Heng Savoeun/2009Heng Savoeun/2009 11

PhotosynthesisPhotosynthesis

RbtikmµrsµIsMeyaK• RKb'CIvitTaMgGs'elIEpndIeyIgenH BwgEpÁkeTAelIBnøÅRBHGaTitü• ruk¡Cati nigmIRkUsarBag<kay‘cyanobacteria’eRbIR:s'BnøÅRBHGaTitü edIm|ITajykplRbeyaCn¾zamBlBnøW enAkñúgRbtikmµKImI EdleKehAza RbtikmµrsµIsMeyaK

• zamBlBnøW E = hν = hc/λ

h = Planck constant(6.626x10-34 Js)

c = Velocity(2.998x108m/s)

Heng Savoeun/2009 2

c = Velocity(2.998x108m/s)

ν = Frenquency

• kñúgdMeNIrrsµIsMeyaKenH ósµ&nkabUnic‘CO2’RtUv:nbMElgeTACakabUGuIRdat• smIkarsrubénRbtikmµKW;

CO2 + H2OBnøW (CH2O) + O2

• elakRobert Mayer(GñksrIrviTüaGaløWm"g'’:nbkRsayBI ruk¡CatibMElgbþÚrzamBlBnøÅeGayeTACazamBlKImI

kørU:)øas (chloroplast)

- kEnøgEdlruk¡CatieZVIRbtikmµrsµIsMeyaKKW kørU":øs EdlCaZatumYyénekasika- kørU":øs manTMhMu RTg'RTay nigragxusKñaBImYyeTAmYy- kørU":øsmanPñasBIrCan'. cenøaHrvagPñasTaMgBIrenHehAza kEnøgcenøaHPñas. Pñaskñúg

ruMBTÆ&RsþÚm"a‘stroma)EdlmanGg'suImy"agxab'. enAkñúgRsþÚm"aman DNA, RNA nigrIbUsUmEdlcUlrYmsMeyaKkørU":øs-RbUetGuInepSg> .

- ekasikamYymanBI1eTA1000kørU":øs

Heng Savoeun/2009 3

rIbUsUmEdlcUlrYmsMeyaKkørU":øs-RbUetGuInepSg> .- ekasikamYymanBI1eTA1000kørU":øs- kørU":øsmanRbEvg 5 µm

kørU":øsrbs'søwk spinach

:nmkedayeRbImIRkUs,ÚbeGLicRtUnic

kørU:)øas

Ù PñasBIrCan'B&TÆCMuvijtMbn'TwkEdleKehAza RsþÚm"a

Ù enAkñúgRsþÚm"aman

Heng Savoeun/2009 4

Ù zg'enHGacekItmaneLIgCaragbMBugEdleKehAzaRkaNa‘grana)

Ù tMbn'énPñasTILakUGuIt EdlminCab'nwgRsþÚm"aKWCabnÞHRkaNa‘granal lamellae)

Ù tMbn'Edlb"HnwgRsþÚma"ehAzabnÞHRsþÚm"a‘stromal lamellae)

Ù EpñkxagkñúgénPñasTILakUGuIt KWCaRbehag‘lumen) .

EdleKehAza RsþÚm"aÙ enAkñúgRsþÚm"aman thylakoid

membrane Edlbt'Ebnbeg,ItCazg'‘vesicle)

dMeNIrrsµIsMeyaK • rsµIsMeyaK KWCadMeNIrRbRBwtþmYyEdlkñúgenaHeGLicRtugénmU"elKulkørU"PIl

EdlRtUv:nrMejakRtUv:nbJ¢ÚnbnþbnÞab'tamry;GñkTTYl‘acceptor’EdltMer[bKñatamlMdab'edIm|IbMElgBIzamBleGLicRtug(electronic

energy) eTACazamBlKImI‘chemical energy’

Heng Savoeun/2009 5

eTACazamBlKImI‘ ’• rsµIsMeyaKRbRBwtþeTACaBIrdMNak' ;

