Photosynthesis in Higher Plants

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PHOTOSYNTHESI

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

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

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

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.

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