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CHAP TER 13 Telegram : prat1bllayadavb1olo9y

.PHoTosYNTHESis IN H10H ER. PLANrs

13.1 Whatdo we All animal s including human beings depend on plants for their food. Have
Know? you ever wondered from where plants get their food? Green plants, in fact,
13.2 Early
have to 'make or rather synthesise the food they need and~ other organisms
Experim ents
depend on them for their needs. The green plants make or rather synthesise
the food they need through photosynthesis and are therefore called autotrophs,
13.3 Wheredoes You have already learnt that tqe autotrophic nutrition is found only in plants
Photosynthesis and all other organisms that depend on the green plants for food are
take place? heterotrophs. Green plants cany out 'photosynthesis', a physico-chemiGa)
13,4 How many process by which they use light energy to drive the .synthesis of organic
Pigments are compounds. Ultimately, all living forms on earth depend on sunlight for
involved in energy. The use of energy from sunlight by plants doing photosynthesis ls
is important due to two reasons: it
Photosynthesis? the basis of life on earth. Photosypthesis
is the primary source ofall food on earth. It is also responsible for the release
13.5 What.is Light
of oxygen into the atmosphere by green plants. Have you ever tlwught what
Reaction?
would happen ifthere were no oxygen to breath? 1bis chapter focusses
on
13.6 Tite Electron the structu re of the photosynthetic machinery- and the various reactions
Transp ort that transform light energy into chemical energy.

13.7 Wherea rethe 13.1 WHAT DO WE KN-ow?

ATP and NADPH
Used? Let us try to find out what we already know about photosynthesis. Some •
J3.8 'The C4 Pathwa y simple experiments yo1:1 may have don~ in
the earlier classes have shown
. . n that chlorophyll ~een pigment of the leaf), light and qo2 are required for
'
13.9 Pfwt.oresp!ratio photos ynthesi s to occur.
13.1 O Fact.ors .You may have carried out the expertment to look for starch formation
affecting fn two leaves - a variegated leaf or a leaf that was partiall y covered with
Phot.osvnthesis black paper, and exposed to light. On testing these leaves
for the presence
. of starch it w~s clear that photosynthesis occurred only in the green parts
of the leaves in the presence oflight. . •

.
r ,I
 ,; 'PHOTOSYNTI-IESIS IN HIGHER Pl.ANra'.•,

Another expertment you may have carried out

where a part of a leaf is enclosed in a test tube
containing some KOH soaked cotton (which
absorbs CO2}. while the other half is exposed to air.
TI)e setup is then placed in light for spme time. Oh
testing for the presence of starch later in the two
pl~ts of the leaf, you must have found _that the
exposed part of the leaf tested positive for starch
while the portion that was in the tube, tested
negative. This showed that CO2 was required for
photosynthesis. Can you explain how this
conclusion could be drawn? ,outube : prat'"'h y d • LM
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13.2 EARl-Y EXPE,RIMENTS Telegram : pratibhayadavbll,lt' ....
It is interesting to learn about those simple
experiments that led to a gradual development in
oi:tr understanding of photosynthesis.
Joseph Priestley (1733-1804) in 1770
performed aseries of experiments that revealed the
essential role of air in the growth of green plants.
Priestley, you may recall. discovered oxygen in
. 1774. f!iestley observed th~t a candle burning in •
a closed space - a bell jar. soon gets extinguished
(Figure 13.1 a. b. c. d}. Similarly, a mouse would
_soon suffocate in a closed spac~. He concluded that le) ~; ::_. ·d
a burning candle or an animal that breathe the air Figure 13.1 Priestl :.?. ()
• . ' ey s experiment
both somehow, damage the arr.1But when he placed a mint >- >-
1
~rune bell jar, he found that the mouse stayed alive and Pthant in the en ::;;
continued to bum. Priestley hypothesised as foUows: Plant e candle ci:l -"• _0
the air whatever breathing animals and burning candle-s s rest0~ > ii
.:...,;.:::-- • - - ... w remov ro -o >
~an you imagine how Priestley would have conducted th ~. ' g;_ -~
using a candle and a plant?· Remember, he would need t e experunent _g

i
o rekindl .c .a cu
candle to test w~ether it ?urns after a few days. H w
9
. e the , i ·.:: @
ways can you think of to light the candle without disturb~ y dlfferent e•
th
Using a similar setup as the one used by Priestley, b:; e set-up? • • ...
once in the dark and once in the sunlight, Jan Ingenh by placing it .8 a §
showed that sunlight is essential to the ~ 9 ) -g -; g>
purifies the air fouled by burning candles or hr thtliat somehow
ea ing - ,
Ingenhousz in an elegant experiment With an aquatic I J 1[ I _
1
in. bright sunlight, small bubbles were forme aroun~ showed tha~ - ;!{14 ~I::,~ '-·~
while in the dark they did not. Later he identified these e een Parts ku~
oxygen. Hence he showed that it is only the green Part ~Ubbles to be of '
0 th
could release oxygen. Youtube : Prat·1..1.. y e Plants th t
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 It t t:il abo ut 185 4 that Juli us von Sac hs prov
was no un ided evidence
for production of glucose when plants grow Glucose is usu ally stor ed
star ch. His late r stud ies showed that the• b as
gree n su stanc e in Plants
(chlorophyll as we know 1t . now) i 1 ted in special bod ies Oater Call
s oca ed
chloroplasts) within plan t cells. He found that th
e green part s in plan ts Is
w·h ere g1uco se is made, and that the glucose is usu ally stor ed as starch
Now consider the interesting experiments .
don
Ufl43 1QQ9J. Using a pris m hes lit li ht into its es by T.W Engelmann
ec~ al com one fit
and Enen illuminated a green a, a op 01;;,
P aced m a suspension
o aero 1c ac ena . e acteria were used to dete ct the site s
of1a;
evolution. He observed that the bacteria accu 1
mul
of blue and re o . 1e split spectrum. A first acti on spec
trum of
p otos yPY 3~s ~as thus desc~-~~8-f It rese
mbles roughly the absorption
spectra·of chlorophyll a and ~ M1sc;,\lssed in
section 13.4).
By ~v~ ~~~ e.., o! .. century the key feat ur:s of plant
photosynthesis were l'mown, namely, that
plan ts could use hgh t energy
to mak e carbohydrates from CO~ and wate
r. The empirical equation
repr esen ting the tota l process of photosy
nthesis for oxygen evolvmg
organisms was then understood as:
•

where [CH p] represented a carbohydrate

. . . . -. (e.g., glucose, a six-carbon
sugar).
A milestone contribution to the understand
ing of photosynthesis was
that mad e by a microbiologist. Cornelius
van Niel (1897-1985), who,
bas ed on his stud ies of purple and green
bac tena , dem ons trate d that
pho t~sy nthe sis is esse ntia lly a ligh t-de
pen den t reac tion in which
hydrogen from a suit able oxidisable compou
nd redu ces carb on dioxide
to carbohydrates. This can be expressed by:
t\\Gt, -~OI'\ pyG.
~. on t St it1 1t
. K_ •~ ltl
cta.do phc,(A, _plants 8i0 is the hydrogen donor and is
A.lit, f>., ~ - lo not release 02 during pho oxidised to 0 2. Some