-Ù RbtikmµPøW ‘light reaction’-Ù Rbtikmµggwt ‘dark reaction’

RbtikmµPøW • RbtikmµPøWeRbIzamBlBnøÅ‘light energy’ edIm|IsMeyaK NADPH nig ATP

• RbtikmµenHRbRBwtþeTAenAkñúgPñasénTILakUGuIt(thylakoid membrane’• zamBlBnøÅRtUv:nykeTAeRbIedIm|IeZVIGuksuItkmµelImU"elKulTwk

Heng Savoeun/2009 6

2H2OBnøW O2 + 4H+ + 4e-

1- karRsUbykBnøW

• kuMepøcRbmUlBnøW‘light harvesting complexes’minRtwmEtmankørU"PIl‘chlorophyll)eTKWvamanCatiBN(CMnUy‘helping pigmente)EdlCasarZatuRsUbBnøWEdr

• mU"elKulkørU"PIl nigCatiBN(CMnYy RsUbnigRbmUlBnøW:nehIy rYcdwknaMzamBlBnøW‘photon’EdlRbmUl:nenaHtamry; resonance-energy-transfer BImU"elKulmYy

Heng Savoeun/2009 7

‘photon’EdlRbmUl:nenaHtamry; resonance-energy-transfer BImU"elKulmYyeTAmYy rhUtdl'mCÄmNÐlRbtikmµrsµIsMeyaK (photosynthetic reaction centre)

• kñúgkardwknaMenH vaRtUvkarEtry;eBl‘10’-10vinaTI bu"eNÑaH• RbtikmµdMbUgénrsµIsMeyaKRbRBwtþeTAenAkñúgmCÄmNÐlRbtikmµrsµIsMeyaK

kuMepøcRbmUlBnøW nigkarbBa¢ÚnBnøW

Ù rMejacEdlRtUv:nbeg,IteLIgedayRsUbykBnøW eZVIdMeNIreTAmk

Heng Savoeun/2009 8

Ù vaeZVIdMeNIrrhUtdl'xøÜnvaRtUv:ncab'ykedaymCÄmNÐlRbtikmµ‘rUbKMnUrmUlBN(exµA’ rWRtUv

:nbeJ©jmkeRkAvij (fluoreszence)

edayRsUbykBnøW eZVIdMeNIreTAmktamry; Resonance energy transfer

rvagmU"elKulénkuMepøcGg'Etn‘rUbKMnUrmUlBN(s’

2- mCÄmNÐldwknaMeGLicRtug (electron transport centre)

• ruk¡Cati nigBYk cyanobacteria eRbIRbtikmµGuksuItkmµénmU"elKulTwk EdlRtUv:nCMrujedayzamBlBnøWedIm|Ibeg,It NADPH

- 2H2OBnøW O2 + 4e- + 4H+

- NADP+ + H+ + 2e- NADPH

Heng Savoeun/2009 9

- NADP + H + 2e NADPH

smIkarsrubKW;2NADP+ + H2O → NADPH + O2 + 2H+

• karbeg,ItGuksuIEsn KWsæitenAeRkamGMeBIénmCÄmNÐlRbtikmµrsµIsMeyaKBIrEdlenAtMer[bKñaCaCYrKW;

1- photosystem I(PS I) CaGñkbeg,It NADPH BI NADP+

2- photosystem II(PS II) CaGñkeZVIRbtikmµGuksuItkmµelImU"elKulTwk

KMnUr Z• smasZatuEdlcUlrYmkñúgkardwknaMeGLicRtugBITwkeTAeGay NADPH KW;1- PS II

2- cytochrom-b6-f-complex

Heng Savoeun/2009 10

KMnUr Z KWCaKMrUbgØajGMBIkMriterdukmµEdlTak'TgnwgkarhUréneGLicRtugqøgkat'GñkdwknaMeGLicRtugrsµIsMeyaK (photosynthetic electron carrier)

