1 3 .

PHOTOSYNTHESIS N
HIGHER PLAINUS
on sunlight 1or energy.
0
liv depend ing forms
OtOSynthesis is a physico-chemical process by which Ultmately, all
yn thesis
-

Importance of Photos
ECCn plants use light energy (solar énergy) synthesisc 1o
source
food on earth.
ofall
l is the primary
organic compounds. into the atmosphere.
Photosynthesis is the basis of life on earth. I t releases oxygen

PHOTOSYNTHESIS
EXPERIMENTS RELATED WITH when plants grow
and it is usualy
Glucosc is produced
1. Variegated leaf experiment o

stored as stlarch.
Take a variegated leaf (or lcaf partially covered with black bodies (chloroplasts).
o Chloroplhyll is located in special
paper) that was exposed to light. of planl.
o Glucose is made in the green parts
Test the leaves for starch. It shows that photosynthess 1909)
T.WEngelmann (1843
occurs only in grecn parts of the leaves in presence of light. Experiments by components
into its spectral
2. Half-leaf experime nt He split the ight using a prism in a
A part of a leaf is enclosed in a test rube containing KOH iluminated a
and alga (Cladophora) placed
green
soaked cotton (which absorbs CO). suspension of aerobic bacteria. evolution.
the sites of Oz
The other half of leaf is exposed to air. The bacteria were used to detect
accumulated mainly in the
Place this serup in light for some time. He observed that the bacteria
spectrum.
-Test the leaf for presence of starch. Exposed part shows region of blue and red bght of the spit is.
positive for starch and portion in the tube shows negative. It was a first described action spectrum of
photosynthes
a & b.
This proves that CO2 is required for photosynthesis. It resembles the absorption spectra of chlorophyl
it is discovered that plants
By the middle of 19th century,
EARLY EXPERIMENTS from CO& H;0.
use ight energy to make carbohydrates
Experime nts by Joseph Priestley(1770) Empirical equation of the process of photosynthesis
is
-Priestley performed experiments to prove the role of air in
co, +,0 nt[CH,01+0
the growth of green plants.
He discovered oxygen in 1774. Where, [CH:0] represents a carbohydrate (e.g., glucose).
van Niel (1897-1985
e observed that a candle burning in a closed bell jar gets
Experiments by Cornelius in
mouse scon suffocated in condusied some stucies
extgished. Similarly, a
an ei (ierubiuiogisi)
thai a burning candle or a
ciose. päce. He conchuded purple and green bacteria.
light-dependent
bralarng.onimal damage Hhe Oust He found tihat
He denonstrated that nhotosynthesis is a

suitable uidisabic
He placed a mint plant n tne same bell jar. reaction in which hydrogen from a

the mouse stayed alive and the candle continued to burn. compound reduces CO» to carbohydrates.
He hypothesised that plants restore to the air whatevei -This can be expressed by:
breathing animals and burning candles remove.

2H,A CO, 2A+CH,O-H,O

(1730-1799)
Experime nts by Jan Ingenhousz
it once in gTeen plants, H»0 c
He conducted the same experiment by placing
902.
-

the dark and once in the sunight.

essential to the plant for /Purple and green sulphur bacteria use H;S as H-donor. So
- He showed that sunlight
animals.
the 'oxidation' product is sulphur or sulphate and no O2 is
purifying the air by burning candles or
fouled
with an aquatic plant. It
produced
He repeated this experiment te inferred that the O2 evolved by the green plant
small bubbles were formed /Thus,
showed that in bright sunlight, comes irom H0, not from CO2. This was later proved by
around green parts while in the dark they did not.
ysing radiosotopic techniques.
to be of oxygen. Thus he
Later he identified these bubbles Therefore overal correct equatiorm for photosynthesis is:
showed that oniy the gree: part of plants release O2.
Sachs (1854) GCO+12H,o cH,0, +6H,O+60,
Experiments by Julius von