1'
tosynthesis. When
A,2.~fobA d,t :{ H2S' .instead
oge n don or for purp le an~ green sulp hur
flt\ .lld "\ .J .lL (, bac teria , the
,roduct is sulp hur or sulphate depending
.,. ,a i \ h u ~ i ~ "'" on the organism
.
LI.: i, l
t\~
.Hence,.he inferred th~t ~he 02~volved by
P, the green plant
l n~t
B\ (J. I J4' Yf NAf' {f I 111 IV\,, J -techniques. from carb on dioxide. 1his was late r proved by using
The correct equation, that would represent
the
I~ o "'.. fr.t s rf'o. HL
~ss of photosynthesis is therefore:
. ob s< ~ ..~ . but <!, ._.. 6CO , +12H,O Light ,c,H .,o, t6H ,0+ 60,
l ,,a JJ ~'1 J.t :Jl
Bl IA e, IR.L 4
fl-~~ L
I' h .
0 s repr~se_nts gl:1cose.
The 0 released from water; this
, f.sing radio isotope te·chni_ques.2Note thatisthis
t

is not a single
f1~0 ~t r) -+ .~ t y,,01, - -. ,~;
·:1
_
_ _,._ ,.. ... ,,- =- ~r
o"-'. "' O
NAOr
4- 11 Jt r! ,-. .-- fl\ .~~ ~~
P£j[_ •
• •
Xtn, ,
 n leaf or 'in the chloroplasts', o O
y0 u would of course answer: in 'the gree ,._ __ _ ~· L J (f(v ,,,_
8. You are definitely right. +-
based on wha t you earlier read in Chapter ( / ""-
leaves of plan ts but it does so
Photosynthesis does take place 1n the green t Jf(_ /Y) f ~.
you name some othe r parts. e R rn1;t
r# \

also in othe r green part s of the plants. Can ~u~rf,r ~f( m f Modi. ti c [ rid\ f''- "7 s 1rJ~ '7
where you thin k pho tosy nthe sis may occ ur?1 - ,/
the me~ phy ll cells 1n the
. You would recollect from previous unit that -h:, tA)C U,,\.
Usl,llllty the cbloroplasts ajign p{).JtPJ..deJ
leaves, have a large'number ofchloroplasts. - W ,
. suc h that they get the
themselv es alon g the wall s of the mes ophy ll cells
t. Wh eTT :do ~yo u:'t hi~ oou:trd:Oflole!Y
/...,o..vJ h..cflt
opti mum qua ntit y of the inci den t ligh ftYi c);Y\~ ,
ed wi~h ~heirflat s':1 'f~e s~~ U.o ._tl u; w'!ifff? Cl ~~znl
chloroplasts will be align vJ~
the incident l~d H,., .., s11 J :sJQ: fm;i~ ·:·
When wou ld they be perpendzcular to ast in Cha pter 8. Wit hln ° O 0 0 0 0

You have studied the stru ctur e of chloropl

em con sisti ng of gran a, the
the chloroplast there is membranous syst
ure 13.2). The re is a clea r
stro ma lamellae, and the matrix stro ma (Fig
The mem bran e syst em is
di~s ion of labo ur within the chloroplast. er:r f
also for the synt hesi s ofATP f~
responsible for trapping the light energy and -fo . I~ .
s syn thes ise sug ar. whi ch in
and NADPH. In stroma, enzymatic reaction / CJ. Ot 5.. - f .
s, since the are directl Ii t r)
turn forms starch. The former set of reaction 7 J. . J , ~ r . : ~ r t_
ermc reactions). The latte r
driven are called lig t reac tion s p o oc h hi
ent on the pro duc ts of ligh t
fI '
c ••
are not directl Ii ht d1iven bu re de end , ~t 7_ - J
and NAD Hen ce, ~o disti ngu ish the a er ey are
reactions ATP . .
tions). However, this sho uld bJ O
by convention, as dark reactions (carbon reac _ g _
darlrness or that they are 1,1ot\.,JX .J~
o '.i

not be construed to mea n that they occur in '.l) •

,
light-dependent.
• (>)