2-3- PS I

• eGLicRtug RtUv:ndwknaMtamry;GñkdwknaMeGLicRtugcl&t‘mobile electron carrier) EdlenAcenøaHrvag PS IInig PS I

dwknaMeGLicRtugeTAeGay NADP+ tamKMrU Z

• bnÞab'BIkarRsUbBnøWrbs' P 680 TwkRtUv:nbMEbkeday Mn-complex ehIykñúgeBlenaHeGLLicRtug GuksuIEsn nig 4H+ RtUv:nbeg,IteLIg

• eGLicRtug RtUv:nRbKl'eGay P 680 rYcRtUv:nbJ¢ÚneTA plastochinon(Q) tamry;Rcvak'dwknaMeGLicRtug ehIy plastochinon(Q)RbKl'eGLicRtugeTAeGaycytochrom-

Heng Savoeun/2009 11

Rcvak'dwknaMeGLicRtug ehIy RbKl'eGLicRtugeTAeGayb6-f-complexrYceGLicRtugRtUv:neGaybnþeTA plastocyanin(PC)

• kñúgeBlCamYyKñaenH RbUtugRtUv:nbJ¢ÚnBI stroma eTAkñúg thylakoidroom

• plastocyanink*eGayeGLicRtugeTA P700 bnþ• eGLicRtug EdlRtUv:nbeJ©jBI P 700 RtUv:nbJ¢Úntamry;Rcvak'dwknaMeGLicRtugrhUtdl' NADP+

3- pUtUpUsVrIlkmµ(photophosphorylation)

• kørU":øsbeg,ItATPenAelIpøÚvdUcmItUkugRDIEdr KWvaP¢ab'eTAnwgkarrMlayénCMralkMhab'RbUtug‘proton-gradient’CamYynwgkarsMeyaKATPEdlman synthase CaGñkCMruj

pUtUpUsVrIlkmµ TamTarnUvPaBlÁ«tx©s'énPñasTILakUGuIt/

Heng Savoeun/2009 12

RbUtug‘proton-gradient’CamYynwgkarsMeyaKATPEdlman synthase CaGñkCMruj

Ù kardwknaMeGLicRtugcMnYnbYntamry;cytochrom-b6-f-complexnaMeGaysMerc:n 4RbUtugBIRsþÚm"acUleTAkñúg thylakoid-room

Ù RbUtugenHhUrcUlmkkñúg stroma tamry; ATP-synthase ehIyvak*skmµeLIg rYcCMrujRbtikmµbeg,It ATP

Rbtikmµggwt (dark reaction)

• kñúgRbtikmµPøW :nbeg,It ATP nig NADPH

«LÚveyIgsikSaza

Heng Savoeun/2009 13

«LÚveyIgsikSazaetIkarsMeyaKkabUGuIRdatecjBI CO2 y"agdUcemþc/

rUbbgØajGMBIsmasZatuénPñasTILakUGuIt Cab'Tak'TgnwgRbtikmµrsµIsMeyaK

sJ¦a‘ ’ KW dMeNIrénBnøWEdlcab':nsJ¦a ‘→’ KWkardwknaMeGLicRtugsJ¦a ‘---’ KWkarbMlas'TIrbs'RbUtug

Heng Savoeun/2009 14

vdþ calvin rW reductive pentose phosphate-cycle

(RPP-cycle)

• dMeNIrEdlruk¡Caticab'yk CO2edIm|Isg'bJ©ÚleTAkñúgkabUGuIRdatRtUv:nBnül'edayelak Melvin Calvin, James Basshamnig Andrew Benson

rvagqñaM1949 nigqñaM1953BYkKat':neZVIkaBiesaZn*edayeRbI kñúgkarciJ©wmsarayéb'tg

Heng Savoeun/2009 15

• BYkKat':neZVIkaBiesaZn*edayeRbI 14CO2 kñúgkarciJ©wmsarayéb'tg

Heng Savoeun/2009 16

vdþ calvin

vdþ calvin begáIt glycerinaldehyd 3-P BI CO2

ÙeKEckvdþ calvin CaBIrdMNak';1- dMNak'karplit‘production phase’

