PHOTOSYNTHESI

S
PHOTOSYNTHESI Si st he pr ocess by whi ch phot osyntheti
c or ganism
harnessr adiantener gyandconv erti ttousef ulf
ormsofchemi calener gy
basical
lyt osust ainnotonl ytheirownexi stencebutal sot heexi stenceall
aerobic non- photosyntheti
c or ganism. Or gani sms capabl e of
photosy nthesi sarev erydiverse,rangi ngfrom simplepr okaryoti
cf ormst o
the largestor ganisms ofal l
,t he gi antr edwood t rees ofCal i
forni
a.
Photosy nthesi sist heprimarysour ceofener gyforal llif
ef orms( except
chemol ithot r
ophic pr okaryotes). The amount of ener gy st ored by
17 17
photosy nthesi sisenor mous.Mor et han4. 2x10 kJ( 10 kcal )off ree
energyisst oredannual l
ybyphot osy nthesi
sonEar th,whi chcorrespondst o
11
theassi mi lati
onofmor ethan10 t onsofcar boni ntocar bohy drateand
otherformsofor ganicmat ter
.
Photosynthesi
scanbedi vi
dedintotheli
ghtdependentr eact
ionsandt he
l
ightindependentreact
ions“dar
k”react
ions.Thesedi
fferentreactionsare
l
ocalized in dif
fer
ent chl
oropl
ast compartments:the l i
ght-dependent
processesoccuri nthethyl
akoi
dmembr anewhereast hedar kr eact
ions
occurinthestroma
TheLi
ghtDependentReact
ions
Thel ight
-dependentreacti
onsusel i
ghtener
gyf rom sunl
ighttogenerate
reducingpowerf ortheproducti
onofNADPH, ATPandO2. Li
ght-
dependent
processesoccuri nthet hy
lakoi
dmembr ane.Embeddedi nthemembr ane
aret hephot osy
stems( IandI I
)thatabsor
blightenergy,elect
roncarri
er
proteins,
protonpumps, andtheenzymeATPsy nthase.
Phot
osy
stems
Phot osyst
emsar est r
uctureswi t
hinthet hy
lakoidmembr anet hathar vest
l
ightandconv ertittochemi calenergy .Itcont ai
nsl argecompl ex esof
proteinsandpi gmentssuchaschl or
ophy l
lsandcar otenoi ds(accessor y
pigment s).Coll
ectivel
y,thepi gmentmol eculescol l
ectener gyandt ransfer
i
tt hespeci alpai rofchl orophy l
l(chl
or ophylla)of t
encal ledspecialpai r
towar dsacent ralpartoft hephot osystem cal ledther eact ioncent er.The
transferisbyr esonanceener gytransfer.Onceener gyr eachest hespeci al
pair,itwil
lnol ongerbepassedont oot herpi gments.I nst ead,thespeci al
paircanact uallyloseanel ect r
onwhenexci ted,passi ng i tto anot her
mol eculeint hecompl excal ledt hepr i
mar yel ectr
onaccept or.Witht his
transfer,t
heel ectronwillbegi ni t
sjourneyt hroughanel ect r
ont r
anspor t
chain.Ther ear et wot y
pesofphot osy stems:phot osy stem I I(PSII)and
phot
osyst
em I( PSI
).Thespecialpairofphot
osyst
em I
Iiscal
l
edP680,
whil
ethespeci
alpai
rofphot
osystem Ii
scal
ledP700.

Notethatl
ightreacti
onisdependentonani
nputofr
adi
antener
gyand
i
nvol
vesfourprocesses/
stages.
I
. Phot
ochemi
cal
exci
tat
ionofchl
orophy
ll
/Phot
oexci
tat
ion
I
I. Oxi
dat
ivecl
eav
ageofwat
er(
phot
ooxi
dat
ion)
/Phot
oly
sis
I
II
. For
mat
ionofATPbyEl
ect
ronTr
anspor
tChai
n(phot
ophosphor
ylat
ion)
I
V. For
mat
ionofNADPH(
phot
oreduct
ion)
I
. Phot
ochemi
calexci
tat
ionofchl
orophy
ll
/Phot
oexci
tat
ion
Inthe fi
rstser i
es ofreacti
ons,li
ghtf al
ls on phot
osystem I
Iand the
chl
orophyllpigmentswi t
hinphotosystem I
Iabsor btheenergy,andthe
el
ectr
onswi thinthem areexci
ted.Theelectr
onsmov etoahi gherener
gy
l
eveland ar et r
ansfer
red tothe primaryelect
ron acceptorwit
hinthe
phot
osystem.