• .• Out er m~m br ifi.J J..,.~ .IMf -

•
~= --- --I nn er mem bran e -;-- ~• [
~, 2.-<
---" ~-: --S tro ma l lamella

<--- ++- -Gr ana

---,< +--S trom a

~~- Rib oso me s
0

Youtube : Pratibha ••• le1 ~y Star ch gran ule

lnsta ID : @pratibhayada •
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p i d drop let
• .,·
• ·,\ ion of
Figu re 13.2 Diagrammatic represen
t~tion 9f an elec tron micr ogra ph of a sect
.• chloropla~t :Pra~gha Yadav Biology
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 ' . Looking at plants have you ever wondere d why
Chloroph yll b and how 1here are so many shades of green In
their leaves - even in the saine plant? ·We can
Caroteno ids look for an answer to this question by trying to
separate the leaf pigment s of any green plant
Cbloropli: through paper chroma tograph y. A
chromat ograp~ separati on of the leaf pigments
shows that the colour th~t we see in leaves is
not due to a single pigment but due to four
pigments : Chlorophyll a (bright or blue green
in the chromat ogram), chlorop hyll b (yellow
green), z:anthop hyU. (yellow) and carotenolda
(yellow to yellow-orange). Let us now see what
roles vartous pigment s play in photosyn thesis.
Pigments are substanc es that have an ability
to absorb light, at specific W,t~J en~~o u
guess which is the 1JJo sr'U8 ,I~t
pigment in the world? Let us study the graph
showtng the ability of chloroph yll a pigment to
absorb lights of different wavelen gths (Figure
' 13.3 a). Of course. you are familiar with the
•• : ; -Rat otosyn wavelen gth of the Visible spectrum o_f light as
. _,Ab
~~~O R.
•f l':l,"hfm _Figure 13.3a can you determin e the
J:V9Pelength /colour oflight) at which chlorophyl!
a shows the maximu m absorpt ion? Does it
show a7?-other absorpti on peak .at any other
~le~gt bs t.DD?.lf yes, which one?
6'61.-,rrru look at Figure 13.3b showing the
~~;... ..~=~ :--=-- :~~ wave hgths at which maximw n photosynthesis
500 600 700 occurs in a plant. Can you see that the
.(c) wavelen
400 gth of light tn nanometres (run) wavelen gths at Which there is maximum ·
ab
sorptlon by chlorophyll a, i.e:. in the blue and
Graph showing the absorptlqn the red
3 regions, also shows higher rate of
Fi,ut"e 1s. • spectrum of chlorophyll a. band photosyn thesis. Hence, we can conclude that
the carotenoids c~Iorophyll a is the chief pigment associated
Graph showing action with photosy nth. t li'in
15-~ s ectrum of phot9synthesis esis·:iBtf.t.;QY &ffl'1iw{1. aectmfd'ie
f'ipfe Gpraph showing 13 3
action - c can you ·s~ th""cthe rb is ·n comnlPft
e is.Sc s ectruro of photosyn thesis one t • •~ ,c--~61li<m·. ui sm,,
- o-one overla.R: .bJM™fJC{si~~b,figfel~W,11
ytglJJ' • ~perimp osed on .absorpti ~n spectrum of chlorop hyll a and ·the action
s trum of chlorophyll a . ~~Ctn..Lm.....of,.JJbnto • \j
spec •. • ·youtube : Praltbna Yaaav t1reiV\:I, synthesi s? ,,Ui ,
: • ,.· lnsta \0 :@pratibhayadavbiology • 1
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 Cl> ·- .,,-p~,..,-;_,;/

~.::....:~:.-.:.::...:.1:.·.:-i..i-,~'::'~rWf':-~~ 211 ffi,1d[S:b"~=--

b->1- to g> <;_
i>- j I g
(J\NvV ':'~__/JL-17r~.,., ' .Q

These graphs, .together, show that most of the photosynthesis takes

ro >- -o co

Glace in t h e ~ the spectrum: some photosynthesis

oes take place at the other 'Yavelengths of the visible.spect:lJ!..m. Let us
see how this happens. Though chlorophyll ts the major pigment
:§
m

a.-~
...:.
ro ..c n:: w
:§
0
responsible for trapping light. other thylakoid pigments like chlorophyll (_l: ?.- ';
b xanthophylls and carotenoids, which are called accessory pigments
' •
(I)
.aOrou
E .!:
also absorb light' and transfer the energy to chlorophyll a. Indeed• they .a:::,-(l)C:
o, .l!l
not only enable a wider range of wavelength of incoming light to be ittilised -§!. 8.
~or photosyntesis but also protect chlorophyll a fn~to-oxidatian. •• .
r • mi g
j
I-~•
'1ti l
l(hl~

T
13.5 WHAT 1s LmHT REACTION? 1 tu ;i. . w~
Jlco ,:..,,-_,._
Light reactions or the 'Photochemical' phase I ! ,.l, t) 9 1,UJ.,UiV'.'.'
jo.~----
include light absorption, water splitting, oxygen . ._ :::: . "' _ra."flf
nt r ,,., •
release, and the formation of high-energy @ -~~r
chemical intermediates, ATP and NADPH.
Several protein complexes are involved in the
process. The pig~ents are org~~ed into two
discrete photochemical light. harvesting
complexes (LHC) within the Photosystem I (PS -<
I) and Photosystem ll (PS ll). These are named Photon c~_,,_ntr~e'_o;,_.':o' ( {)tJ, A)
in the sequence of their diicovery, and not in :
the sequence in which they function during the ~: ? g .
light reaction. The LHC are made up of ..a~ •' ' Pigment
hundreds of pigII?-ent molecules bound to . molecules
proteins. Each photosystem has all the pigments
(except one molecule of chlorophyll a) fonrung • •<8L.,
a light harvesting system also called antennae
. to make
W ~aw
-.

(Figure 13.4). These pigments help . ure 13.4 The light l (;-: - ~-
-· .,., •
1
photosynthesis more efficient by absorbing ,~~;complex
different wavelengths of light. The single chlorophyll - -<
the. reaction centre .. '.fhe reaction centre 1•8 dif'e a rn~lecule forms
• , • • 1• rent i b
photosystems. 1n PS I the reaction centre chlorophyll ah n °th the
peak at 700 run,. h ence• is called P700; while in PS II itasanab
h SOrption
maxima at 680 run. and is called P680. _ as absorption
•• , Youtube : Pr;,1:"ha Yadav
13.6 THE ELECTRON TRANSPORT lnsta ID •r;_ · c1tibhavswt~
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• 1

In photosystem II th~ reaction centre chlorophyll a ab •~y .

wavelength of red light causing electrons to becoiii"e ) ...1r1:r-2.011
into an orbit farth.~r from the atomic nucleus The·
• • •
exci e
seele tr0
'~
A>
\"P,
up by an electron acceptor which passes them to c ns are Pi k
. an electrona c ed • •
·. . . !ranaport •
 212 l--~~- ....._•..,.- ...
' ..J;:;. ..11,:-, --·.;

.
.
~~1 -
B~o~Y.,~
l
system consistin g of cJochro mes (Figure
Photosyste m n Photosyste m I 13.5). Tilis movement of electrons is downhill,
I ----- k,c{lf\J..t·f / in terms of. an oxidatlon- reductio~ or redox ,
e acceptor ".&.(NADPH potential scale. The electrons are not used up
ADP+IP ATP NADP+ as they pass through the electron transport

.t chain, but are passed on to the pigments of

photosyste m PS I. Simultane ously, electrons
in the reaction centre of PS I are also excited
. when they receive red light of wavelengt h 700.
-1.,,:,.~ • • ,: si. run and are transfe rr~othe r accepter
• ; :..:_:: molecule that has a ~ e d o x potential.
, ;, i
:-.:=;-< These electrons then are moved downhill
·: tn • again, this time to a molec1:1Ie of energy-ric h
7,:::::;;;;;;;;.;.,..,_ _ _ _.;,~~~~· NADP'". The addition of Uiese electrons reduces
~~.a:,:;~•.:.:·,,_, H20-+ 2e·+ 2H+ +~or? NADP+ to NADPH + H+. Th.is whole scheme of
l""-
r
J:!• .It, ..0

• Nl=,Cr"'t - 1o transfer of electrons, starting from the PS II,

~ure 13.5 Z scheme of light rea~~01s;- uphill to the acceptor,
g g' 8 ~-
the electron
1:f g~ transport chain to PS I, excitation of electrons,
.Q g transfer to anotfih~c ceptor, and finally down hill to NADp+ reducing it to
'ero -oi:u :.o·. .Q0 ' >- NADPH + H+ is called the Z scheme. due to its•charact erstic shape (Figure
"fil Cl? cc :g C;D 13•5). Tilis shape is formed when all the carriers are placed in a sequence
2
.c: ,g .c UJ .g .g
-o C1J ca
on a redox potential scale.
,
,f!ro:QZ ~~"g
a. ro "-'
ro
a.. =
,._ r.:::a,
a. (1)
....•• E
a - .c.o :::-
'"'
@1§
,u
..c ·- .c
:g
13.6.1 Splitting of Water
'" E _, C1J a. co You would then ask, How does PS II supply electrons continuou sly?The
.oQ O::©)c.
3~g>"E •••••• .electrons that were moved from photosyste m II must be replaced. This is
=>cn-o E
~.E~U _!gt! achieved by electrons available due to splitting of water. The splitting .of
'§; g> water is associated with the PS II; water is split into 2H... [OJ and electrons.
.-f!s{! Tiiis creates oxygen, one clfue
net products of photosynt hesis. The
electrons needed to replace those removed from photosyste m I are provided

Go
. by photosyst epi II. _______ ·

NEET- d O.,J ON Q_ . ~4W +O, + a •:,:,

()· f)..tl in NO'O .. ~c..lt (, ".'to emphasise here that thewate~ splitting com~lex is associated
t,· (N~f#\, (;}rII, whi~h itself,~ physically located on the mner side of the
,t\~ho oft I pf the thylakoi~ Then, where are the protons and 0 Jormed

f os..-
1 ~~n. 1k'
... ,JL . • 'M¼
- ~Pleased - in
1
V or on the outer side ofthe membr,ane?•
the.Jilmen?
2

~® •.
\N c... .<I

iJ. t ~OU-<ct yclic and Non-cyclic Photo-ph osphoryl atio~

' I. U•4"'k- tisms have the capability of extracting energy from oxidisable
md store this in the form ofbond. energy. Special substance s like
l
