Integration of Carbon and Nitrogen Metabolism

8/3/2019 Integration of Carbon and Nitrogen Metabolism

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58 HUPPE TURPIN

he Algae . 596Th Photosynthtic Hihr Pnt C. 597eerorophi iue 598

CONCLUDNG REMARKS 599

INTRODUCTION

Th meai pahay r a ad irg ume h grear par he phyhei e ar ad eergy reure p 55% e paar i mied irge aimiai ad meaim i me iueCar ad irge meaim are ed eaue hey mu hare rgai ad eery uppied direy rm phyhei ee apad C0 ai, r rm repirai d a ia gyyi, heriarxyi aid (TCA ye ad he mihdria eer rapr haimiETC The iegrai hee impa meai pree muie exeie reuai eee he pahay

hi re ue he ieme ar aimiai ad repirai i pmay m f trg d, cry th ct nrge aimiai he rae ad rue ar meaim i he e

May rerie urrudi eiher pahay i e umaied h

h ur urre uderadig ih h reuireme meai ierai We hae reried hi reie he aimiai e irge irgai maer i phyhei euary

The priary aiain igai irge in amn aid euirear ee i he rm ead ad eery i he rm A T adrdua Figur a I h phyhi ad hrrphi he ai are iemeiae repiray meaim 76 Repirai reda/r aae phya pvie ery i hepi herea phyhei uuay uppie eergy i phyhei e (3 8, I a e a ireaed rae primary irge aimai reuire

Abbreviaton 20G: 2oxolutarate; 3-GA: 3-phospholcre; Asp: apaa DM:3-(3.4-dichoopheny).dmethylurea DHP dhydoxyacetone phosphate 26P fructose,6-bisphosphate 6P fructose 6-phosphate FBP fctose 1,6bisphosphate BPase fctose16bphosphaase F eeoi FNR Fd-NADP odoeductase G6P glucose 6-phosphateG6PDH luos 6phophate ddros; GAPDH rdd 3phosphate ehyogenase GIn gutamne Gu gutamate GGAT glutamine 2-oxoglutaae amioanseae GS:glutamne synthetase IH soctate dehyrogenase MDH maate ehyrogenae mETmitochonal eetn rpo han NH4+ mmoium NR nre reuae N0 nrateNR rte ece OAA oxoacette OP: oxtve peoe phophte PD pyruvaeehyroee compex PP phosphoeopyruve PPcae: phophoenopyrvae car boxyase F phophouoknas PF pyophosphate ftoe6phophate 1-phosphoanese noranc phospha P pyruve kae yr: pyuvae Rubso uoseI,bsphophe crboxye RuBP: boe bphosphae SPS ucre phophte syhete TCA: crboxyc ac oe popha.

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NTERTN N N ET 59

sme nceased w carn thruh respratry pathways whch aects themetasm the whe ce (8 )

AN APPROACH TO STUDY

he pneern studes carn and ntren nteractns were cared ut ySett (1511), Bassham, Kanaawa and cwes (, 0, 1) pmarusn unceua aae. n eay seatn that s centra t the understandin and esearch ths pem was that ntrensucent aae requediht t assmate ntren, ut ntrenimited ces rapid assiated ntr en in the dar (, 5, 13) ntaceua carhdate stes deceasedduin da assimiatn and assiatin ceased when these stes wee de peted (15, 10). Exenusy supped carhydrate cud as vercmedar nhtn (39, 15). he at t induce ntren assmatn n thedar awed phsica studies this prcess n the asence the cm pexities phtsnthetc as exchane urther, t ndcated tat carhdrate respiratn s mprtant t the assmatn ntren

he use uniceuar aae emains centa t u undestandn hwrespiratn, phtsynthess, and ntren assimiatin nteract. Aae can erwn as a hmeneus ce ppuatin in chemstats (1) where nutrient

avaiait sets rwth rates s that nerm and shrtterm eects dier n medum cmpstn can e cmpared. he rapd unrm respnse aa ces t chanes such as ncreased avaait nranc ntrenaws detemnatn shrte uxes that hhht the enymes andreuaty actrs cntrn the respnse. pecc enymes reuate meta c pathwas he nhtn r actvatn these euatry enymes rad ca chane the ux in the whe pathwa Determnatin sustrate/pd uct ats ca e used t dent euat enmes minn in vitrstudies indvidua enmes with eaminatin hw metaites and en

me eectrs chane durn the activatn a metaic pathwa prvdesan indicatin what actrs sustan the actvty that pathway.

Mutceart and an derentatn adds cmpext t the stud can and ntren nteractns n hher pants ( 113) Man asc questns aut nitren metaism in hiher pants are sti answered cmpedescrptins Pants can assmiate th ntrate (N0) r ammnium (NH4+

he ste assmatn and tpe ntren used vares dua, seasnay,ecca, devepmenta, and wth speces (3, 11). Pants as nitr en thuh the rts, and mst i nt a NH+ ased s assmated n therts ( ), whereas much the N0 s transprted t and assimiated n theshts (3 15). Sme pants assiate ntren n the dar in th ts andhts (3 1 133) Mst pants stre are amunts N0 n eavacues s the nten avaiae r assimiatn n a ce can exceed the

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80

amunt avaiab fr uptak frm th nvirnmnt (5 inc nitrn i ambi nurin pan ar ab mv ntn nt n rth by ncindr tiu. Finay hih C rati quat t nitrn mtabim actin

crbn mtabim in hihr pant han in aa (65. h nurint tau a hihr pan chan radua hich mk i dificu dmin hrdr in hich nzm bcm acivatd (in cntr ntad mphaiinadutmnt f nzym cnnt car cntr. Ln-trm diffrnc inmtabit v a affctd b nutrint tau can faciiat idntificatin fruatry it imprtant in th imutanu mtabim f carbn and nitrn in bth hhr pant and aa

