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Photosynthesis in Higher Plants
Photosynthesis in Higher Plants
Photosynthesis in Higher Plants
SINHI
GHERPLANTS
POI
NTSTOREMEMBER:
Photosynthesi
s:Photosynthesisi
sanenzymer egulatedanaboli
cprocessofmanufact
ure
oforgani
ccompoundsi nsidet hechl
orophy
llcont
ainingcell
sfrom car
bondioxi
deandwater
wit
ht hehelpofsunl
ightasasour ceofenergy.
Hi
stor
ical
Per
spect
ive:
JosephPr
iest
ley(
1770):
Showedt
hatpl
ant
shav
etheabi
l
ityt
otakeup
CO2
fr
om at
mospher
eandr
eleaseO2.
JanI
ngenhousz(
1779):
Rel
easeofO2
bypl
ant
swaspossi
bleonl
yinsunl
i
ght
andonl ybyt hegr eenpar tsofpl ants.
Theodor e de Saussur e( 1804): Wateri s an essent i
alr equirementf or
photosynthesist ooccur .
Juli
usVonSachs( 1854):
Greenpar tsi npl antpr oducegl ucosewhi chis
storedasst arch.
T.W.Engel mann ( 1888):
Theef fectofdi f
ferentwav elengt h ofl ighton
photosynthesisandpl ottedthef ir
stact ionspect r
um ofphot osy nthesis.
C.B.Van Ni el( 1931): Photosy nthesisi s essent i
all
ya l ightdependent
reacti
oni nwhi chhy drogenf rom anoxi disablecompoundr educesCO2 to
form sugar.Hegav easi mplif
iedchemi calequat i
onofphot osy nthesis.
Hill(
1937): Ev oluti
onofoxy genoccur sinlightreaction.
Calvin(1954-55): Tracedt hepat hwayofcar bonf i
xati
on.
HatchandSl ack( 1965):
Discov eredC4pat hwayofCO2 fi
xation.
Si
tef
orphot
osy
nthesi
s:
Phot
osy
nthesistakesplaceonlyi
ngr
eenpar
tsoft
heplant
,most
lyi
nleaves.
Wit
hinaleaf,photosynthesi
soccur
sinmesophyl
lcell
swhichcontai
nt he
chl
oropl
asts.
Chl
oropl
astsar etheactualsit
esforphotosynthesis.
Thethyl
akoidsinchloropl
astcontainmostofpi gment sr
equir
edf orcaptur
ing
sol
arenergytoiniti
atephotosynt
hesis.
Theme mbr anesy stem (
grana)isr esponsi
blef ortr
appingthelightenergy
andforthesy nt
hesisofATPandNADPH.Bi osynthet
icphase(darkreacti
on)
i
scar
ri
edi
nst
roma.
Pi
gment
sinv
olv
edi
nphot
osy
nthesi
s:
Chl
orophy
lla:
(Br
ightorbl
uegr
eeni
nchr
omat
ogr
aph)
.Maj
orpi
gment
,act
as r eaction cent re, invol
ved in trappi
ng and conver
ti
ng li
ght int
o
chemi calener gy.
Chlorophy l
l b: (
Yell
owgr een)
Xanthophy lls:
(Yell
ow)
Carotenoid: (
Yellowtoy el
low-
orange)
Inthe blue and red
regionsofspect
rum showshi
gherr
ateofphot
osy
nthesi
s.
Whati
sli
ghtr
eact
ion?
Li
ghtr
eact
ions ort
he ‘
Phot
ochemi
cal‘
phase i
ncl
udes l
i
ghtabsor
pti
on,
spli
tt
ing ofwat er,ev ol
ut ion ofoxy gen and f ormati
on ofhi gh energy
compoundl i
keATPandNADPH.
LightHarvesti
ng Compl ex es (LHC):
The lightharvesti
ng complexes are
made up of hundreds of pigment mol ecul
es bound to pr
otei
n withi
n the
photosyst
em I(PSI)andphot osystem II(PSII
).
Eachphotosystem hasallt hepigment sexceptonemol ecul
eofchlorophyl
l‘a’
formingali
ghtharvesti
ngsy stem (antennae).
Thereacti
oncent r
e(chlorophy ll
a)i sdiff
erentinboththephotosy
stems.
Photosyst
em I(PSI):
Chlorophy ll‘
a’hasanabsor ptionpeakat700nm ( P700).
Photosyst
em II(PSII
):
Chlor ophyll‘
a’hasabsor pti
onpeakat680nm ( P680).
Pr
ocessofphot
osy
nthesi
s:
I
tincludest
wophases-Phot ochemicalphaseandbiosynthet
icphase.
Photochemicalphase(
Lightreact
ion):
Thisphaseincludes-lightabsor
pti
on,
spl
i
tti
ngofwat
er,
oxy
genr
eleaseandf
ormat
ionofATPandNADPH.
Bi
osy
nthet
icphase(
Dar
kreact
ion):
Iti
sli
ghti
ndependentphase,
synt
hesi
sof
f
oodmat
eri
al(
sugar
s).