1 1- • tis energy in their chemical bonds., The process through which
;J · 0t~jfN'
;3- kl~
~¥() >~ f
 PH01:0SYNTHESIS IN HIGHER Pl.Ams a

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ATP Is synthesised by cells (in mitochondria and lnsta ID : @pratibhayadavbiolo9y ,.,., ---:
chloroplasts) is named phosphorylation. Photo- Telegram.: pratibhayadavbioltiJytosyst~~ 9_
phosphorylation Is the synthesis of ATP from · I t ,. .,.

e
ADP and inorganic phosphate In the presence of
light. When the two photosystems work in a $ ( e-aecep'?[ ', : ::
sertes, first PS II and then the PS I, a process called
~n-cycUc photo-phosphorylation occurs. The
two photosystems are connected through an • 1-,-----, .J J•
Electron
electron transport chain, as seen earlier-in the transport
z scheme. Both ATP and NADPH + W are system
synthesised by this kind of electron flow (Figure ...... -
13.5).
When only PS I is functional, the electron is Chlorophyll
circulated within the photosystem and the P700
phqsphorylat~on oc;:~urs due to cyclic flow of
electrons (Figure 13.6). A possible location Figure 13.6 Cyclic photophosphorylation
where this could be happening is in the stroma tfmrn"--
lame'!:Iae. :rWhil~e:..l·t~h~e~'l:n~·e~m~!W,l~LJJ:U.W=~........~~""'-'"41/.C-.JUUJ.LU~
and W, II the troma l~ell~cfflembranes lack PS II as w
W-: pr-a. •
- '<a /), J t - t ~
educ e'e -~~ctron does not pass on to NAD~but is NfrPr ,....,..
cycled back to the PS I compl~x through the electron transport chain
(Figure 13.6). The cyclic flow hence, results only in the synthesis ofATP,
but not ofNADPH + tt•. Cyclic photophosphorylation also occurs when
only light of wavelengths beyond 680 run are available for excitation.
. . . Youtube : Pratibha Yadav B~Y
13.6.3 Chem1osmotic Hypothesis lnsta IO :@pratibhayadavbiok>9Y
Let us·now try and understand how actually
chloropl~st. Toe chemiosmotic hypothesis has been put forward to explain
the mechanism. Llke in respiration in ph~tosynthesis too, A1P synthesis is
!inked to develoe...ment,of a proton gradient across.._a membl'a.fil. This time W
these are the membranes of thylakoid. There is one difference though h
. ere
th roton accumulation is towards the inside of the membrane · , th
, .e., m e
1
In respiration, protons accumulate in the intennembrane space of
the mi,tochondria when electrons move through the ETS (Ch t
aper 14).
Let us understand what causes the proton gradient
across the
membrane, We need to consider again the processes that take
1
pacecturtng
the activation of electrons and their transport to determine th t
es eps that
cause a proton gradient to develop (Figure 13.7).
(a) Since splitting of the water molecule takes place on th inn
e_ ersideof
the membrane, the protons or hydrogen ions that ar ,,___....;;;.;.:=
the splitting of water accumulate within t h ~ f theproduced by
' • . e thylakoids.

,,J
 ...,~, • B10LOGY';._

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Elec,-,..~ .
-"'-"lem1ca1
Po~ntiaJ
• Gradient

.Stroma ·11,,J.~}
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«I· ~ropi,M.t ff ..

hi~.,u+ 'YlUW\bUf Gfr,3.7 ATP ~ynthesis through c~e~iosmosi~

S Ql,A, /,.., • ntA, :trons move thr~ugh the pho~~tems: protons are transpo~ed
• • I s' th
(' ~,"',me,
. ., mbrane. This ·happens because the prtm~rv
'#•_!JlH'V accepter of
Ih •
-J

fPlli~~!:h i~!i?, n s the • e of them . e

• " - - knt\(Al, Gom ,J .1 "- ,f~t"lifl.,1':-1~l~~t1:er _not to an electron canier but to an H cam~r
I ITJ I r~' [e, this molecule removes a proton from the strom hile
l · t +to
f
Ml\- rporting an electron. When this molecll:le passes on its electron
l
l • Lu.MW\ 't "Hl•
M..
l
1.,, • J
~o, OI,
electron carrier on.theJnoec~ oftbe.J!lembr~e, the proton
ased into the inner si~e or thdJgmen s~of the ~embrane._
'4 • "JIIJ.l'f M(~), • Wk.c ADP reductase enzyme is_ located on the ~~l;ide of the
---------------:11..--=::_Jbrane . Along With electrons that come f r ~acceptor of.
• ele~trons of PS I, protons are necessary for the reduction of NAO?- to
·._A NADPH+ ff+. These protons are al~o re~oved from t h ~
.3..V';- Hence, Within the chloroplast, protons in. the strom d~se in
·.; . ~C.L.!!umber. w • e in the lumen ere is accum a ·on o pro ons.
1
·s crea es
7
• . a proton gradient across the thyl oi mem rane as we as-a ~easurable
decrtase in pH in the lumen. .
Why are we so interested in the ·proton gradient? This gradient is
important because ~t is the breakdown of this gradient that leads to the
synthesis of ATP: .Toe gr~dient is bro OWi) due to the movement of
protons across the membrane to th throu h the e
 215
PHOTOS\'l'ITHESIS IN HIGHER PLANrS

cnannel of t h ~ f the ATP thase. The ATP synthase enzyme consist s

of two_parts: one ccµled the F0 I embedded in the thylakoid membr ane
and fonns a transm embran e annel that carries out facilitated diffusi on~
of proton s across the membrane. The other portion is called CF, and
protrUdes on the outer surface of the thylakoid membr ane on the side
that faces the stroma . The break down of the gradien t provides enough •
energy to cause a confqmiational change in the CF1 particle of the ATP
synthase, which makes the enzyme synthesise several molecules of energy-
. packed ATP. ~---i ~------.

Che·m.1osmosis require s rane a ~roton ,pump } a proton i' .

gradien,Ll!pd ATP syntha se. nergy s used to pump proton s acrostLa
membrane, to create a gradien or a high concentration of proton s within
the thylakoid lumen. ATP syntha se has a channe l that allows diffusion of
protons back across the membrane: this releases enough energy to activate·
ATP syntha se enzyme that catalyses the formation ofATP.
Along with the NADPH produced by the movement of electrons, the
ATP will be used immediately in the biosynthetic reaction taking place in
the. stroma , responsible for flXing CO2 , and synthe sis of sugars .

13.7 WHERE ARE THE ATP AND NADPH USED?

We learnt that the produc ts oflight reaction arelATI>. NADPH and oJ Of

these 0 2 diffuses out of the chloroplast while ATP and NADPH are used to
drive the processes leaqing to the synthesis offood, more accurately, sugars.
This is ~1:J>Jasynthetic ph~~(?Jip~otosynthesis. This proces s does not
directly Wpend on the pre~~~c~p!Ji!gpt but is depend ent on the produc ts
of the·ligh~~~£tj.99.! !,-,Y ·~~~ ~~H, besides CO2 and Hp. You may
wonder how this could be vertlie : is simple: immediately after light
becomes unavailable, the biosynthetic process continu es for some time,
and then stops. Ifthen, light is made available, the synthe sis starts again.
Can we, hence, say that calling the biosynt1ietic phase as the dark
reaction is a misnomer? Discuss this amongst yourselves.
Let us now see how the ATP and NADPH are used in the biosyn thetlc
pha~e. We saw earlier that CO2 is combined with 1¾0 to produc e (CH2 0)n
or sugars . It was of interes t to scienti sts to find out how this reactio n
proceeded, or rather what was the first product formed when CO2 is ta.ken
into a reactio n or fixed. Just after world war'II, among the several efforts
to put radioisotopes to bene fict~ , the work ~elVi n Calvin is
exemplary. The use of radioac tive 14
C by him m algal photos ynthes is
studies led to the discovery that e first CO2 fixa on produc t was ·a
3-carbo n organic acid. He also contrib uted to working out the comple te
biosynthetic pathway: hence it was called Calvin cycle after. him. The
frrst produc t identified was. 3-phosphoglycerlc acid or in short PGA
'How many carbon atoms doesit h ~ - - - = /
. yl2(,.'t1
 all pla nts have PGA as the flt
. Sci ent ists also tried to Imow whether .. St
oth er pro duc t was fon
pro duc t of co2 fixation , or whether any d •'"ed lt1
r a wi e ran ge of pla nts 1
-oth er pla nts. Experiments conducted ove
where tpe first stab le PrOdu:~ to
the discoveiy of ano the r group ofplants,
, but one which had 4 carb of
CO2 fixation was again an organic acid
ato ms in it. This a_cid was identified ta.be
oxaloacetj.c acJd or OAA, Stn~n
esis was said to be of two Ill e
the n CO2 assimilation dwi ng photosynth
duc t of CO2 fixation is a ca acid
atn
types: tho se plants1rrwhich the first pro
....."f , which the first pro duc t was a C
(PGA), i.e., u'f C3 pathway, and those in
acid (OMJ, i.e., -the C' pathwa-y. ~, These two gro ups of pla nts shoWed•
••
will disc uss late:i:-.
oth er asso ciat ed characterlstics tha t we .
R,y'r),,
13. 7.1 Th e Primary Ac cep tor of CO2
t was asked by the scie ntls ~ Who
Let us now ask ourselves a question tha
tion'. How man y carbon atoms
were struggling to und erst and the 'dark reac
epting (fixing) CO2 , would have 3
wou ld a molecule hav e which after acc
car bon s (of PGA)? 2.-
tha t the· acc epto r molecule