HSOLOGCA EVDENCE OF NTACTON

Ga xchan maurmnt rct th ct f nitrn n carbn Additin f a nitrn urc t imitd r tarvd aa c in th darkincra piratry C ra dmntratin th activatin f rpiratrycarbn f (58, 77, 79 h hihr rductant dmand f aimiation requires that moe cabon be mobilized to assimiate vs Thiscau highr raiv ra C0 ra dung N03- aimain hanhn h H i h nirgn rc (7

h diffrin rductant dmand f H and a ad t diffrntiacupin btn th CA cyc and th mtEC. Durin H aimiatinin aa cnumptin and C vutin incra prprtina hinthat rpiratr carbn fr amin acid ynthi i cupd t th mitECand i th rpirar ctrn accptr(77, 79 n cnrat uptak iny ihty hihr durin N0 aimiatin dpit th arr incra inC ra hi rut ndcat hat m rpiratry ctrn f t rathr than t th mitEC (79 n br rt th rpiraty qutintincra in rpn but nt t H hich rct a ar drp in0 upak and i chan in C0 ra Ovra thi rpn hthat th ruirmnt fr H and aimiain diffr imiar in pantand aa athuh th nry paritinin ma vary mha. ira aimiatin by bary ht a rut in an incrad rpiratry qutint hichh that a inificant prtin f ithr rpiraty r phtynthtic ctrn i cupd t rductin in hhr pant ( h r f a aink fr rpiratr ctrn i a indicatd by th r inhibitin f bthrpiratr carbn and nitrn aimiatin brvd hn v i th nitrn urc undr aarbic cnditin (7

tudi in -imitd aa durin th acivatin f rpiratin fr aminacid ynthi d n uppr a r fr th atativ xida in uppyincrbn fr binhi h cchrm ida paha mut cuph nirgn-imua espiatoy n the mitETC in



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RAI AD N METAIM 1

mnuum cus this lg cs lt pthwy ctivity (178. n Clmy domon end, dspit its high ltt pthwy cpcity 8 discrimintion dmonstrtd no ltrntiv oids ctivity during H+ ssimi

ltion (76.Rspirtor rls nd consumption cn diffrntitd during

photosynthsis using mmrn inlt mss sctromtr nd mdi contining 8 nd (15. n gl cls, th rspirtory rspos is similrduring photosynthtic nd dr nitrogn ssimiltion (77, 79. Dspit stimulting grtr rls of thn H ssiiltion, ssimiltioncuss onl slight incrs in rspiror consumption (9. n imitd minuum rsirtor cron flow provids con for mino cid synthsisduring photosynthis s wl s som of th rductnt for 0 ssimiltion

(179.itrognsufficint g do not ccumult lg crohdrt rsrvs

(.g. tah); herefore, he suppl of cabon o amino acid synhesis is dpndnt on cnt hotosyntht ( 16. Withholding ithr light or inhiitsnitrogn ssimiltion (, 9, 16, 163, 169. Inhiitors of photosynthtic lctron trnsport nd cron fition [.g. 3-(34-dichlorophnyl1,1-imthyl ur (DU or D-glycrldhyd rspctivly] ic this ct(, 63. f clls providd with mtolil sourc of orgnic cron(.g. gucos, ctt, or intl crohydr (39 6 nitrogn ssimiltion com indpndnt of rcnt photosyntht (, 161.

itrogn ssimiltion y highr plnts nd lg is oftn ccopnid stimultion of volution cus nitrogn mtoism provids nothrsin for th nrg hrvstd in photosnthsis (, 9, . sit thincrsd volution, th rt of fiion is somtims supprssdduring nitrogn ssimilion, spcily t nonsturting light nnsitis (4,53, 94 13. In som -imitd lg, this photosynthtic supprssion isdrmtic vn t sturting light (44 nd strch is moilid to provid cron

(1 6

ARBON RQURMNTS FOR NITROGNSSIMITION

Cohydrts fmd i photosynthtic fition r ithr trnsportdfrom th sit of ssimiltion to othr pts of th plnt or usd in th chloroplst to supply nrgy production, ioynthsis, or storg. uch wor incron tois s focusd on undrstnding th lloction of cronwithin th cll, nd twn th sourc nd sin tissus of plnts (3, 151. Amjor dmnd on th cron ssimiltd in th chlooplsts is th synthss ofmino cids rom inorgnic nitrogn. Whn th rt of nitrogn mtoism isrducd y nitrogn limittion or y dcrsing th mount of nitrt rducts

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582 HPPE & TRPIN

(NR), carbon storage compounds accumlate nd photoynthetic eciency

my be educed (51 52, 132). Activaton of nitroen assimlation conumes

TCA cycle ntemediates wic must be relced by anpleurotc C02 fx-

Sur

ytosol c ch pasid

'PieI l ! A i fc)1P r---.-, G6OH

/ .. ---------' : l ox: b) ,F . , imn-: ( ' ',___IU,= -.' 1: L-'"T1 tFB: " li.-.I jDHAP tDA .

.

GA

!E

NAD}H

Gn r +!EtPi ND()fGlu @ a tDHAP

,2-0G Glusp w yrMa W . N02

Mal_A.Asp "fam iy Aa fmily

mitochondron GIUfamilY}+G II gcyCo

(

O A A --:l -.. _+- tnFigure 1a The conol of cbon partitiog o prmay irogen assimaon. This gure diaeshe geeal feaues assoaed with h povsio o ab ro sth, suos ad rt

phoosynhae o he sythess o amo aids Te sgnfa ah sre depeds o the sse

n qestio ad is develpea ad physiologial saus Th raos boxed by dashed li ddesigated ad c ae detaed i Fgures Ib ad Smal ows pdng a abot datewhher t s an incease (i) or decas () i h lvl of the metabolie Abbreatio/symbolae n ext xcept tyl-Co: atyloezye A; Ma aat ativae 0 ; deaat0 ;hoshoylated, nt phsphylated; .






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TERAT AD MEAIM 83

tion Fiu a Th ulation of abon ow as fftd b niton assimilaion ovids nst into how abon dmands a baland in t ll

Photosynthetic Carbon Fixationnt hotonthat ud n amno ad nthss 2 8) f abohdato a aalabl, how, nton an b amilatd b lnt and alallls in t an ofC02 n faC02 faton s sud n omlls duin nion assimilation Altou otd in som ass fo ilans 51, 1) t most damati and wll-studid amls of this do inC02 fiation a in ala wh h oima aus is t limitaion ofblos 1-ishosat abolas ubiso) b ibulos 1, -bishoha

uB) 42, 44) Th ulato mhanims a unown, b dlin isolatd wt th ont of tah mobilaton 146) and ma nomatblt bwn abon dution and siaion in th hloolast Tusion n hotosnthti mtabolism is tanint, lastin until th niton assimilaion as dlin o t avaiabl stah ss a dltd 16 161)

th nt o sod hotonthat od abon fo amno adsntsis dnds on th aai of otosntsis and t aailabili ofstod abodat 11) Fo aml ivn th otosnthti 0 fiation

at and aio of otin in N-limitd S. mnuum abon fom hotosnthsis would susain a at of N+ assimilation of mol m ChI h

Q

iOQN

+EGA

tF6P

E(A

BP.--.