I
I. Oxi
dat
ivecl
eav
ageofwat
er(
phot
ooxi
dat
ion)
/Phot
oly
sis
Astheseelectronsmov et oanel ectr
onaccept or,theyneedt ober eplaced.
Enzymesont het hylakoidmembr anebr eakwat ermol ecul
esinto,hydrogen
i
ons,oxygenat oms,ener gizedelectrons.Theener gi
zedelectronsr eplace
theexcit
edel ectronst hathav ebeent ransfer
redt ot heprimaryel ectron
acceptor
.The oxy gen gas exi t
st heleafthrough t hest
omat aand i s
rel
easedintot heenv ir
onmentandt hehy dr
ogeni onsar ereleasedi nside
the thyl
akoid membr ane.The r eplacement of t he electron enabl es
chl
orophyll
tor espondt oanot herphot on.
I
II
. For
mat
ion of ATP by El
ect
ron Tr
anspor
t Chai
n
(
phot
ophosphor
ylat
ion)
Theel ectronsr el
eased f r
om phot osystem IIent erachai n ofpr ot
eins
knownasel ect r
ont ransportchai n(ETC) .Theymov ef r
om PSIItoasmal l
l
ipid-solublemol ecule,plastoquinone( Pq),andthent oapr ot
eincompl ex
call
edcy tochr omeb6f .Theel ectronsar ef i
nal
lyt r
ansf er
redtoacopper -
containingpr oteincalledplastocy anin(Pc)beforebei ngacceptedbyPSI .
Ast heel ectronsf lowbet weenPSI IandPSI ,
theyloseener gytotransl
ocate
+
Hi onsf rom t hest romai ntothet hyl
akoidlumen.Si ncetheyhav elost
energybef orear r
ivi
ngatPSI ,theel ectronsarer e-energi
zedatPSI .This
processal lowst heabsor pti
onofanot herphotonbyt hePSI .
Theaccumul atonofH+i
i onsinthethy
lakoi
dl umencreat
esaconcentr
ation
+
gradientofH i onsacr
osst hemembr ane,thusgenerat
ingaf or
ceknown
ast heprot
onmot i
veforce.Atr
ansmembr aneprotei
ncalledATPsynthase
+
helpstheH i onstoret
urntothestr
oma, produci
ngATPf rom ADP+Pi.
Thef ow ofH+i
l onsf r
om t
helumen,anar eaofhi ghconcent r
ati
on,tothe
stroma,anareaoflowconcentr
ati
on,ofchloropl
astthroughATPsy nt
hase
i
sknownaschemi osmosi
s.ThisprocessofATPsy nthesi
sisknownas
photophosphor
ylat
ion,as li
ghtprovides the energy to carry outthe
procedure.
I
V. For
mat
ionofNADPH(
phot
oreduct
ion)
Ast heel ectronf r
om t heel
ectrontransportchainar
rivesatphot osystem I
,
i
ti sre-energizedwi thanotherphotoncapt uredbychlorophyllandreleases
electr
onst hatf l
owt oani r
on-contai
ningpr otei
ncal
ledf er
redoxin(Fd)and
+
thent oNADP r eductase,afl
av opr
oteinlocatedonthest r
omalsi deoft he
membr ane, t
hatcat al
y zest
hef ormati
onofNADPH.
Chemi
calEquat
ion
2H2O+2NADP++3ADP+3Pi
→ O2+2NADPH+3ATP
The energy-
carr
iermolecul
es,ATP,and NADPH produced i
nthe l
i
ght
react
ionareusedint
hesecondphaseofphot
osynt
hesisortheCal
vi
ncycl
e
toassemblesugarmolecul
es.
TheLi
ghtI
ndependentReact
ions/
Ther eactionsar epower edbyt heATPandNADPHf ormeddur ingthel i
ght
react i
onsofphot osynthesis.Here, car
bonat
omsf r
om CO2endar efi
xedor
i
ncor por atedi nt
oor ganicmol eculesandusedtosynthesizecarbohy drat
es.