The stu die s veiy unexpectedly showed
bisphosphate (RuBPJ. Dia. any
was a 5-@ Lbn ~et ose sug ar - rtbulose
woriy; the scie ntis ts also took
of you thin k of this pos sibi lity ? Do not
ents to reac h this conclusion.
a lon g tim e anchc;onducted man y expertm
d~ct a C3 acid, the pr1mruy
They_ also believed tha t since the first pro
they spe nt ma ny years trying
acc ept or would be a 2-carbpn compound;
re they discovered the 5-carbon
to iden tify a 2-c arb qn compound befo
RuBP. Youtube : Pratibha Yadav Biology
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13 . 7.2 Th e Ca lvin Cy cle Telegram: pralibhayadavbiology •
the whole pathway and showed
Calvin and his co-workers.then worked out ~
7 d in a cyclic manner; the RuBP was r~ ge
-tha
-- t the pat h~a y.9aerate
operates and where the sugar is
Let us now see how the Calvin pathway
and veiy clearly ·that the Calvin •
syn the sise d. Let us at the out set und erst
nts; it does not mat ter whether •
pa th wa y~ 11;_all pho tosy nth etic pla ,
s (Figure 13.8).
the y hav e(S ,6{ c)o ran yot her J pathway
le can be described under
For eas e ofu nde rsta ndi ng, the Calvin cyc
regeoeratlao ,
thre e stag es: ~bo xyl atio n. reductj_on and
tion ofCO2 into a stable organic
car box yla tion - Carboxylation is the fixa
crucial step ofthe Calvin cycle
intermediate. Carboxylation is•~ mo st
n of RuBP. • This reaction is
whe re CO2 is utilised for the carboxylatlo
which results in the fonnatton •
• catalysed by the enzyme RuBP carf;>ozyiase
yme also has an oxygenation
of two mo1ec:ules oC3 PCA Since this enz
RuBP carb~xylase~oxygenase
activity it would be m~re correct to call it
orRuBisCO.
• : : .:s f ( ~·

liiA ¥ iffi? 556 if§, t

 Atm osp her e

I
0 carb o:x ylat ton
ADP

\ ' '

• • : I ,
'I

,.,,, .- ATP
Red uct ion \ +
'-=' \- NA DPH

l
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P, +NA DP-
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Suc rose , star ch

.
Fig ure 13. 8 The Cal vin cycle pro cee ds in thre e stag (

CO com bine s wit h rtbu lose -1,5 -bis phoes : (1) carb oxy lati on, dur ing wh ich
carb ohy drat e is form ed at the exp ens sph ate; (2) red uct ion , dur ing wh ich
2
\ .
e
and NADPfI; and (3) rege nera tion dur of the pho toch emi call y ma de ~TP
ing whi ch the CO2 acc ept or ripu lose
1,5- bisp hos pha te is form ed agaiJ:_l so -
tha t the cyc le con tinu es
2. Red uct ion -Th ese are a serie13 of
rea ctio ns tha t le.a d to the for ma tion
of ·glu cos e. The step s inv olv e util
isat ion of 2 mo lec ule s of AT P for
pho sph ory lati on and two of NA DPH
for red uct ion per CO mo lec ule
fixe d. The fixa tion of six mo lecu les 2
of CO2 and 6 tur ns of the cyc le are
req uire d for th~ form atio n· of one mo
l~u le_o fglu cos e fro m the pat hw ay.
3. Reg ene rat ion - Reg ene rati on-
of the CO acc ept or mo lec ule Ru
cru cial if the ·cycle is to con tinu e 2 BP is
uni nte rru pte d .. Th e reg ene rati on
step s r e q u t r e ~ r p h
~f;rm ~:u .:J

E
-\:..
• - ••··-· - ,,. 218 f....~.....:.:.•..
...... ' -· -;;::~
.... - ~ -• .J........ -- ---~ '.
',,
r<'\\"' ' . BIOLC>Qt
. ~Q
\-1...(1\,
-\., -_ " \..'\v
......C\"", , every co molec ule enter ing the Calvt n cycle , 3 molecules
Henc e ,or
6 t " ({"'.,. 2
of ATP and 2 ofNADPH are requir ed. It is proba bly to meet this
difference
\l }j. \,.
v-r in numb er of ATP and NADPH used in the dark react
,,
ion that the cyclic
r .uth't-- ~hosphorvlatjnn...takes place.
""~ . .
To make one molecule of gluco se 6 turns of the cycle are
reqmred,
\..J 1/1 Work out how many ATP and NADP H mole: ules will be
requi red to make
o<' -¥ one molecul.e of glucos e throu gh the Calvm pathw
ay.
-~;J . • It migh t help you to unders~and_all of this ifwe look at what
goes in
and what come s out of the Calvm cycle.

In Out - -7
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Six CO2
18 ATP
12 NADPH
One gluco se
18 ADP
12 NADf._ .1.,r0
l -Ji

g g> ~8 Mtt ,{fr PQ <)~ ~~ I

-oo
as.~ ~ g UC.
THE c ~ T ~ P~
t--?~yY· .
_g :§
13.s
11 ·:4 ,l\rOr)TrJ,,,1,w..a
,n.H.."
-g
>- ro -g O?
>-t- Plant s that are. adapt ed to dry tropic al regio ns have the c·
co :§ ro t±1 4 pathway
.c ·- .c menti oned earlie
. r. Thou gh these plant s have the C4 oxalo acetic acid as
,g "§ :§ the first CO2 fixation produ ct they use the C pathw ay or the CalVin
£ @5 5--; as the main b~o syn the~ Tne n, in3 what way are they
cycle
• • ••. E _s from C3 plants'? This 1s a quest ion that you may reas<?nably ask.
different
.g Q • ~- g __ c 4 plant s cl!e special: They have a spe;.ial type of leaf anatomy, they
:5, ,ffi %§ tol_srate higher temperatures, they show a r(rtB P~t9 .\n& \lygJ
at~~ i~,
E: r- <..> they ~ack a pfocess called photarespiratiofnBMRl-~f4W-Je.i-~
~o~¥g@ty

~a ~J'bJ1~~~!i
of bioma ss. Let us under stand these one 01rR u b d 9.::::1.
• • • Study vertical sections ofleaves, one of a1t~·pi~t 1 1C
_at l , I• LA'f '}) plant. Do you TI?tice the differences? Do both have t~ same
types j
•E.--- ,).. . rJ o f (.J mesophylls? Do they have stmilar cells arowuithe vascu lar bundl 0
e sheath?
:i .' Th HJ<. w,J,l Toe partic ularly large cells aroun d the va,scu lar bundle_s of
the C
• • plant s are called bundle sheat h cells, ~d the leave s which
~, •• N O
_ u .L:~ J., anato mv ~rP. r,ai·ct t O h ave 'Kranz' ~at omy, ·Kranz· m~an s 'wreat
have such4
h' and
~i:.'.. -101~ PY!, ;;::-~~tf~ ~Oh. P'f&. ~:~:.:~
ll -~ - ~tt h-< ~,!I'~ large °I®. 'ho soh o~ '"""' vl.ttr v1ou s to
QC. • ~ • Hf ,M M,r ange r ll...
I 7 "~ I'• 0 ~~:a1 (,P Ef) ,·nt fi111•~ 1M"
,Q \to· cut a
·erve the
'
{JJ-.y 1a1, • 5
r,otl f I • e tnter~
l I
(JJ IA(..f_J • ~ IV\ :
l-, .... - - f, 1-I
pec1es of
"o
, ~ ;. , : r I\ ; you ' , iJ,alA..t.. ve under
"
•UI 01~\ ...V
I
. ~.~J ;;r~·s e1,. l r "'~ IT' • ~ascular
@ otO O""- oo;O I C ,,,.,.,.
'f
11,f ~C
. ()(11. 2- lidentify

pho.sphoj~ttrrt 1'\· (Je el~~

ep~os,ho~ul fdr-t{;Jt)~ ,. l1L<'t pl•wat -
 , PHOTOSYNrHESIS IN HIGHER f>u.ms '

pathway that has ,' _

Now stud y the pathway shown in Figure 13.9. This l'ET• ul 1 / 101 :J-
a cyclic process. ~t
been nam ed the Hatch and Slack Pathwar, is again N~ f d., .
us stud y the pathway by listing the steps. phos phoe nol t •
cule
The primary CO2 acceptor is a 3-carbon mole f
phyll cell~. The enzyme N E' T
pyruvate [PEP) and is present in the meso ). 2,3' PV£
PEPcase. It is important
responsible for this fixation is PEP carb oxy ~ or 0
theb esop hyjj £vi½ lack RuB isCO enzy me.Jrbe c acid ll'llff • 'UJ,,.J
to register ehat 1 1
,9AA I~oq ned in the rnesophvll £ell§& PY4. / Ul,I ,
He n form s othe r 4-carb0n
LI
.com
liJ\ ,TI
pounds likecrnalic acid or aspa rtio
ported to the bundle
r •
\l

aciQ_ in tJ?-e.,p~s?e!!pl ~~q~ ~~~ lsfhi ch are trans are bmken down
shea th eel . the bundle shea th cells these 4
f1
C acid~

t w..,eleas CO d a 3-carbon mole cule .-, tU. VI l

- n molecule is tr~s port e~ back to the mesophyll where it
ft/
is converted to P~P again, thus , completmg
the cycle.
s th C r Cabdo •
The CO2 released in the bund.le sheath cells enter
bundle s eath cells are
p ~ a pa~w ay common to all pl~t s. The
,, J. l ,N<'J ~l, t'" Atmospheric CO 2
VN"" fY

J1.. · Mesophyll •
-: V cell

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Bund,le
~h.e~th
.cell •

•~fT--2-0J-O P'II.
~--Gl«t fhc CMVn2 s~ :
!· RuS,sco
ATP l-NADPl1 •
~. RuBi£lo -<.A b,{~v,c,-h'O'rl~ Gu.cl
U \~ ;_~.. •
P(tvrd" 1 tw, •• . . · ·
b· r1.

{), ; ! t"' , Mt.JJ )f½ Jl representation of the Hatch d

"Ir: .I)/ {, W ,IA /'...,- - an Slack Pathway

7h J.w bS ~ ~·a o1 _
RJJKi' .C,A 6 l (j.rv.( '
 '
220 t- -~ ____·-:.__ - -- -----------; ---.;_ ~---:--·:--·, -- --- ,~-: ·-... ..~-- ;__ . .:...:___
~----- ~)A ) .. ,____ _._. ___ ._,.___, ...~-- · ~:=: =
BioWOY
c,lV4 v i ~ck ..\tt1- •
.
rich in an en e Ribulose bispho sptiat e carbo xylase -oxyg en:se
~t- c., pl A;/'\ • sC ). but lack PEPcase. Thus. the basic pathw ay that
result s~
(._,3 f) • \ the formation of the sugars, the Calvin pathway, is common to the C3 an
J; 1 C4 plants. •
N\~ {., 11:,£ C... Did you note that the C...,alvin pathway occur s in all the mesop
~ll
cells of the C 3 plants? In the c plants' it do·es not take place lo lhe
4
mesophyll <:_ells but does so ortly in ~e bundl e sheat.1?: censJ
Youtube : Praiibha Yadav Biology
13.9 PHoTORESPIRATION lnsta ID : @pra\11 hayadavbiology
• rtavbiol
~~~el? anoav
~tihh
Let us try and under stand one morJfJMLitrni efi'trr~ nnpo ~t
diff~rence between C3 and c plant s- Photorespiratlon. To understand
.
photorespiration we have to4know a little bit more about the fi,rst step of
.
the Calvin pathw ay - the first CO fixation step. This is the reaction
where RuBP
·-
combines With co t~ form 2 molecules-of 3PGA, that is
catalysed by RuBisCO. 2