Fgur b Inhton o cytosoc guconeogenes and acvaton o gycoys: a he ucose6 pospae (F6P)/uctose 16-bsphosphate (B inteconveson dung ntogen assmlaonAbbevaon e as n the et ecept 26PK ucose2,6bsphospate knase; 26Pasefrctose ,6-bisphshaase

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58 HUPP TURP

Without nitrogn, illumintd clls ccumult strch f th H+ ssimiltion rt incrss, h rt of stc ccumultion dcrss Whn th dmnds for cron y nitrogn ssimilion cd th rt of hotosynthtic

C ition strch dgrdtion is osrvd in th light vidnc oth intgrtion of cron nd nitrogn ssimiltion hs lso n rortd inhighr nts. oyn lnts grown t thr nitrogn lvs 3 nd 18 rtitiond lss hotosyntht to strh s th nitrogn suly incrsd 12.Th css cron ccumultd in th first dy tht soyn lnts wdrivd of ll nitrogn quld th mount o cron ncssry to suort thdily rt of nitrogn ssimiltion in thos lnts 132.

n th d, stord crohydrt must rovid cron for mino cid synthsis 81 t hs n dmonsttd in S. minutum tht th stoichiomtrytwn th rt o strch dgrdton nd th rt of H+ ssilton isconsistnt with th C rtio in this lg Clls moiliz strch t highr rtduring ssilton his lrg cron dmnd is rctd y th highrrt of rsirtory C rls whn ssimltion rthr thn H isth nitrogn sourc 17

h t of wht oots to ssmt nton s otd drt wththir ohdt stts, nd th ron stons fo nitogn ssimitionr drvd from th stord cron 28 h rovision of ognous cron to

lvs, roots, nd lg in th dr significntly stimults th rt ofitrognssimiltion 7 13 Th grtr vilility of crohydrt in th lightccouns, in rt, for th diul cycl of nitrogn ssimiltion in oth htrotrohic nd hotosynthtic tissus 133. This indicts gnrl rltionshi

AD()

OS

6PG

6x

@v

G6r

.

ADH

AD

i A aaac repesenan he cascade nvlve n he hrexn acvatn fhe ave pentse phsphate pahway prve reucn pwe r N0 reucn Abbrevn e a in he text exce x edx; r edced; x xdzed






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NRAN AN N MABM

eteen arodrate tat and te differing dark aimilation rate reportedin plant

Anapleurotic Carbon Fixationtudie of dark C0 fiation in plant o tat anapleroti aron i required to replae TCA le intemeiate onumed i ioei Teonet of nitrogen aimilation in ot iger plant tie 8 5, 15 and algalell 8, 15, 171 relt in a large timlation of dark C0 fiaton ere ia liear relationip etee te rate of itroge aimilation ad anaplerotiaron fiation 5, 171 In mnuum 3 mol aron i fied per molntrogen aimilated, a toiiometr oepondng ell to te reqired anapleroti HC0 aton allated from te amno ad ompoition of totalelllar proten (11 If anapleurt reation are loked trngentC0C0 removal dark H aimilatio top (2 Gu et al avedemaed he aiai f aapeui a fixai popoeopruvate arolae (ae during pototeti nitroge aimilatioing te differential dirimination Rio and Eae againt (relatve to

The Source of2-0xoglutarte for Glutamate Synthesis

Amino aid are teied from te tranamiation of ketoaid it glutamate Gl and oter amino donor ketoaid e all intermediae ofrepirator pata 76 e ompleitie of amino aid ntei are detailed eleere (22), t of ke importane to t anal te proviion of2-oogltarate 2-0G for Glu ntei Tree potetial pata for teprodution of 20G ave een derid It a een tougt tat itrate iproded an idized t 20G te eqential ation of aonitae andAD-ioitrate dedrogenae AD-IDH itin te mitoondria Te 2-OG i ten eported to te tool and loroplat for Gl ntei va

gltamine nttae and gltamine 2-oogltarate aminotranferae G/GOGAT 13 e idetifiation of ooli aonitae (21 and AD-ID(21, 3, 31, 0 mean tat itrate eported from te mitoondra old eonverted to 20G in te tool efore tranport to te roplat Aloroplati iozme of NAD-DH a alo een found (2 22

Cen & Gadal 31 ave propoed tat te tool i te major ore of20G for amino aid ntei aed on e reltive ativitie of te tooland mitoondrialD ioze, te ig ativit of te mitoondrial 2-oogltarate dedrogenae, and te ig apat 20G tranporter in te loroplat evelope 12 e 23fold ger eport rate of itrate over 2-0G iolated pia mitoondria oidizng malate i onitent it te tool rater ta te mitoondri eing te more igniant ore of 2-0G(0 Te loroplati enzme i not togt to e a gnifiant ontritor to

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58 HU TUR

2-0G prodution. No soure of isoitrate for the hloropasti NADPIDH isknown ecause neither a chloropastic acoitase (41) nor a chloroplastic transpot systm or isoitate has been idntifid (30).

UTION O CON FOW TO T TC CYC

T deman or TC yl intmiates inrass wit t rat o nitrognassimilation To understand how tis increased emand for ron from teTCA ycle is baane wit th storage and transloation o rbohydates inthe cell we must examine the reulation of enzymes that irect cron durnnitroen assimilation In eneral, the effetors iscusse here have been found

to act at physiologically relevant oncentrations on the isolatd enym

Phosphoenolpyruvate Carbolase

Posphonolpyruat aoylas is major enym involve in rplacingt TC intrmediats removd for iosynthesis in hiher plants and aga (41 10 We know less aout PEPcase reuation in C3 plants than in C4plants (, 512440142)

Key inhitors of the S mnuum PEPcase re the ano ais Glu anaspatat sp) an th TC yl intmiats 2G an malat (140.Malate inhiits PEPcase from Euglea gcls (11 communs otyeons Post, manspt in prparaion), a nol nyms rom soyan(142) nd upin (100) Asp and Glu iniition has een repored for all ut theugea PEPase whras 20G affcts th nymes fom noules Ohrinterediates includin citrate isoctate succinate and pyuvate yr) havebn inhibitoy to som but not vy PEPas amnd

The onl activators of anapleuroti PEPcase monstate to dat a lutamne Gn) an yroyaetone pospate P) in S muum (0

an DHP in soybean nodul PEPase (142). The rsponse to fetors canv ramatically with pH (142; Podest manuscript in preparation) andthe presn o glyerol (142). Tese prlimina inings iniat a ntralole for sp Glu, an te C yle intemeiates in te feeack iniitiono anaproti PEPase T reatr pospoylation o wat PEPas inillumnate vs rkned leaves has led to speculation that phosphoyation alsomay b involvd in th regulation of C PEPcas ativity (27.

