygn oop on h xhng o on ox wn pln n h oph. In Sl Ioop n Pln Con/W lon ( J.. Ehlng,.E. Hll & G.D. Fquh), pp P, Sn Dgo. Fquh G.D., O Ly M.H. & By J.. (982) On h lonhp wn on oop non n h nllul on ox onnon n lv. uln Jounl o Pln Phyology 9, Fquh G.D., Ehlng J.. & Huk K.T. (989) Con oop non n phooynh. nnul vw o Pln Phyology n Pln Molul Bology 40, Flx J., Ouño M.F., -Co M., Dz-Epjo., Floz-S I.D. & Mno H. (2007) Mophyll onun o CO 2 n op hln. Nw Phyolog 75, Gl., Thkz G., Puk.D., Ghhgh J. & Fquh G.D. (2008) Expnl vn o l von o h on oop opoon n l, n phlo p ogn n nu oun. Pln, Cll & Envonn 3, Hly P.C., Loo F., Do G. & Shky T.D. (992) Thol onon whn ng h ophyll onun o CO 2 lux y nly o h pon o phooynh o CO 2. Pln Phyology 98, Hwoo K.G., Gllon J.S., Gh H. & Boow M.S.J. (998) Dunl von o D 3 CO 2, DC 8 O 6 O, n vpov nhn o H 2 8 O n Pp unu un l onon n Tn. Pln, Cll & Envonn 2, Jn P.D. (974) Th uon o on ox n w vpou hough o. Jounl o Expnl Bony 25, Lngn G.J., Bon N., Gh H. & S U. (2008) Con oop onon ung phoopon n oxylon n Sno. Pln Phyology 48, LI-CO Bon (2008) Ung h LI-6400 Pol Phooynh Sy. LI-CO Bon, In., Lnoln, NE. Loo F., Hly P.C., Do G. & Shky T.D. (992) Eon o ophyll onun o CO 2 lux y h n ho. Pln Phyology 98, Luz B., Bkn E., Y. & Shh. (2009) Fonon o oxygn n hyogn oop n vpong w. Goh Cooh 73, Mo T.. & Mon E.. (972) Gou uon on. Jounl o Phyl n Chl n D, 3 8. Mlv L.J. (978) Molul uv o H2M 6 0, HD 6 0, n H2 8 0 n g. Jounl o Chl Phy 69, Moow P.. & Sly.O. (97) L pu on un o uv n o w vpo n. Pln Phyology 47, Mo K.. & Pk D. (20) lnv ppv on h onol o npon y on. Pong o h Nonl y o Sn o h Un S o 08, PP Sy (2006) TPS-2 Pol Phooynh Sy: Opo Mnul. PP Sy In., Hhn, UK. Shky T.D., Bnh C.J., Fquh G.D. & Sng E.L. (2007) Fng phooynh on ox pon uv o C 3 lv. Pln, Cll & Envonn 30, Tzo Y., von C S., Evllo G.M. & Evn J.. (20) Ung unl o l poopy o u on oop non n ophyll onun o CO 2 uon ynlly n CO 2 onnon. Pln, Cll & Envonn 34, Vál D., Všková M., Honková M., Flx J. & Šnůčk J. (2009) Mophyll onun o CO 2 npo y wo npnn ho: o vl CO 2 onnon n. Jounl o Expnl Bony 60, Wlz (2005) Pol G Exhng Fluon Sy GFS- 3000: Hnook o Opon. Hnz Wlz GH, Elh, Gny. Wlp L.., L X., K K., G T.J., Bllk K.. & Bk J.M. (2008) 8 O o w vpou, vponpon n h o l w vpoon n oyn nopy. Pln, Cll n Envonn 3, Wng L., S U., Mon J.B., Jv P.G. & Lloy J. (2007) Von n 3 C non ung CO 2 xhng y P hn nh n h l. Pln, Cll & Envonn 30, v 2 D 20; v n v o 23 Jnuy 202; p o pulon 23 Jnuy Blkwll Pulhng L, Pln, Cll n Envonn, 35, 22 23