-He provedthat
PHOTOSYNTHESIS: SITE AND PIGMENTS
The membrane system traps light energy and synthesise
ROotS Photosynthesis occurs in green leaves & other green parts. ATP and NADPH IL s caled light reactions.
of
ave Chloroplasts present in the walls of mesophyll cells I n stroma, ntymatic reactions incorporate CO: into the
Chlotou leaves. It helps to get optimum quantity of incident ight.
plant for synthes izingsugar,which in turn forms starch It
Chloroplast contains a membranous systeèm. It consists of
is caled dark reactions. It does not mean that they occur
frce pa grana, stroma lamellae and luid
stroma.
in darkness or that they are not light dependent.
membrane-bound sacs called
Each granum is a group of
TnaSfor thylakoids (lamellae). They coptain leaf pigments.
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To-a eoPyun ACt 0n pe CHtum
SPace NADPH>2 SAT?
top al tek ma Pho Sythaet¢1 FA H f:TAz

Melt USA Temp

 O

Cha-Css a205 N
Outer membrane 2 Rte
Innor membrane

Stromal lamnella Graph showing

* SFY Grana actiorn
spectrum of
.
ronia photosynthiesis
** RIbosomes

Starch granule

Llpld droplat oluvavelength of Hght (uun)

PIGMENTs INVOLVED IN PHOTOSYNTHESIS R n l e of plotosynthesls|

51plon
Graph showing
action
Pigments are substances that have abilty to absorb light, at
spectrum of
specific wavelengths. photosynthesis
Chromatography shows the folowing leaf pigments: superimposed

o Chlorophyll a (bright or blue green in chromatogram) on absorption

spectrum of
o Chlorophyll b (yellow green) chlorophyll a
Accessory 700
o Xanthophylls (yellow) pigments
400 SOO
Wavelengh of light iu nanometres
o Carotenoids (yelow to yellow-orange *
- Functions of accessory pigments: Photosystems
called
Pigments are into two Photosystems
organised
o They absorb light at different wavelength and transfer I1 (PSI). These
ae

the energy to chlorophyll a. Photosystem I (PSI) & Photosystem

named in the sequence óf their discovery
o They protect chlorophyl a from photo-oxidation. a and accessory /PST -C3On
- The absorption spe ctrum & action spectrum coincide Each photosystem has a chlorophyll
pigments bound by proteins.
show ing that photósynthesis is maximum at the fom
closely molecule of chlorophyl a)
bluc & red regions of the spectrum.
-All pigments (except one
- The graphs also show that chlorophylla is the chief a light harvesting Pdmar.accepleu
pigment associated with pBotosynthesis.
complex (LHC or
antennae).
Chlorophyll a Single chlorophyl a acis Phot Reacuon
Graph / cenlre

A -Carotenoids showing
absorption
as reaction centre.
- In
centre
PS I,
absorbs
the rcaction
ngni a
Piment
molecuies
Chlorophyll b chlorophyll 700 nm, and so called

A 7K)
, b&
carotenoids P700.
In PS I, the reaction
Light harvesting compiex

(n) Wan elength af light (nm) entre absorbs light at 680 nm,
and so caled P680.

LIGHT REACTION (PHOT0CHEMICAL PHASE)

Photosystem I
Photosyslem
reactions include light absorption, vater splitting, n ,CL
-Light
and formation of ATP & NADPH (high Eacceptor NADPH tH'
oxygen release
energy chemical intermediates). acceptor ADP+ ATPA
NADP
The Electron Transport
wavelength, Electron
II absorbs red light of 680
nm
When PS transpor
electrons are excited and trans
fèred to an electron acceptor. system
them to a chain of electrons
The electron acceptor passes
trausport system
consis ting of cytochromes. LHC
This movement of electrons is
downhil, in terms of redox
potential scale. LHC
I.
-