Pyruvate Kinase

nitiation of de novo ano acid synthesis requires the provision of cron toth TC l via pyuvat inas P Tr ae at last two isoyms o PKin a given plant tissue th cytosoli fom PKc) and the plastidi fom PKp)w sow makly irnt rglatory kinti r nr






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NTERATN C AN N MEAM 58

sandn P reguatn s mted despte the evdence ts mptance nmduatng plant metabsm under a vaety physgcal cndtns

s wk n e eguay ppeies of plan PK has demonsaed

inibiin by ATP ciae (9, 117, 164) and acivain by AP (9, 164).nly fu sudies ave relaed PK euaon wh s roe in provdng crbonr amn acd synthess (, 8, 1, 186), and these sudes have shwn thatamn acds ae as centra reguatrs Pc n tssues acve n prtensynhesis Guamae nhbed spinach ea PI (n lcalizain deermned)weeas Asp evesed Glu inibiion indicaing a ole fo e Gu/Asp aio incnllin is enzyme (9). Glu was also a pen inibi f PKc fom nd reinhrdii(98, 86) HAP was an activar an Asp weaklyacvated algal PKc, bu nhbed the plastidic szyme (98)

In coun ctyledns, Glu, Asp, and agnne egulan wa depen den n p a p 6 hey inhibied nd a pH hey acivaed P (EPodes, manuscip in pepaain) Cyslic PK from devepin seed endspem of c swe n esponse amin acids a eie pH (11,118) The reuaory derences in commun PKc iszymes are cnsistenwh the re the endsperm as a substrae exprer and he ctyedns as atssue actve n bsynthess and nrgen assman.

yruv Dhydrgn Compl

he pyruvate dehydrenase cmpex (PDC) cntrs entry cabn nt the CA cyce Athugh ths utenzye cpe has been puied romplans (1a), Randall and cowokes ave elucidaed is eguaion mainyfom impue pepaaions Liinduced inibiion of PC by osola ion is affeced by e relaive aciviy of b PC inase and PC psporyase enzyes that rm part the cmpe gh APADP rats asavr nactvatn PDC, whch ndcates energetc cnr the CA cyceand spprts the hypthess that phsphyatn plays a e n te cnt o

PDC (5, 6) Pyvae acs as an eecive PDC aciva by oveidin APinhibiin f PDC inase (141)

Phophofructokn nd Pyrophopht Fructo-6phophtPhophotrf

Te ineconvesin of frcose 6pospae (F6P) and fcose 1,6bisphos pae (FBP) is a ey feaure in plan respiay meaboism (Fiue b)ypcay, psphrctnase (P) cnvers 6P t P we ructse,6bsphsphatase (Pase) catayzes the reverse reactn n th pastdsand the cyts 3 A cytslc pyphsphate fcose6phsphae ps phoransease (PP) ha opeaes in eihe diecion has been puiied om awide rane o ighe pans (14) he eguaion o cabn ow ugh hesepahways is crcal r prvidin F6P t gycysis and the TCA cyce (36)

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588 PP TUN

All lls in high plants hav a tosoli and plastidi PFK (PFK andK spiv) hw hs isoms ma a wihin h sam planbtwn issu p and dvlopmntal stag (35 36 38) In gn alga it

appas ha h is a ingl pasidi nzym (17 82 83 87) PK andPFK a distinguishabl immunologiall kntiall and b thi stabilitduing puiation (16 35 82 86) Both iszyms shw mpx bhaviowad h ubsaATP and 6Pad his bhai an han dpndinn h pH usd h nnatin inani phspha (Pi) psn (16735558682) Phosphofosknas an undgo hangs in its agggation stat (3) and in ao oot ths hangs alt th gulato andinti poptis f PFK and a afftd b th lvs of ATP and/o 6P(82) Inoani phosphat ativats both ytosoli and plasidi PK a low

nnains (5 mM)whas hih Pi nnains > 2 inhibi hplasidi isozm (16 35 82) PFK and PFK a allostia inhibid bphsphonlprua (PEP) and 3phspha (3-PGA) (55). Aivainb low lvls of Pi an ovid th inhibition by PP and PGA and thPi/PGA and PPP ais a hh b in ulain P and PTe eglaoy pope of PFK ae conen wh he nzm's signifian in onting F6P to FBP to ovid abn fo nirgn assimilation(16 17 35 82 86) PFK should at in th poison of abon fom stah

whas PK shoud b impotant in tissus hat impot suos vsus hosha dpnd n phspha (P) pt m h paidTh ol f th PP has bn dbad sin is disr in plan issus

(35) PFP is full vsibl (27) and is ativatd in both ditn bftos 26-bisphosphat (F26P) (88 15) an funions hav bn poposd f is nym inluding h podution of pophsphat fo suossynhsis (5) and as a lyolyi bpass f PK duin Pi dfiiny (38a)Sd PP in issus i in nn ssmln s q mnwha if any ol his nzm has in poviding abon fo amno aid synth

sis

FrusJ,bspospaas

Both osoli and hloplasti FBPas hav bn pufid and haaizdfom sva pans and alga (8 143 88) Th psn of pastidi FBPashas n bn unisally prd in hphi plan iss and h appas o b a olation btwn th tp of stoag abohdat and thpsn of this nzym (4)

Th tosoli FBPas is gulatd by two majo fatos F26P and APwhih a bth inhibis of h nzm (5 147) Ths mabolis hav aky in ntolling suos snthsis and dgadain Th synthsis anddadain f F26P isf is ontolld b mtabolits F26Pinas aivit



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INRAION O C AN N MABOM

s btd b 3GA, E, d T d s ctvtd b 6 d , w t6pspts s btd b d 6 (3, 47, 4)