- 3mU"elKul ribulose 5-PeZVIRbtikmµCamYy3mU"elKul CO2beg,It:n6mU"elKul glycerinaldehyd 3-P

- RbtikmµenHeRbIGs' 9mU"elKul ATP nig6 NADPH

2- dMNak'karekIteLIgvij‘regeneration phase’

Heng Savoeun/2009 17

2- dMNak'karekIteLIgvij‘regeneration phase’- 5mU"elKul glycerinaldehyd 3-P RtUvrgRbtikmµbnþbnÞab'eT[t edIm|Ibeg,It3mU"elKul ribulose 5-P eLIgvijEdlCakEnøgcab''epþIménvdþ

- RbtikmµenHmann&yza KWCakareZVIBI 5C3 eGayeTACa 3C5

• smIkarsrub KW;3CO2 + 9ATP + 6NADPH + 5H2O → 9ADP + 6NADP+ + Triose phosphate

(Triose phosphate = Glycerinaldehyd 3-P rW Dihydroxyacetone phosphate)

Ribulose bisphosphate carboxylase

• Gg'suImenH CMrujRbtikmµEdlcab'yk CO2 BIbriyakas. dUcenH vaCaGg'suImmYyd*mansar;sMxan'bMputkñúgBiPBelakeyIgenH eRJHza CIvitTaMgGs'elIEpndIenHBwgEpÁkeTAelIskmµPaBrbs'vaEtmYyKt'

• Gg'suImenH mancMnYnrhUtdl'15PaKryénRbUetGuInkñúgkørU":øs• tamkar:"n'sµan RbEhl40lanetan kñúgmYyqñaMRtUv:nsMeyaK

Heng Savoeun/2009 18

• tamkar:"n'sµan RbEhl40lanetan kñúgmYyqñaMRtUv:nsMeyaK• RbUetGuInenH RtUv:nsMeyaKeday chloroplast-DNA nigmYyEpñkeT[ténGg'suImenHRtUv:nsMeyaKBIéNVyU"

glycerinaldehyd 3-P KWsMrab;sMeyaKKøúykUs nigsarFatuepSgeTot

• glycerinaldehyd 3-P EdlCaplitplTI1énrsµIsMeyaK RtUv:neRbIenAkñúgkarsMeyaKsarZatuCaeRcIn

• glycerinaldehyd 3-PRtUvbMElgeTACa fructose 1-P rYckøayCa glucose 1-P eRkamGMeBIglucose phosphate isomerase

• glucose 1-P KWCasarZatudMbUgénkabUGuIRdatsMrab'ruk¡CatiÙkñúgenaHman;

Heng Savoeun/2009 19

ÙkñúgenaHman;- su)yRkUs EdlCas,rdwknaMd*sMxan'sMrab'dwkCJ¢ÚneTAkEnøgekasika Edlmin

skmµkñúgkareZVIRbtikmµrsµIsMeyaK- GamIdug KWCasarZatubMrugd*sMxan'sMrab'ruk¡Cati- EsluyLÚs KWCasmasZatuTMrg'rbs'PñasEsluyLÚs‘cell-wall)énruk¡Cati- karsMeyaKGasuItxøaj' nigGamINUGasuIt

su‘yRkUs nigGamIdugRtUv)ansMeyaKecjBI RPP-cycle

• enARKb'ruk¡Catizñak'x¬s'TaMgGs' kabUnEdlvacab':nzµI> RtUvbMElgeTACaGamIdug rWsu)yRkUs

• su)yRkUsRtUvbMlas'TItampøÚEGm‘phloem) eTAsrIrag<ruk¡CatiEdlminGaceZVIrsµIsMeyaKxøÜnÉg:ndUcCa rWs RKab' nigsøwkBnøkxøI>