Theel
ect
ront
ranspor
t:
I
nphot
osy
stem cent
rechl
orophy
llaabsor
bs680nm wav
elengt
hofr
edl
i
ght
causingel ectronst obecomeexci tedandr eleaset woel ect
ronsf r
om t he
atomicnucl eus.
Theseel ectronsar eaccept edbyprimar yel
ect r
onaccept ori.
e.
ferr
edoxin.
The el ectron f rom t he f err
edoxi n passed t o el
ectr
on t ransport
system consi sti
ng cy t
ochromes.
Theel ectronmov edi ndownhi l
lint ermsofr edoxpot enti
albyoxi dati
on-
reducti
onr eactions.
Finall
ytheel ectronr eachedphotosystem- I
.
Simultaneousl yelect r
onreleasedfrom phot osystem-Iisacceptedbyelectron
acceptor.
Electr
onhol ecr eatedi nPS-Iisfi
l
ledupbyt heelectronfrom PS-II
.
Electr
onf rom PS- Ipasseddownhi l
landr educeNADPi ntoNADPH+ +H+.
ELECTRONTRANSPORTCHAI
N
Phot
oly
sisofwat
er:
PS-
IIl
ooseel
ect
ronscont
inuousl
y,f
il
ledupbyel
ect
ronsr
eleasedduet
o
phot
olysi
sofwat er
.
oH+, ghtandMn2+ -
Wateri
sspliti
nt (O)andelectr
onsinpresenceofl
i andCl
.
Thi
salsocreatesO2 t
hebi-pr
oductofphotosynt
hesi
s.
Photol
ysi
stakesplaceinthevici
nit
yofthePS-II
.
2H2O→ 4H+ +O2 +4e-.
Phot
ophosphor
ylat
ion:
Thepr
ocessoff
ormat
ionofhi
gh-
ener
gychemi
cal
s(ATPandNADPH)
.
NonCy
cli
cphot
ophosphor
ylat
ion:
Twophotosy
stemsworkinser
ies–Fir
stPSI
Iandt
henPSI
.
Theset
wophot osy
stemsareconnect
edthr
oughanel
ect
ront
ranspor
tchai
n
(
Z.Scheme)
.
+
ATP
and
NADPH +H
aresy
nthesi
zedbyt
hispr
ocess.
PSI
and PSI
I ar
e
f
ound i
n l
amel
l
ae of
gr
ana,
hence t
his pr
ocess i
scar
ri
edher
e.
Cy
cli
cphot
ophosphor
ylat
ion:
Onl
y
PS-I
works,
theel
ectr
onci
rcul
ateswit
hint
hephot
osyst
em.
I
t happens inthe str
oma lamell
ae
(possi
blel
ocati
on)
because i
nthi
s
r
egi
on
PS-
II
and
NADPr
educt
ase
enzy
mear
eabsent
.
Henceonl
yATPmol
ecul
esar
esy
nthesi
zed.
Chemi
osmot
icHy
pot
hesi
s:
Chemi
osmot
ic hy
pot
hesi
s expl
aint
he mechani
sm ofATP sy
nthesi
sin
chl
oropl
ast
.
I
n phot
osy
nthesi
s,ATP sy
nthesi
sisl
i
nked t
o dev
elopmentofa pr
oton
gr
adi
entacr
ossamembr
ane.
Thepr
otonst
hatar
epr
oducedbyt
hespl
i
tti
ngofwat
erar
eaccumul
ated
i
nsi
deofmembr
aneoft
hyl
akoi
ds(
inl
umen)
.
Ast
heel
ect
ronmov
est
hrought
hephot
osy
stem,pr
otonsar
etr
anspor
ted
acrossthemembr ane.
NADPr eductaseenzy mei slocatedont hestr
omasi deoft hemembrane,
alongwi t
helectronsfrom theacceptoritremovesH+f rom thestr
omaduring
reductonofNADPH+H+.
i
Thiscreatespr otongr adi
entsacrosst hethyl
akoidmembr aneaswel lasa
measur abl
edecr easeinpHi nthelumen.
ATPasehasachannelt hatal
lowsdiffusionofprotonsbackt ost
romaacross
themembr ane.
Thisrel
easesener gyt
oact i
vateATPaseenzy met hatcatal
ysesthefor
mation
ofATP.
Bi
osy
nthet
icphasei
nC3pl
ant
s:
ATP and NADH,
the pr
oduct
s of
li
ght
react
ion ar
e used i
n sy
nthesi
s of
food.Thef i
rstCO2
fi
xati
onproducti
nC3pl anti
s3-phosphoglyceri
caci
dor
PGA.
I
n some ot herplantsthe f
ir
ststabl
e productis an organi
c aci
d cal
l
ed
oxaloacet
icacida4-Ccompoundhenceiscall
edC4pl ant
s.
TheCal
vi
ncy
cle:
TheCO2
accept
ormol
ecul
eisRuBP(Ri
bul
osebi
sphosphat
e).