Crpstc s s ctv t k pH d f t prt s

rducd v t frrdx (Fd)trdx sstm (54) Ts tw crpstc cngs ccur wt umitn (3) T ctvtd zm rqurs dvnt ctn (Mg+2, C) s cfctr (54, 43). Rdx rgut f tFs d tr zms f t rductv pnts pspt ptw surs gst fut ccg f crb btwn t crpstc rductv dxdtv pnts pspt (OPP) ptws durng ptsntss (4) tt s knwn but w ths ptws r rgutd f strch s mbzdduring ptsthss wvr, durng htsnttc N0 ssmt Nmtd g tr s n xdtv chg t ctvt stt f k

zms f ts ptws (H Hupp, muscript rprt) Ts mdct tt t Fdtrdx sstm c b vvd n gutng nzmctvt n spns t tbc dnds vn n tutn ht nvnmnts

uose Phosphae ynhease

Sucrs pspt sntts (SS), t mjr nzm f sucrs bsnthss, s ctvtd up trstn fm drk t gt (7, 7) Actvt d

crss trugut t d s t zms sstvt t nbt crss (7) In cntrst, t bt f gucs 6phspt (G6) t vrrdSS bt s crss during t d Atug t dctvtn f SSs crrtd n vs wt n ccumut f sucrs, thr s n drct ffctf sucrs n t nzm (7, 53). Instd, g sucrs vs m btT trnsprt frm t crpst dg t rs t SS btr,

Th rgutn f SS rts t t phsprt sttus f t nzm;sprtd SS s mr sstv t nbt (6, 7) T SSpspt gt ctivd b mcnsm nvvng prtn snss (8)

Irgc pspt d G6 r cts f t pprtn mcsms w s drct stc ffctrs SS Irgc pspt btsSSpspts d G6 bts t ks Ts cgs r cssttwt dpsprtv ctvt f SS up trst frm drk t gt(8)

ah Meabolsm

Irgc pspt bt nd GA ctvt f ADgucs prpprs r k t t rgutn f strc sthss () Ltt s knwnbut th rgut f strc dgrdtn (5). Strch m b dgrddcntnu n th gt, bt t vr w rt rtv t ts bstss(5). Atug T s usu prsumd t b t prduct f stc dgrdt xprtd frm t pstd, mts r tr xss mgt s b pr

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590 UPPE TURPIN

dued and de epoed 48) An undeandn o o e euaon oa deadaon neaed no nen meaom l eque undamena adane n ou noede o a deadaon

A MODE OR ACTVATO O RESPATOR CARBOOW TO TRO ASSATO

Te deand eaed an neae nogen alao ee ome poe o elua eoue Te oodnaed epone o man enme a equed o mee ee demand mu e medaed eulao gnal aedamon oe enme

I le noen eng alaed e amnaon o Glu y G oud elo Gn pool ould e o and Gu and TA yle nemedae ould e uae knae and Eae ay ould e ned. H eome aalale e ee o G and GOGAT uae Gu and -0G ldop ue a) Ineaed a o GGOGAT eale a ne alanen Glu and Gn y e demand o aon an e andued o o and Eae o neae gyoy and anapeuo aon o epeelyTe dp Gu non n o enyme eea Gn may aaeEae n ome yem. An deeae n e TA nemedae oud alo

ep o aae o ae and Te podued an eee nono e D y ATAD ao Te omned aaon o Eae andD ould pode aon o e TA le depeng lel o E Tedappeaane o E ould elee non o F. In o a anduoe-dependen ue an ae aae e lo o aon oadgoy ae an uoe yes Toe popae epo om eoopa n eane o ould neae pad u upponpad F aaon and alon e deeae n GA nn ane ee den aon o epao meaolm

An ned yoo Bae equed o deeae e dan on ee epoed om oopla o o nmze ule ylng o mpoed uoeTe lee o 26 an o o yool Bae oud e eeaedeaue deeaed E and GA ould aae e F26-nae le a leel o ool ould n e 26popaae A leel oyool oud alo n n o ay: a de aloe ee andan neae n popolaon ae a non o e popaae

Te eul o maole meaemn dung aaon o H+ amlaon n a aey o ue ae onen many ape o enao

ee elo) Mo o e meaole ue ndued y 0 ae mla uloe an H+(70 Tee ae ao ane n pydne uleodeand O paay omponen dung e one o 0 amaon aele e equemen o edue 0 o H+ (5 10).



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IRAI A ABIS 9

RY CORATO

The carbon requiremen for amino acid ynhei ae independen of he form

of inoranic nioen aimilaed The major dierence beween he aimila on of inoanic N and NH+ ae he enery co aociaed wih hereducon of N o NH+ Compamenalaon of N aimilaory pahway enyme ncreae he compleiy of negraing and conrollng nogenand carbon meabolim durin aimilaion How enery requreen are medepend on he ype of ue and i phyoloical crcumance Ulmaelyphooynhec lh reacion upply all enery n he pan cell Alhouh hepreence of ligh requred for carbon amilaon nroen amlaon canoccur whou he drec npu of lh energy relyn inead on carbon andenery allocaed niay o carbon amlaion and ored n carbon bond

The Enzymes of Nitte Reduction

The enyme of he nrae ailaory pahway have been revewed een vely 89 ) The NR a cyoolc enyme ha reduce N oN by he addon of wo elecron om NADH ( 72) An addi onal NR ha ue eher NAD and ADH ha been dened n everalorganim and conned o roo n mo higher plan ha have been

eamned (33 897) Nre reducae (NR a plad enzyme n algaeleave and roo rduce N o NH wh elecron donaed fom d(8) S/OAT raher han luamae dehydroenae i acceped a hepahway reponble for primary ammonia amlaon ( 3) Mulplefor of GS ncludng boh pladc ad cyoolc ozyme have eendenfed n phooynhec cell and roo erac ( 9 73) Molecularan ndcae ha GS ozyme are epreed n eparae ue of heplan and hu may have dfferen funcon () S he fir ep innoen amlaon ha requre AT. Boh a ddependen and NADde

penden GOAT have been found n leave roo and green algae (9 )he chloroplac ddependn pece he predomnan form and acv y hher n green han n heerorophc ue

Of hee enymeonly he regulaon of NR ha been uded n deal NR hough o he rae of N amlaon becaue any decreae n NRaciviy reul n a correpondng decreae n N alaon 2)Trangen obao plan ha Qverexpe NR however do no amlaeubanially more nirae (52) indicain ha NR aiviy may be opimied ohe eneral plan meabolim.