The electrons are transferred to the pigments of PS H20- 2e+ 2H"+ [O

in PS I arealso excted when
Simultaneously, electrons Z scheme of light reaction
700 and are transferred to
they receive red light of nm

another accepter molecule having a greater

redox potential. Splitting of Water (Photolysis)
molecule of - The water splitting complex in PS II is located on the inner
These electrons are moved downhill
to a

NADPH+H'.
NADP". Asresult, NADP" is reducedto
a sideof thethylakoid membrane. g n eoloton
Transfer ofelectrons from PS II to PS I and finally
downhill Water is spli into H', [O] and electrons.
NADP' is called the Z scheme, due
to
to its zigzag shape, 2H04H' + O + 4e
in
This shape is formed when all he carriers
are placed a
-
These electrons are needed to replace the electrons that are

sequence on a redox potential scale. moved from PS I1.

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 Tn ho0gvtieesrH,o isoneted to form 0, and Co Heduted to 5ora s 1°g orn
Carn bohyo ratel Non CHcue it movLe e bkecH Ve
from the
renovcs a prolon
mnolecule
Thc prolns (H°) are used to reduce NADP to NADP 01 Carricr. Sothis an
clectron.
When this

wlilc trunsporting on
Oxygen is liberaled as a by-product ofphotosynthesi. stroma
cleciron to the eleciron
carrier

on its nto
Inolceue passes is rclcascd
The PS I1 provides electrons needed to replace those proton CnytyLoha
Ihe inner sicde of the niembranc,
reimoved from PS 1. membrone.
the lumen of the FHD Mo
is locatcd Ontne suSPemONe,
Photo-phosphorylation o The NADP reductase enzy1mc comjng
Along with elkctrons
The synthesis of ATP by cellk (in mitochondrin sicde of tle meTmhTane. ncccssary to rcducc NADP
chloroplasts) is called phosplhorylation. Tau Plosei ron sprolons are
ADI' removcd from the Slroma.while
An Ma Photo-phosphorylation is the sythesis of ATr from Thesc protois arc also in
thic stroma decrcase in numbcr,
u o no in chloroplasts in presence of light. Tlence, protons in
This crcatcs a proton
M toChon d a

It occurs in 2 ways: Non- cyclic and Cyclic. the Jumcn, protons

arc uccumulatcd.
and decrcascin pH proton
SoeMo a) Non-cyclic photo-phosphorylatlon
It occurs when the two photosystlems work in a scrics, (fist
gradicnt neross thc
in the lumcn
thylakoid membranc
8radfent
helP+o
PS I and then PS 1) through an electron transport chain as ATPsynilcsis through NADP A4afe
chemlosm9sls prtoton
Parth
seen in the Z scheme.
Here, ATP & NADPH + H' are synthes is ed. otvsystc1t ytorhrunics
PlUsySte
**ADIH
MOHe,
I t is non-cyclic process because the clectrons
lost by P'S fonce
AP ATP Siye
a
to NADP",
II do not come back to it but pass on

b) Cyclic photo-phosphoryiation
I n SHtoma

lameU a e
- It occurs in stroma
when only PS
Photosystcn1 I
umen

ATp
only PaL lamelae Thylakold
foIis functional. cacceptor membrane

The electron
6 t7
cirCulated within theishi
Suroma
-F
eo6 ATV synthgse*
Larnella
photosystem and the DP
A T P

Electron to release ofenergg

boSn T ATP synthess occurs
tiansport Breakdown of proton gradient leads
due to cyclic flow of L System to the movement ot
PSoA The gradient is broken down due
electrons. across the riembrane to the through h
stroma
protons
- The leme!lee of grana ofthe Foofthe enzyme ATPase.
Chlorophyu trans-membranechannel
have PS I & PS L. The P700
nwo pmrtk-
TheATPase (AT? syntlhase) membraneofand
consists
lamellae forms a trans-
Stroma
o Fo: It is embedded in the
membranes lack PS 11 and NADPreductase. membrane channel that carries out faciltated diffusioFn
on to NADP" but
is cycled back
not
-