• kñúgkMLúgeBlézá enaHGamIdugRtUv:nrkSaTukkñúgsøwk

Heng Savoeun/2009 20

• kñúgkMLúgeBlézá enaHGamIdugRtUv:nrkSaTukkñúgsøwk• kñúgkMLúgeBlyb'EdlKµanrsµIsMeyaK enaHGamIdugRtUvbMEbkeTACasu)yRkUsnigeRbICaRbPBzamBlsMrab'søwkenaHxøÜnÉgpÞal' nigsrIrag<epSg>eT[ténruk¡Cati• su)yRkUsRtUv:nsMeyaKenAkñúgsuItUsulBI glycerinaldehyd 3-P nig dihydro-

xyacetone phosphateEdlcab'kMeNItmkBI RPP-cycle nigxøHeT[tmkBIGamIdug• GamIdugkñúgkørU":øsRtUvbMEbkeTAKøúykUs rW glucose monophosphate nigbMElgeTACa triose phosphate tamKøIkUsIl

1-sMeyaKsu‘yRkUs kñúgkørU:)aøs4mU"elKulén triose phosphate(4C3)BI RPP-cycleRtUv:nRbmUlpþúMbeg,It:nCa1mU"elKulsu)yRkUs (C12) kñúgsuItUsul

Heng Savoeun/2009 21

2-sMeyaKGamIdug kñúgkørU:)aøs

Ù Rbtikmµcab'epþImedaykarbMElg triose phosphate

Heng Savoeun/2009 22

bMElg triose phosphate

eTA fructose 1,6-bi-P

Ù fructose 1,6-bi-P bMElgCafructose 6-P rYceTA glucose 1-P

Ù glucose 1-P køayCa ADP-glucose

karRtYtBinitüvdþ calvin

• eBlézá ruk¡CatiTTYlzamBltamry;RbtikmµPøW nigRbtikmµggwténRbtikmµrsµIsMeyaK

• pÞúymkvijeBlyb' ruk¡CatiRtUveRbIR:s'GaharbMrugrbs'vaedIm|Iplit ATP nigNADPH tamry;KøIkUlIs pUsVrIlkmµGuksuItkmµ nigdMeNIrpUsVatb"g'tUs

• edaysarEtenAkñúgRsþÚm"amanGg'suIménKøIkUlIs Gg'suIménvdþ calvin nigGg'suIméndMeNIrpUsVatb"g'tUs dUcenH ruk¡CatiRtUvTukdakzamBltamry;karRtYtBinitü

Heng Savoeun/2009 23

dMeNIrpUsVatb"g'tUs dUcenH ruk¡CatiRtUvTukdakzamBltamry;karRtYtBinitüEdlBwgEpÁkBnøW

ÙÙ karRtYtBinitüénvdþ calvinmanbIkEnøg;- ribulose bisphosphate carboxylase (RuBP-carboxylase)(RbtikmµTI2’- fructose bisphosphatase (FBPase) (RbtikmµTI7 )

- sedoheptulose bisphosphatase (SBPase) (Rbtikmµ10’

skmµPaBén RuBP-carboxylase GaRs½ynig kkþa2EdlBwgEp¥kelIBnøW;

1- vaERbRbYltamtMél pH

- kalNamanBnøW eZVIeGaytMél pH énRsþÚm"aekIneLIg RbEhlBI7eTA8 eRJHRbUtugénRsþÚm"aRtUv:nbUmbJ©ÚleTAkñúg thylakoidroom

- RuBP-carboxylase man pH RbesIry"agcegá{tcMeJH pH=8

2- vaRtUv:neZVIeGayskmµtamlk¡N; eday

Heng Savoeun/2009 24

- man RbesIry"agcegá{tcMeJH2- vaRtUv:neZVIeGayskmµtamlk¡N; allosteric eday NADPH

- RuBP-carboxylase RtUv:neZVIeGayskmµtamlk¡N; allosteric edayNADPH EdlRtUv:nsMeyaKeLIg kñúgRbtikmµBnøWén PS I