A diual boynhei/deradaion cycle regulae NR decreang i poenial aciviy hroughou he day (72) Spermpoed on he coae cono ofenyme quaniy i a rapd ncreae of eniivy o M

nhbon upon

darkenin (878798) Seniiviy o M

diappear wihn an hour of

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P & TUN,

expong leae o lgh, bu hen nceae lowly dung he day (7 Seeallne o edence lnk lgh acan dphophylan h la nydeceang eny o Mg+ (8, 79. n o applcaon o okadc acd, a

phophaae nhbo, peen h dehopholaon and opleely blockgh acaon o NR (71

Teaen o leae wh cyclhexde ndae ha NR phophaaeel lgh aaed by a poe dependen on poen ynhe and ha heenye u be ynheed onnually (7 The NR knae nehe uedoe apdly no aeed by lgh, bu nhbed by G6 NR aaon,heeoe depend on egulaon and ynhe o he NR phophaae Addon o ylohexde o llunaed eae lead o a low deeae n NRaaon (71ndang ha he apd dak deaaon o he phophaae

nole oe han a hal n poen ynh.

Cytoolc Rdctnt Spply

oble ouce o cyool eduan o NR nlude yool dehydogeae ohonda A ay an/o phooynhe hehe obeaon o he nduon oN alaon by lgh C and he poon o an/o epao needae n he dak ued o ague o a paulaoue depend on nepeaon ( 97 99 16 11 1 15 184 Foexaple, he lgh dependency o N0 eduon uld plae equeen o phooynhe enegy o phooynhae o be eped o oxddo o lgh nhbon o he TC o ake educan aalable o NR (97

n he plan cell he low NADNAD and NADNAD ao n heyool he hloopla o ohonda, epeely, ao he oganellea he oue o yool duan (61 6 89. Nehe NAD no NADcan e eped decly chlpla chnda he acy he hule beween hee copaen u be egulaed o anan hee

edox deenalhe hloopla ha wo anpoe o eduan (61 anpoed

o he plad a a DA/-GA hule an upply NAD and A ahe cyoolc NAD-glyealdehyde 3-phophae dehydogenae (GAD and-GA-knae. NAD an alo be expoed a alae by he alae-oxaloaeae (OAA hule, whh egulaed by NAD-alae dehydogenae(MD he dual equeen o educed hoedoxn and hghNADAD ao o NAD-MD acaon ec expo o eduana alae een n he lgh 8

The ochonda expo eduan a a alaeOAA hule o upplyeleon o peoxoe o he eduo o-hydoxy pyuae n phooepaon (79, 9. Malae oul be oxded n he cyool by he yoolNAD-D, bu only OAA eoed a enough o pull he eaon n



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ITERATI AD METAIM 5

aor o NA rdtio A trasloator it a ig aiity or AA an dntd n t toonda o pa dling and root )

lnt tndrl tt drn ptt rlt t t potntl

ro 0 aiiatio g at yt dig potytirqirs tat A aro o oti i t ligt 5 t as agdtat t itE i iiitd i t ligt y t ig ll o A prodd ypotopoporlaton ror t A lprodd rdant aal t R 3 35) r tntal dn, r, tat tondral rpraon not lgt ntd 38 Rprator 0 ptk an asrd i t ligt t opliatig potorspiratio ad t Mlrratios a iiitd 1 Ftr, iiitio o t itE y oligoyisos tat itoodria otrt sigiiatly to ytooli A lls i t

lgt 8 nrad rprtr qtnt drn 0 laton ad t t

latio o da N0 rdtio s o tE lto s iiitdially or y aaroiosis spports a rol or a itoodrial spply orgy to N0 rdtio 58 1 A igiat oplig o rspiratio toda 0 aiilatio i aly plats a n dotratd 12 additon, llto o t ltron rqrnt nry to pport t rat o0 altn rd n -ltd tat potoyntt ltr ppl ndqat to t t nd o 0 rdton andtat so rdtat t proidd y rpiratio 19

rdo qilii t plat ll opatts is aitaid ytgt rglatn o lorplat and tndral rdtant tranton (36) tor tng on or o tol AH old trggr otror to pnat r t n trt rdtn o o ny rto in t total ytooli rd q t ay o t ytooli NADprodig patays a pottially pply rgy or N0 rdtio g 45,

Enegy Supply n he lad

THE HLRLAST N THE LGHT AND DARK o o t tr rdttp in 0 alaton NR and GOGA tak pla n t loroplat andrqr Fd gt tlaton and M ton o 0 rdtion i grtiss idiat a li t liar potoytti ltro o ad itrogassiilatio 58 123 18 Bas ot N0 rdtio ad aro assiilaton or in t lorplat, t pattonng o ltron tn t topata t ontrlld n tr prnt a dontrtd tat taddton o NR to a rntttd ligt yt lor t rat o AHoraton y Fd-A odordta FR) ) atrat lgt, tpply o ltron aptor ratr ta rgy lit potoytti ltro

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59 HUPPE & TURPN

tapt 4) Ehaemet li elet tapt duig N0amlat 34) idat d a l at bau th a w k phtht l lhu l w tmuatd du ph

ttheti N0 aimlati, C0 xati i uppeed i Nlimted agae43, mmium au le uppei 0 xati h twbea demtate a det eet the eg demad N0duti the b aimilati at uppei C0 at b amlat l qut tat h plt 12 12) papbaue C0 aailablt athe ha eeg te limt the ate C0 xati hgh plat

Nitate aimlati i a phttheti ell de t ala uie lightg ) Rat N0 aimlati i daked leae a high a 76%

the ate the lght hae bee pd ) Reduti N0 ha bdemtated i the dak ude aabi ad aebi diti 7) Dkhbiti 0 aimat a te be eme plat ad alae the hae a alteati ab ue 7, 3, 63)

Ctplam b xdat uld uppl hplat dutat thx hu tht xp c phhic dutt h cl weeed i the dak Hwee, tw the hlplati ezme iledNADMDH the malate/AA huttle ad the NDGDH th