Theelectron does pass of protons across the membrane.

transportchain. surface of the thylakoid
toPSI complex through electron NADPH +I). o F: It protrudes on the outer Vnte
-

Here, only ATP is synthesised (no membrane. The energy due to-breakdowir of gradient U
also ocurs when only light Fi particle. It T P
-

Cyclic photophosphorylation causes a

conformational change in the
available.
'avelengths beyond 680
nm are
ATP molecules.
of makes the enzyme to synthesise
Chemiosmotic Hypothesis Thus, chemiosmosis requres a membrane, a proton pump,
syathesis in chloroplast. proton gradient and 4TPase.
It explains mechanism of ATP
a
n across a membrane, to
linked to development of a prot
Energy S used to pump prötons
ATP synthesis is concentration of protons within
across thylakoid membranes. create a gradient or a high
gradient membrë e.
occurs on the inner side of the the thylakoid lumen.
Spltting of water channel that allows diffusion ofprotons
back
in the lumen of thylakoids.
accumulate -
ATPase has a
So the protons are to activate
the photosystems, protons across the membrane.
This releascs energy
As clectrons move through
membrane. lt is due to
the removal that catalyses formation of ATP.
ATPase enzyme
transported across the reasons:
stroma for the following
ofprotons from the located towards tlhe
electron accepter is
o The primary and transfers its eleciron to
duter side of the membrane NADPH
PHASE) - USE OF & ATP
DARK REACTION (BIOSYNTHETIC folows: Immediately after light
and O. It can be verified as
NADPH
- Products oflight reaction are ATP, drive the becomes unavailable, the biosynthetic process
continues
Dark reaction is the use of ATP and NADPH to
If ight is avalable, the
for some time, and then stops.
-

processes for the synthesis

of food (sugars).
synthesis starts again.
on the light but is
-This phase does not directly depend reaction. - CO combines with H0 to form (CH2O)% or sugars.
dependent on the products of the light CO assimilation during photosynthesisis 2 types
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CATtbS
Siorton S4ephen 4 a c k w i

Owicdud e phosPhorat on An naly

 SLACK PATHVIAY
PATHWAY (HATCH &
o Ca pathwny: In this, fiust stuble moduct of CO, fixntion A regons.
is a Cacid (PGA). al'hn Cu to drytropical
*
K prcscnl inn planta adapteel paillhwuy.
mnin hiosyntlhetic
Melvin Calvin studicd nlgnl photosynheis usimy C hey nio (' pnthwny ay ofthe (4plantw
use

le discovcied that the fiust CO, finatiou proudhuct wny he hrpe cell anound tie vrcular bundles
arc
Callca meloPAH
cell, Such anatomy
phosphoglyceric neigl (PGA) n J-cuhon n pganie ne e cnlleel Inrllr shenth Sutune
'wrentl'k runn une aroUnd
o
Cepmthway: In this, fiust stnlhle product is uNNlontgtle Itnn' mntomy('Kran'
forin several layers
acid (OAA), a d-enmlhon (C) upm e Thie bulle shentlh cclh may
the vcular lundlle.
thick a l s
C PATHWAY (CALVIN CYCLE) hnve wpe nnher of chkroplast7,
They spacc.
ll occurs in all photosynthetie plants (C'or pathway) and no intcrcclluar
npcr vinua to gns cxclhange
- Ihas 3 stages:eanboNylation, teduction nnd iepcnetnti01. Slack Pathway
Stops of Hatch and a
Atmosphere
CO; ncceptor is phnsphoenol pyruvate
(PEP)
Prinary 1or
The cnrync
3-carbon molkcule scen in mesophyl cels.

this fixation is 1EPcarhaxylase (P'EPcuse).