Rl$P + CO2
RuBisCo >2 x 3PGA
RuBisCO that is the most ~bundant enzyme in the world (Do -you
wo~d y?) M>ar actert sed by the fact that Its active site can bind l<l
bo CO d ~ - hence the name. Can you think how this could be
poss· ?RuBisCO has a much greater affinity for CO when the CO : 0
is nearly equal than for 0 2. Imagine what~~1c10 ,h~gp~ 2 2 2
p. if thi~ 'Y~re.not
so! This binding is competitive. It is the rel~·ve c;:pnc~ntration of 0,2 ~d
C0 2th~t determtnes which of the two willI•HDTC,C1 ;l J'('lilU ·
d to.~¢.- e!l:ZYIIle, . ~. . f
.. ' C Plants
:-:-:'. some
~-;-;- ~~ O2 does
- -bind
-:- toRuB
--- isCt1,' ~crri.~'h~e'coa· ~tion is
-
decreased. Here e RuBP instead of being converted to 2 mole)\JJ~~ of
ttJ.1 PGA binds With 0 2 to form one molecule of pho
Tlh l )bpho sphog lvcola te (2 Carboni in a pa way c e p _o orespiration. In
rck~"lnd
the photorespiratoiy pathway, there is neither synthesis of su ars nor of
~O ATP. Rather it results in th,e release of c~ tli. e utilisation ofATP. In _
ffiephotorespiratory pathway there is no synthesis of ATP o:c._NADPH.