DHA/3GA hutle ae lightegulated ezme ad huld be iatie ith dk 4, ) Rdutat uld b pdued i tu b th hplat pathwa, tw ute albl he degadati th 5)Chlplat gue phphate dehdgea GH) whih lthe patha, i ihibited b bth l NADNADH ati ad edut a th dthdx tm 4 36) th dk G6DH xidd the pathwa a peate 37; TJ mauipt i ppaati)

limted aga hage DDH ad phphgluatG6 ati ad the xidatie atiati G6DH demtate atiati

h athwa up uppl ihe th dk liht 75 7HC Huppe maupt i pepaat) igue l ) h eed eg aapptl mulat ab xidat th dk ad d ihbit the patha the lght Hw te NADH pdud edu d tla C b lea NR tgl a eduti NADH b deg the NR eat wuld qu a hgh tu d 45)

REUTN UL N TE ROOT h pahwa pbab upp N0 amla t t 34) Al m th pawaG6DH, 6phphgluate dehdgeae, taketlae, ad taaldlae) ae pt i bh t platid ad tl 46), ad hi ateiead he t ee iubate ith N0 34 he ati C



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NERAN AND N MEAM 55

relesed from C nd C beled glucose fed to these roots durng Nssiltion ws lso consistent with incresed O pthw ctivit )

Reduction of N to lu in isolted plstids nd the plstidic loction of

bo NiR nd OAT ) indicte tht te plstidic pthw is nvolved solted root plstids trnspor nd T , 5, ) There is stoichiometrc reltionshp between the rte of N reduction nd odtion in isolted plstids from pe nd ble , , 08). ee 7) hsestimted tht the potenti O pthw ctivit in the plstid is insufficientto support the potentil for N reduction 97); however, this is bsed on invito knetics, which m not ccurtel predict the in io potentils of eithepthw.

The O pthw produces NAD, wheres NiR nd OAT re dde

pendent enzmes Fd hs been isolted fom roots o rdish, mung ben, ndspinch 3, 0, l , 7) A FNRlike potein hs been detected in both co7) ad pa oos 0) ad solad fo spac ad adsh oo 0, 0)The identicton of hese two poteins povides mechnism to reduce d inroots

Root FNR ctivit ws detected b NAD oo rther thn redction 7) Fd nd FNRlie components in mize roots could substtute for lefd or NR, respectivel, in ctochome c reduction b NAD ut not in thephotorduction of NAD 7), which is the kineticll fvored direction of

lef FNR Ben sprout FNR wored in both directions, but there ws kineticdvntge i NAD redution when the spinch d ws used, wheres theben sprout d wored better to reduce ctochrome c ). he pimrtcture of the pinch root FNR nd d differ from ther lef counte05). t wil be interesting to le how tese observtions relte to thepprent role of the root enzme in Fd reduction rther thn oidtion

AP Suy

Whether the mitETC, glcolsis, or photophosphorltion supplies AT tonitrogen metbolism depends on when nd where nitrogen is ssimilted. Drnd heterotrophic tissues must rel on glcolsis nd the itETC, wheresphotophospholtion cn contribute AT to both the chloroplst nd ctosolof photosnthesizing cells Although chloroplsticll produced AT tnsported cross the chloroplst membrne s T is thought to be mor supplierof ctosolic AT, it is now pprent tht mitochondri contribute significntmounts of AT in the ctosol in the light 0).

imr nitrogen ssiiltion reuire AT t the step nd polinvolves the chloroplstic S isozmes, but the source of AT in drkenedchloroplsts hs not been identified The onset of N+ ssimilion cuses trnsient drop in AT levels tht is corelted with n ctivtion of sevidenced b metbolite chnges ), but the comprtmentliztion of these

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RN

cange unnwn. lpla tanpt aenlae ecl a a lw ae.ac eraan ma e ale upp AT pucn ecaue anm f e e lclc enzme are peen n e clrpla (36) A

uppl f A a n een emnrae n e r pla eer u man e reque ecaue e cc G pra e mar me nr ()

n e l cell ma mulae e capac fr an NH prucn frm pne cnrllng e relae ace f cclc annn-cclc elecrn flw ae ann (8). ae rann ma aler an NH prucn n repne cane n carn (66) annen mealm (6). Wen NH+ amlae e cellularNH requremen ncreae ramacall relae e f C

fxan. Fllwn 2 mn f NH amlan e Kurecence pecra we a ecreae n e ra f II I flurecencepcal f a ae 2 rann pl farng A prucn cclcelecrn ranpr. Reucn an alan f N wc eman mreNH an NH+ alan n ne a mlr ae 2 rann T can ml e nr eman f en amlan can mulae e quanum el f S!.

GUTION DFNT YTUneaning e eglan f ae-lming enme ung e negatinf nigen an can amla equie eamnan f e mealiefue a ccu n elan e plgcal epne cmne wicaaceiain f e ilae enme. Te pilgical an mealecange a ccu win ecn f e ne f ngen amlan aveeen meaure n uncellular alae. ee ean can e apple elnge me cue meaual in ige plan nieain f e iffe

ence in e mealic equemn f e cell allw peculan f w eegulan mig iffe an pvie cle f fuue w.

The Algae

In N-le alae e ne f NH+ amlan caue a rap rp n Gluan A an an ncreae n A AMan Gn (4666)). eel f 2-Geclne reflecng e mealm f Gn GOGAT (66) e f cangen lcc meale a rp n E an an ncreae n r wnecn f NH+ an cnen w ap acan f PK. Te

cncenan an eul f uncuple epermen ncae a K aenlae rerce n ee cel efre NH an (8) erefre K acan ma reul al fm e ranen re n eel () fe eel ece e rp n Glu an nceae n T wul uan acivan.