Thecycl
icpathofsugarfor
mati
oniscall
edCal
vincycl
eonthenameof
Melvi
nCal vin,t
hediscover
eroft hi
spat
hway .Calv
incyclepr
oceedsint
hree
st
ages:
Carboxyl
ation:
o Ca r
boxylat
ionist
hefixat
ionofCO2
i
nt oastabl
eorgani
cinter
medi
ate.
o CO2 combineswithRibul
ose1,5bi sphosphat
et ofor
m 3PGAi nthe
presenceof RuBi
sCoenzy me.
Reducti
on:
o Theseareaseri
esofr
eacti
onsthatl
eadtothefor
mati
onofgl
ucose.
o 2 molecul
es ofATP forphosphory
lat
ion and t
wo ofNADPH for
r
educt
ionperCO2
mol
ecul
efi
xed.
o Thef
ixat
ionofsi
xmol
ecul
esofCO2
and6t
urnsoft
hecy
clear
e
r
equi
redf
ort
hef
ormat
ionofonemol
ecul
eofgl
ucose.
Regener
ation:
o Re gener
ati
onoft
heCO2
accept
ormol
ecul
eRuBPi
scr
uci
ali
fthecy
cle
i
stocont
inueuni
nter
rupt
ed.
o Regener
ati
onst
epsr
equi
redoneATP f
orphosphor
ylat
iont
ofor
m
RuBP.
Hencef
orev
eryCO2
mol
ecul
eent
eri
ngt
heCal
vi
ncy
cle,3mol
ecul
esofATP
and2mol
ecul
esofNADPHar
erequi
red.
TheC4
pat
hway:
Plantsthatareadapt
edtodr yt
ropi
calregionshavetheC4
pat
hway .
C4 oxal
oaceticaci
disthefi
rstCO2
fi
xationproduct
.
The se plant
s have speci
alt ype ofl eafanatomy,they tol
erat
e hi
gher
t
emper
atur
es.
Thel
eafhast
wot
ypesofcel
l
s:mesophy
llcel
l
sandBundl
esheat
hcel
l
s
(Kr
anzanat omy ).
Ini
ti
all
yCO2 istakenupbyphosphoenolpy r
uvate(PEP)inmesophy l
lcell
sand
changedtooxal oaceti
cacid(OAA)i nthepresenceof
PEPcarboxyl
ase.
Oxaloacetateisr educedtomal ate/asparatethatreachesint
obundl esheath
cel
ls.
Inthebundl esheat hcell
stheseC4 acidsar ebrokendownt orel
easeCO2 and
a3-carbonmol eculei
.e.pyr
uvicacid.
TheCO2 r
eleasedi nthebundlesheat hcellenterstheC3cy cl
ebecauset hese
cel
ls ar e r ich i n enzy me Ri bulose bi sphosphate car boxyl
ase-
oxygenase (RuBi sCO).
The pyruvat ef ormed inthe bundl e sheath cel
lt r
ansport
ed backt ot he
mesophy l
l cell
,getphosphorylat
edt oform phosphoenol py
ruvat
e.
Phot
orespi
rat
ion:
Thel
i
ghti
nducedr
espi
rat
ion(
evol
uti
onofCO2)i
ngr
eenpl
ant
siscal
l
ed
phot
orespi
rat
ion.
Act
ivesi
teofRuBi
sCOhasacti
vesi
teforbothO2
andCO2.
I
n C3 plant
s some O2 bi
nds wi
th RuBisCo and hence CO2
fi
xat
ion i
s
decr
eased.
I
n t
his pr
ocess RuBP i
nst
ead of
bei
ng conv
ert
ed t
o 2mol
ecul
es of
PGA,
bi
nds
wit
h O2
to f
orm one mol
ecul
e of
PGA andphosphogl
ycol
ate.
I
nthephot
orespi
rat
orypat
hwayt
her
eisnei
thersy
nthesi
sofsugar
,norof
ATP.Ratheritresultsinther eleaseofCO2wi thutil
i
zationofATP.
I
nthephot orespirator
ypat hwayt her
ei snosy nthesisofATPorNADPH.
Ther
eforephot orespir
ati
oni sawast eful
process.
I
nC4 pl
antphot orespirat
iondoesnotoccur :
o RuBi sCOenzy mei spr esentint hebundlesheat hcel l
s.
o Pr i
mar ycar boxylationi st akespl aceint hemesophy llcel
lbyPEP
car
boxy lase.
o CO2 suppliedtobundl esheat hcellbyC4 acidinter
medi ate.
o He nceC4 plantsarephot osyntheti
call
ymor eef fi
cientthanC3plant
.
LawofLi
mit
ingFact
ors:
I
fachemi
calpr
ocessi
saf
fect
edbymor
ethanonef
act
or,
theni
tsr
atewi
l
lbe
det
ermi
nedbythefact
orwhi
chisnearestt
oitsmini
malv al
ue.I
tist
hef
act
or
whi
chdi
rect
lyaf
fect
sthepr
ocessi
fitsquanti
tyi
schanged.