9 Tny on CO 2 oopologu 229 PPENDIX L o yol ol on o nng g oh hn w vpou n CO 2 wgh onon o h ouny ly n o n onon o h ouny ly onon o h o 3 C onon o nlung oluon n uon n lqu w 8 O onon o nlung oluon n uon n lqu o CO 2 lon o lon o h no oopologu o CO 2 C oop onon ung oxylon n C 3 pln (nly, u no olly, uo) 3 C oop onon y uo 4 v onon gn 3 CnCO 2 y PEP oxyl ol on o CO 2 n h n quvln ol on o CO 2 h o oxylon n h hloopl o lul quvln ol on o CO 2 h o oxylon n h hloopl n h n o ny oon ol on o CO 2 n h nllul p ol on o CO 2 h l u vg o n ol on o h no oopologu o CO 2 o n n h n ol on o h no oopologu o CO 2 o n n h nllul p vg o n onon ung y pon (oxylon) onon ung phoopon (oxylon) E npon g onun o uon o n CO 2 (n o CO 2 n ) g onun o uon o n h no oopologu o CO 2 (n o h no oopologu o CO 2 n ) g w onun o uon o w vpou n CO 2 (n v v) g w onun o uon o w vpou n h no oopologu o CO 2 (n v v). g ophyll onun o uon o CO 2 (/ ) p n pl pu o CO 2 p pl pu o CO 2 n h nllul p p pl pu o CO 2 h o PEP oxylon n h ophyll ll yopl o C 4 pln p pl pu o CO 2 h o ubp oxylon n h unl hh ll o C 4 pln P oph pu ophyll n o uon o CO 2 (/g ) o lul ophyll n o uon o CO 2 gnong ny oon o oop quon oop o o h CO 2 n h n oop o o h lon y pon oop o o h CO 2 n h nllul p o lul oop o o h CO 2 n h nllul p n h n o ny oon o h oop quon n k (ngh) pon C oop onon o wh lkg o nogn on o h unl hh o h ophyll ll o C 4 l ny oon on ( Eqn 7) V o uo oxylon w ol on o w vpou n w ol on o w vpou n nllul p w vg o w n w onon o o h oopologu o CO 2 ung n onon o o C 3 (nly uo) oxylon onon o o y pon (oxylon) wh p o n lon onon o o phoopon (oxylon) wh p o n lon 3 on oop onon o oop opoon n x x/ -, wh ng h oop o o n oop opoon o CO 2 n h n oop opoon o CO 2 h ng l oop opoon o CO 2 h ng p ngh oop opoon o CO 2 h o oxylon n h hloopl oop opoon o CO 2 n h nllul p o lul oop opoon o CO 2 n h nllul p gnong ny oon D 3 on oop non (oly u o png C oop non y C 3 pln) D 3 4 C oop non y C 4 pln D 3 on oop non y C 3 pln w zo Δ 3 o ov on oop non G* CO 2 phoo-opnon pon popoon o CO 2 x y PEP oxyl h uqunly lk ou o h unl hh o C 4 pln 202 Blkwll Pulhng L, Pln, Cll n Envonn, 35, 22 23

10 230 G. D. Fquh & L.. Cnuk PPENDIX 2 Dvon o Eqn 0, h ny o quon o on oop non W hv h ny o quon lng h oop o o lon hough ouny ly n o o h o h CO 2 n h n n n h nllul p,. ( + ). α ( ) ( + ) (6) Ou gy () o u quon o lqu ph CO 2 uon o h ophyll n,, o l o, h on oop o o h on ox h o oxylon, xp n o n quvln g ph ol on,, whh h quvln pl pu p, v y h oph pu, P. Thn () o u quon o ol on o l o. Fnlly, w () u h non o non o on Eqn 0. α () Δ / ( non) 3 3 Th o lon o 2 CO 2,, gvn y P p n h o 3 CO 2,, gvn y P p α wh + h onon o wh oluon o CO 2 n uqun uon. hn gvn y /, o h. () α ( ) () Th o lon o 2 CO 2 qul h o ubp oxylon, V, nu h o phoopoy CO 2 lo, F, nu h o y pon,. V F Th o lon o 3 CO 2,, gvn y h oponng 3 C lux V F V wh V α ; F /α ; /αn + F h uo onon, + h onon y phoopon wh p o h n lon, n + h onon y y pon, gn wh p o h n lon. Fo h quon V F α α α V F n y vng houghou y V n olvng o kng F u o Γ * / (von C & Fquh 98) n V o n w on α α α α * α Γ α α. () α α V Nx w uu o n () ung (), n hy olv o * ( )+ α Γ α α α. α α V Th xpon o nx uu ( wo pl) n h nuo o Eqn 6 ov, lvng only wo oop o, n whh p { α( ) ( + )+ α * α ( ) α( )+ Γ α α α V } ( ). () pplyng h o ou non α Δ / 3 3 w on 3 α ( ) Δ + α ( )+ αγ * α α α αv n ng h α ( ) wh h wgh onon o h ouny ly n o n, w n w +. plng y + n, ppn, y +, w ln o ngn yl Δ α α V α Γ *. α () (0) W no h n h on l o Eqn 0, Γ * (von C & Fquh 98). In h V + onx o ng, h long o k o n (Wng l. 2007; Lngn l. 2008; Bou l. 200; Tzo l. 20) h vlu o onn o wh ov, n h o oxylon V o un h 202 Blkwll Pulhng L, Pln, Cll n Envonn, 35, 22 23

11 Tny on CO 2 oopologu 23 vlu pn on h vlu o p n hn on h o. So Eqn 0 o Δ α α + Γ* α Γ*. α PPENDIX 3 Dvon o Eqn 3, h ny o quon o ophyll (wll) n n o p, h p(co 2) h o oxylon Ou ppoh o uu (0) / P () n Eqn 0 n hn olv o.to on h oluon, uul o n D h vlu whn zo, h,. Thu + * α Γ α Γ * α + α Δ o o p ( Δo Δ 3 ) α α + Th vlu o x h o o, h uno vlu, D > D o, Th x gvn y ( Δ 3 Δ ) ( Δ 3 Δ ) p α α + p 2 3 Δ Δo + Δ o. + α α + o o o o n uung h xpon o D n D o n nllng w l wh. + α Γ* α Γ* α + α Δ n o Eqn 0 o 3 Δ Δ + + α. α + Ung () Δ o 3 + p α α + 2 o + 3 Δ p α α + o o whh uul ough ppoxon Δ Δ + α P α + 3 n o h ophyll n gvn y p ( Δ Δ 3 ). α α + wh p P. In n xpnl uon, u h D 3 goo pnon o h ov non, Δ 3 o. No h n opng h ul wh pvou on, h uno vlu o, o, pn on h uno vlu o D, no D o. 2 p o (3) wh p o h uno o p. n o h l n wn o n vlu, o, whn o zo, (o () ov) o 2 3 o + Δ α α + n h oponng ough ppoxon (3) o 2 o. (3) 202 Blkwll Pulhng L, Pln, Cll n Envonn, 35, 22 23