(.,,0
._
Ltht< oJi"
: -~ •
•
t
1l!,e biological function of photorespiration is not known

h~ve
In C4 plants photorespir.atlon does not occur.• This is because they
. a mcreases e concen a on of CO at e enzyme
yet. . ,.

\ sit s es p ace w en e 4 afiq P.T tht,.w,e~~93::11 is broken

• ·r- .,~ dow nin the~ tore leh' se Jtbftf~lillts_inincreasing
), the ~trace llular concentration of CO • fn t~. this ensur es that
the
)0\ RuBisCO functions as a carboxylase minim 2
ising the oxygenase actiytty.
Now that you know that the. C plants lack photorespiratlon, you
probably can understand why prod~ctlvtty and yields are better in these
plants. In addition these plants show tolerance to higher temperatures.
dav Bi<ffffi:ased on the above discussion can you compare ptwt~ showing
Youtube : Pratib~a Ya davbi~~c3 and the C4 Pathway? Use the table format given
lnsta ID : @prat1bhaya 'b· I YLJ<Jr~n. and.fill in the
• •
. ratibhayadav
10 og_
Telegram •P NEET-Biology C9urse •
Contact me for

•
 Youtube : Pratibha Yadav Biology
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.~:...:...PHorosmn1Es1sJN HloHERPl.A.Nrs .. ---.....Telegram : pratibhayadavbiology_._,:..___.· ~.::......;J
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Ntt T-,:1016 er a 2 and 3 in this table to highlight the differences

Planb ·

.
CD. Tht IJrO~
•
--~-
whi~ f~fs '+ i
C, Plants
• '
Choose from
~ : ; ,
• •
~cr(fU OJ,/~\(._ , Si( Mesophy~/B~ndle sheath/both- .
l
f ,.,, (rC. -. tv\SG .
' ,..,._,
IXPM~ ( &,
is ·.
y ,

Mesophyll/Bundle sheath /both

Y.i
f • U1-1 J -(J
1 tAu) I ·--f"L
I
O Two: Bundle sheath and
"-L. th/ mesophyll :
turJA, .
. r,.. I \J'tn One: ~esophyll •1
Three: Bundle sheath, palisade, J.
. J,. -J
• CLO J&-«-&t i YA,1,d'ft ~Gp spongy mesophyll
fll RuBP/PEP/PGA ti
"' . Ru pi-rA -hoYl 1
3 5141 3 '1
WhiC.::h ts·the primary coz
fiXS.tl<?~ product
r<n(;C)
fr OIt-fr
C..4L). PGA/OAA/RuBP/PEP
l'

No, of carbons in 'the primary U

f co2 f}xatio~ ·produ_ct . 7 3 I4/ 5 .j
poes_ the p~ant have RuBis~O? 'ti/.:> 1'
I ooes tbe. plant have_PEP G~e? ..
NO \J. _
u.o
Yes/No/Not always
1
W}lich cells in the plant have Yes/No/Not always
? • ,. '3~ l. M h
esop yll/Bundle sheath/none 1
... •
I
Rubisc9~ . -
fiX;tlbn rate under high :
ugbt s.o~ditions . N

Whethei· photoresptratlon }s •
- •
·\:Q,~oi8•, ,.
\rooloi~ . ,
~7ct... 1~ / medium
l)f''i : 9duruoY
HI fl' .ni
i -sent at:tow i.. \I.,'
Jight irltenslties ,tiolOitJ . g /neg(tgm ~J•etimes
p,,,. . .... - q • •
. wtl~ihe~•. photores.ptr~tio~ Js" • .
... - .1],..,1.i-1IJ11tilq: fnG1ot~a1'
....
p ~ b j g h !001.t.mtens!U~s~, ,/ . . , . .. _ tt·igh /negligible/ sometimes
,e_'"f-'J()l,O f '/ A . f .. It••~,-~=~
~-015 ((1(,\ t,'(JYI ~ii,.~ 1 t •.• _'
High/negligible/sometimes

rl" 1
n., i(O en~
0 l\•·. JIV\,nJ [ . Iii
,gh/negllgible/some~es
-cUfircrHtm
a

~-1-o U\IA4 l&.l~·i$,· • ao.:- .

·.£.<tr.lA.n,t\ Wi/ •. - 40 C/20-25C/above 40 c
1v· I .-n
a7 C f' Irv.I\ lh' Ii
Cul Vertt a1
difti c sections of leaves of
-

I • 1 n J\'1 • 0 " \.,,(/' 'T"'f "'""'. . . . .,~"~_:_:;;-""' th erent plants and observe wider
e microsco fi Kranz
,q .
;.,.
"' . . 8 • c,L
1 ••,Vi -D
( _ (, 11
, T. . . . . . . . . . . .,

...._l..!E.,t~ ·••"':~.'!-'h~~
-
.,and list •'h pc or anatomy
c.ol~ ..::'~em in the appropriate
~· d/J- 4- (, t i m9/ . Youtube . P ----==s· ••. •
1 · 1m r - "~ . oo lnsta ID ;@~ Yadav Biology
o-fr_ vO VV'1
v' iele Pflti>havrut.......
'1 ·'.;I, MM . .3 --C Ctfr,.fD~ gram :P r a ~ ~ Y
 • i

. _ _. BroJ,OGY

13.. 10 FACTORS AFFECTING PHOTOSYN THESIS

An understand ing of the factors that affect photosynt hesis is necessary.

"(he rate of photomi,th esis is vety in!Bortant in determint ng the yield of
piantis incluging crop plants. '.Photosyn thesis is under the influence of
several factors. both internal (plant) and external. Toe plant factors include
the number, size:' age and.~ctent ation of leaves, mesophyl l cells and
chloroplasts. internal_9...02 concentrat ion and the amount of chlorophyll.
The plant or internal factors are dependen t on the genetic predisposition
and the growth of thet plant. •• ' '\
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The external factors would include the availabilit y of sunlight,
.temperature, CO2 concentration and water. As a plant photosynthesises,
all these factors will slm.ultanedu~ly affect its·rate. Hence, though several
factors interact and simultaneously affect photosynt hesis or CO fixation,
2
usually <;>ne factor is'tlie major.dlu se or is the one that limits·the rate.
Hence, at any P?int the rate~~~ determine d by the factor availabl~ at
sub-optliiial lev9s.. , . ,. •
• When several fa<_:tors affect any [bioi chemical process, Blackman's
(1905) Law ofLimiting Factors comes into effect. Tilis states the following:
---U a chemical process is .aJfected by more than one factor, .then its
rate wiH· be·det~ d by ,t~ factor which is nearest to its minimal
value: it is the factor ~hich !1-ii~ctly affects the process if its quantity is
changed. :·- :J ••\ • •
For example, despite the presence of green a
Y~be : Pratibha.Yadav Biology leaf and optimal light and co; conditions, ilie
lnsta ID : @pratibhayadavbiology plant may not photosynthesise if the temp~rature
Telegram :_ pratibhayadavbiology is· very low: This leaf, if given the· optimal
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tempera~ e, start photosynth esising..