Unliket hel ightreactions,whicht akeplacei
nthethylakoidmembr ane,the
react i
onst akeplacei nthest r
oma.I npl
ant
s,car
bondi oxide(CO2)ent ers
thei nteri
orofal eafv iapor escalledstomat
aanddiffusesi nt
ot hestroma
ofthechl oroplast.
React
ionsoft
heCal
vi
ncy
cle
TheCal
vi
ncy
cler
eact
ionscanbedi
vi
dedi
ntot
hreemai
nst
ages:
a.The fi
xati
on of car
bon by t
he car
boxy
lat
ion of r
ibul
ose 1,
5-
bi
sphosphate
b.Thereduct
ionoft
hef
ixedcar
bont
obegi
nthesy
nthesi
sofhexose
and
c.The r
egener
ati
on of t
he st
art
ing compound, r
ibul
ose 1,
5-
 bi
sphosphat
e.

1.TheCar
boxy
lat
ionReact
ion
The firstst
ep oft he cycl
ei nvol
ves the carboxyl
ati
on ofribul
ose-1,5-
bisphosphat
e,RuBP, a5-
carboncompound, bycarbondioxi
deinatwo-step
reacti
ont oproducesanunst abl
esix-
car boncompoundt hati
mmedi ately
spli
tsinto2mol ecul
esof3-phosphogl
ycerate.
The r
eact
ion i
s catal
yzed by r i
bulose 1, 5-
bisphosphate
car
boxyl
ase/
oxygenase (
RUBI SCO).Thi
simpor
tantreact
ionist herate-
l
imit
ingst
epinhexosesynthesis.
2.Reduct
ionPhase
Theenzy mephosphoglycerat
ekinasecat
aly
sest hephosphor ylati
onof3-
PGA byATP.1, 3-Bi
sphosphogly
cerate(
glycerate-1,3-
bisphosphate)and
ADP ar e t he product
s. The enzy me gl yceraldehyde 3- phosphate
dehydrogenasecat
alysesthereducti
onof1,
3BPGAbyNADPHt opr oduce
Glycer
aldehyde3-
phosphate
3.Regener
ati
onst
ep
I
nt hi
sphase,t her eact
ionsconsistofaser iesofal dolcondensat
ion,
dephosphor
ylat
ion,andtransket
olasereacti
onst hatl
eadtothegenerati
on
ofafive-
car
bonsugar -
phosphate–r i
bulose-
5-phosphate(Ru5P).
Some mol ecules of 3-phosphoglyceral
dehyde are conv er
ted into 3-
dihydroxy
acetone,whi ch combines wi th 3-phosphogly
ceraldehyde t
o
produce f r
uctose 1, 6- bi
sphosphate. Fruct
ose 1,6- bisphosphatei s
conv er
tedintofructose6-phosphateinpresenceofenzy mephosphat ase,
someofwhi chi sconv ert
ed i
nto glucoseorst arch( i
nchl oropl
ast
)or
sucrose (in cytosol)and some ar e used tor egenerater i
bulose 1,
5-
bisphosphate.
I
n additi
on to sugar and st
arch,G3PandDHAPcan be conver
ted
i
ntoaminoaci
dsandfat
tyaci
ds,whicharebui
l
dingbl
ocksf
orpr
otei
nsand
l
i
pids.
Not e:Eacht urnoft heCal vincy cle“ f
ixes”onemol ecul
eofcar bonthatcan
beusedt omakesugar .I
tt akest hreet urnsoft heCalvi
ncyclet ocreateone
mol ecul
eofgl yceral
dehy de-3phosphat e.Aftersi xtur
nsoft heCal vi
ncycle,
twomol eculesofgl yceraldehyde- 3phosphat ecanbecombi nedtomakea
glucosemol ecule.Eacht ur noft heCal vincycl ealsousesup3ATPand2
NADPH i n t he pr ocesses of r educi ng ( addi
ng elect r
ons t o) 3-
phosphogl yceric acid t o pr oduce gl yceraldehyde-
3 phosphat e, and
regeneratingRuBPsot hatt heycanacceptanewat om ofcarbonf r
om CO2
from theai r
.Thi smeanst hattopr oduceasi nglemoleculeofgl ucose,18
ATPand12NADPHar eused