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NAIN AN N MABIM 7

Ecas is stimulatd rapidl praps rspondin to t dcrasd GluGnratio (71)

itron nricmnt causs a dcrasd ow of fixd to carbodrats

and can induc starc dradation and supprssion of potosnttic fixation (8 16 16) A drop in B and F6 indicats slowin of trnraon pas of t rductv pnos pospat patwawl incrasn FB and T lvls dmonstrat rroutin of fxd carbon to t ctosol( 16166) T drop in and -A mdiatd activation of K andEcas sould rliv iniition on t plastidic FK (17) and ad t xportof carbon from t cloroplast in xcan for i Incrasd cloroplastc ifavors starc bradown

atitionig caon in t ctosol of nicllular aa plfid bcaus

tr is no translocation of sucos to or tissus S for xamplas no ctosolic FK (17) An drain on T b t ctosolic FBa would biniitd F6 wic incrass initial at t onst of nitron assimilation n (18166)

itrat asimilation inducs simlarut slowr cans in most mtaolits in S (17) Tr is alo a rapid drop in G6 lvls and tADH rducton stat and an incrs in 6-pospoluconat lvls andG6DH activit at t onst of 0 assimilation wic dos not occurdurin H

+ assimlation (717) Ts cans indicat tat stimulation of

t O patwa to provid rductant occurs in bot t lit and t darfor carbon ow incrass to suppl amino acid iosntsis

he Phoohec Hher Pla Cell

imilar rulator sinals oprat in ir plants and ala to activat carbonow to amino acid biosntsis durin nitron assimilation In laf discs andsoatd lnitron stimulats incrad lablin into amino acids rad TA inmdiats (8 185) wic indicats incrasd flow of caron

from potosnttic fixation and activaton of anaplurotic fixationpossibl involvin posporlation of t Ecas (27) orr tim courssusin otolasts sowd a transint drop in lu laln wit a ris in n3 In -limitd co plantsan incrasd /u ratio was found witn 1 of rsuppln niton 3

T mos consistnt ffct of nitron assimilation on laf caron mtaolism is tat sucros sntsis dcrass wil starc or rductiv pntospospat patwa comonnts ar oftn not actd (8 1 . -limitation pants inducs an ovrall incras in carbodrat lvls tat is oftnratr in roots tan in lavs (1) T major function of matur lavs is tosuppl sucos for t rst of t plant T ctosol is t pima sit for trulation of carbon partitionin in lavs; ncpotosntsis can b unaffctd vn wn tr ar lar cans in t dirction of carbon ow in t

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598 HUPPE & TURPIN

cytoo (151) Wheher cel export ucroe or tore trch ht hepexplin why leve hve een repored to tute to uppre nd to notchne 0 evolution or photoynthetc 0 xtion upon niroen reuppy

he ey reultor drectin caron o uconeoene nd repirton nthe le cytool re F6P nd P whc ect SPS nd the enye reultnhe lnce o 6P nd BP (, 7 51) ew det o he reutory uxen ture ource leve durn nroen iton hve een repored In thedevelopin cotyledon o whch lo ucroe in, nyi oetolte chne durn nitroen ion deontrted conupon oornc cid (0G nd Pyr) n uppot o ino cd ynthe (56) dropin PEP nd P ndicted ctvtion o PK nd PEPce Reievin the PEPnhition o 6Pne increed P therey encourn lycoytc

cron flow vi the PP nd nhitin ow to luconeoene vi cytoolicFBPe

Dun N iltion n ource leve, hih level o 6P nd Pi woulde expected to nht the ow o caron to ucroe v BPe nd SPSDecrein SPS ctvty would ower con rnlocton ro ource tuewhere n inhted SPS in in te woud vor the port nd derdton o ucroe

he reultion o SPS nd NR phophoylton y hve ole indrectn caon ow duin N0 iltion (7) Detched leve ccu

ute photoynthte Excion cue ore rpid phophoryltion (nhiion) o SPS durn the dy thn n ttched leve where the NR ctivtydoe not decree (7) Greter vllity o cron y vor N0 iton he eect o crohydrte ttu on the htrelted dephophoryltono SPS nd NR coud e portnt in undertndn the derent drtion rte ound n plnt

Heephic Tissues

Root ut ue dierent ttee thn do photoynthetic tiue to uppotnitroen iton ecue trnlocted photoynthte provde oth croneleton nd l the enery to root Cytoolc enery or ntoen ltion i proly provded y ycoly the cycle, nd the tE iti n the le (5) he cytoolc OPP pthwy lo could e ipotnt in ootwth NR tht ue oth NDPH nd NDH (16)

Root contin oye o l the reutor enye tht interte cronand nten eto n eve n le; however there h een ereearch done on the reultion o roo ioye (, 6) In uceton

root, oe reulto echn ht vr coniderly ro thoe nource leve It will e interetin to invetite the reultion o he lihtreulted enye, uch SPS nd NR, to deterne crohydrte ttu the portnt reultor n in root






23/31

NGATON OF C AND N MABOLSM 5

The platdc OPP pathway the mot lkey ouce o eductant o ntteeducton (5 and a mao oute o deadng tach whch upple cabono amno ad ynthe n ome platd yet te eguaton o oot G6DH

not undetood The mechanm o Fd educton a the oot R alo equeuthe haacteaton

Although ou knowledge o ntogen amlaton n oot mpongthee much wok to be done on the metaboc egulaton n th ue

CONCUDIN REMARKS

We ae only begnnng to eale the omplexty o abon and ntogen nte

aton n photoynthetc eukayote Depte deent abon ouce tachucoe, o ecent photoynthate) o ntogen amlaton n aou tue,m unn atu mn ntal t n nn amatneut n meabolte change that actate glycoy at the leel o P andPPcae Th actaton caue metabolte change that gnal the need todect cabon to eatn om th tach, ucoe, th euct pentoe phophate pathway, dependng on the tue Dung photoynth, themetabolc demand o ntogen amlaton ae tanduced to the photoyn

thetc appaatu to baance ATP and NADPH poduton wth the enegyequement The exta eductant demanded n amlatng hange thental poty o abon ow n tue whee photoynthet enegy nadequate to meet the eleton demand n uh ae, the O pathway atated to pode edung powe The nogen an abon etab pathwayae nteconnected though egulato gnal that ncude alotec metabolteeecto, phhlaton, and d eulatn. abn and ntgn mtabolm ae ntedependent pocee, o nethe can contnue to opeate to thedetment o the othe Nutent tatu, ethe o cabohydate ee o ntogen

compound, ae mpotant acto n detemnng the poty o eouce uebetween thee two pathway The dea and conept that ae emegng omtude o cabon and ntogen nteacton n plant wll contnue to podeuch nght nto how each o thee poee act nddually and how theynteact n epone to the enonment o the oganm.

KOWLDGTS

We wh to thank D M-L hampngy, Foye, Hube, Oak, and W

Platon o odn unpublhed manucpt and pepnt We ae ndebtedto D R uy and W Platon o ctcal eadng o the manucpt to ongdon o atance n pepang th manucpt, and to TJ Fa o oealluppot and o dong o much, o wel

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