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8 13.10.1 ~ight·
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} ,--:-~ ~~::= :=:=:= :! ·We_need to distinguish between light quality, light
.8 - intensity and the duratioB of e.wosure to light,
a. . while discussing light· as a factor that affects .
'a photosynthesis. TI)ere is a linear relationship •
• tweeu incident lfght i!J)d COa fixation rates.at .
W~es.~At. higher light intensi~,
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• wfut ttlcf increase
g (Figure 13.10
• •. '
 ' ' i

223
PHOTOSYNTiiES1$ IN HIGHER !>wrrs

lllctdent light bey~nd a point causes the breakdown of chlorophyll and a

decrease in photosynthesis.. _ •

13. 10.2 Carbon dioxide Concentration

carbon dioxide ,is the major limiting factor for photosynthesis. The
concentration of CO 2 is very low in the atmosphere (between 0.03 and
o.04 per cent). Increase in conc~ntration upto 0.05 per c~nt can cause .an
1ncrease in CO2 fixation rates; beyond this the levels can become damagmg
over longer periods.
The C and C4 plants respond differently to CO2 concentrations. At
3
low light conditions neither group responds to high CO2 conditions. At
·highlight intensities, both c3 •and C4 plants show increase in the rates of
photosyn~esis. What is important to note is that the 4 plants show
saturation at about 360 µ1L· 1 while C3 responds to increased CO2
concentration and saturation is seen only b.,eyond 450 µ11· 1. Thus, current
availability of CO2 levels is limiting to the C3 plants.
The fact that C3 plants respond to higher CO2 concentration by
showin~ increased rates of photosynthesis leading to higher productivity
has been used for some greenhouse crops such as tomatoes and bell
pepper. ~ey are allowed to grow in carbon dioxide eru'iched atmosphere
that leads to higher yields. .Youtube : Pratibha Yadav Biology

13_10 _3 _Temperature lnsta ID :@pratibhayadavbiology

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The dark reactions being enzymatic are temperature controlled. Though
the light reactions are also temperature sensitive they are affected to a
. mu~h lesser.extent. Tha<;'blants respond to higher temperatures and
show higher rate of. pho~;uthesis while~lants have a much lower
temperature 0Dtimum. JI'"lv/\\~ \., Q ,'I ,, -. C'r
Th \tV V • •v 1'1. sc·iluoY
• ov f7'/ v
e temp~~ature optimum for.photo~vnthesis of different plants also
depends o th h<>:b ta , • UI GlcTtl
hi n <\~ L .ltll-8'.L~~¥:¥im1n~fM>ted to. Tropical plants have a
_gher temperature optimum than the plants adapted to temperate
climates.

l3.l0.4 Water

Even though water is one of the reactants in the light reactlo th fli
water as a factor is more through its effect on the plant rath nth, e edirect of
on phot th • •
osyn esis. Water stress causes the stomata tO 1 her an .
ectly
.
the CO c ose ence reducing
2 availability. Besides, water stress also mak
reduc1n th es 1eaves Wilt th
g e surface area of the leaves and their met b li . ' us,
. a o c actiVity as well.

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 SUMMARY

is. During this proc ess carbon

Gree n plan ts make their own food by photosyntheS
thro ugh stom ata and used for
dioxide from the atmosphere is taken in by leav.es
h. Photosynthesis take s Place
mak ing carbohydrates, principally glucose and starc
leaves. Within the leaves, the
only in the gree n part s of the plants, mainly the
sts that are resp ?nsi ble for co2
mes ophy ll cells have a large number of chloropla
are sites for the light reaction
fixation. Within the chloroplasts, the membranes
a. Photosynthesis has tw~
. while the chemosynthetic pathway occurs in the strom
tions. In the light reaction the
stag es: the light reaction and the carbon fixing reac
the ante nna, and funnelled to
ligh t energy ts absorbed by the pigments present in
re chlorophylls. Ther e are two
special chlorophyll a molecules called reaction cent
absorbing chlorophyll a P700
phot osys tems , PS I and PS II. PS I has a 700 nm
reaction cent re that absorbs
molecule at its reaction centre, while PS II has a P680
are excited and transferred
. red ligh t at 680 nm. After absorbing light, electrons
NADH. Duri ng this process a
thro ugh PS II and PS I and finally to NAO forming
the tllylakoid. The breakdown
prot on grad ient is created across the membrane of
F part of the A1Pase enzyme
of the prot ons gradient due to movement through the 0
ting of wate r molecules is
rele ases enou gh energy for synthesis of ATP. Split I
ns and tran sfer of electrons
associated with PS II resulting in the release of 0 2, proto
to PS II.
utiJ?ZYIIle, RuBisCO, to a 5-
In the carb on fixation cwJe, CO2 is added.by :Pt1.1~
tW.~s of 3-ca rbon PGA. _This·
carb on com poun d RuBP th~ s converted to ~1moJ~
is the~ converted to suga r by t\le.Ccuvio.o/de
;:~d@!~&~¢i3P is regenerated. Owi ng
reaction are utilised. RuBisCO
this process ATP and NADPH synthesised in the light
plants: photorespiration.
also cata lyse s a wasteful oxygenation reaction in C3
thesis ~ed C4 pathway.
Som e tropicai plan ts show a special type ofphotosyn
takes place in the mesophyll,
In thes e plan ts the first product of CO2 fixation that
is a 4-ca rbon compound. In the bundle shea
th cells the Calvµi path way is carried
out for the synt hesi s of carbohydrates. Yout~be : Pratibha Yadav Biology
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EXERCISES •
Contact me for NEET-Biology Course

her a plantis c or c ? Why and

!

I. By looking at a plan t extemapy can you-tell whet

bow? / 3 4 •

t can you tell wheth~r a plant ts

2. By looking at which internal structure of a plan
C3 or c4? Explain. •
out the biosynthetlc _ Calvin
3. Even thou gh a vefY fe~ cells in a C4 plant carry
discuss why?
path way , yet they are highly productive. Can yo~
 a carboxylase and oxygenase. Why do
4. .RuBisCO is an enzyme that acts both as
atlon inc, plan ts?
you thin k RuBisCO carrtes out more carboxyl
conc entr atio n of Chlorophyll b, but
5. Suppose there were plants that had a high
tosy nthe sis? The n why do plan ts
Jacked chlorophyll a. would it cari y out pho
ents?
have chlorophyll band·othe r accessoiy pigm
dark frequently yellow, or p~e gree n?
6. Why is the colour of a leaf kept in the
• •
Which pigment do you think is more stab le?
shad y side and com pare it with the
7. Look. at leaves of the ~ame plan t on the
ed plan ts kep t in the sunl ight with
leaves on the sunn y side. Or, compare the pott
es that are- dark er gree n ? Why?
those in the shade. Which of the'm has leav
the rate ofphotosynthesis. Bas ed on the
8.. Ftgure 13.10 snows the effect ornght on
graph, answer ~e foll o~g questions:
(a) At which poin t/s (A, B or C) in the curve
a
is light limiting factor?
(b) What could be the limiting fact or/s in
region A?
curve?
vpdniflc)·Wliat 'd~ cang='.P.: {_epresent on the
:
XFg_ Give comparison betvlctn the following
~~:.,,c(a) c;~ d e~"path¾iy/ ,. • .
. (b) Cyclic and non-cyclic photophospho
iylation
(c) Anatomy ofle af in C3 and c, plants
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