CmboruoThe CA acid OAA is formed
The mesophyll cclls lack RubisCO cnryne
in the mesophyl ccls
acid or
ADP
RUB

I0 then forms olhcr 4-carbon acids like malic

transportcd to bundle shcath
cells.
Aspartic ncid. They arc
molelea 3- PGA Atmnsphe ric CO
ROreneOn
emr
ATpC1) Mewplhy
ReuEton NADPJ
1 ATP PhaaphoTol.
HCO pTU
Triose A
phospbate Fxation Regeneratia
P+NADP
C ncid C acid
Sucrose, starch
Plaso
1. Carboxylation of RuBP deamnta

- RuBP (ribulose bisphosphate - a 5-Carbon ketose sugar) Bunde

sheath
=
s theprmey CO; acceptor. two 3-
ccll
-

It is the most crucial step. CO2 is fixed by RuBP to Transpor

of the enzyme RuBP carboxylase. Fixation by
PGA in presence
oxygenation activity It s qck
alta
also has an ncid
-Since this enzyme C
called RuBP carboxjlase-oxygenase
(RuBisCO).
2. Reduction |Decarboxylation+ C acid

I t is a series of reactions leading to the glucose formation.

phosphorylation and two of
Here, 2 ATP molecules for
NADPH for reduction per CO2molcculeare used. In the bundle sheath cells, Ca acids are broken down to
Fixation of 6 CO2molecules and
6 turns of the cycle are
-

release CO2 and a Ci molccule.

the pathway.
needed toremove glucosemolecule fron
one -
The Ca molecule is transported back to mesophyil where it
3. Regeneration of RuBP is converted to PEP again.
It iscrucial for continuation of the cycle. The released CO2 enters the Ca pathway.
to form RuBP. are rich in RuBisCO, but hck
I t requires one ATP for phosphorylation Bundle sheath cells
molecules and 2 PEPcase. Thus Co pathway is common to Ca & Cplants.
Hence for every CO2 molecule, 3 ATP
NADPH areTequired. und
Ca plants are special becanuse:
Js probably to meet this difference in number of ATP o They have a special type of leaf anatomy (Kranz2).
reaction that the cyclic
NADPH used in the dark o They tolerate higher temperatures.
phosphorylation takes place. o They show a response to highlight intensities.
Tomake 1glucose molecule, 6 turns ofthecycle are needed. o They lack photorespiration.
What does go in and come out of the Culvin cycle?
o They have greater productivity of biomass.
Out
6 CO glucose
PHOTORESPIRATION 4E|
rehon 18 ADP In Calvin pathway, RuBP combines with C02.
on18 ATP
12 NADPH 12 NADP RuBP C0, Rublsco2x 3IIGA
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 Rather it
1uRC) is the mosI abundant_enzyine-in the world. Its jence pholorespiration js
a vastefulprocess.
acne Sie can bind to both CO and 0-hence the name.
Causes the relcase of CO:by using ATP.

LOhas 3 ereaicr affinity for COthan for O. This

T In Ca plants, photorespiration doesatnot ocenzyme site. Ths
CO2
the
concentration

competitive. Relative concentration of O2 and increase broken

Dihdng is Can from the mesophyl s
Ca acid
when the
CO detcrmines which one wl bind to the enzyme. takes place mnmises
Ths
In Caplants,some O: bind to RuBisCo. Hence C0% fixation downinthe bundle
cells to release CO2.
Oxygenase activity RuBisCO. productivity and yields
of
form
s decreased. Here RuBP binds with O to one
Due to the lack photorespiration, show

molecule of phosphoglycerate and phosphogbycolate. This are better in Ci plants. In_addition,

these plants
pathway is caled photorespiration. Carb NADPH. (od tolerance to higher lemperatures.
In this, there is no synthesis ofsugars, ATP and
Differences between Ca and Ca plants
plants
Ca plants and bundle sheath
cells
1. Pholosynthesis OCCUTs In nesophyll cells In rresophyll

Kranz anatom yis absent Presen

RuBPis the primaryC0; acceptor PEPisthe primaryC02 acceptor
firststable product
4 3.PGA a 3-C compound is the frst s table product OAA a 4-C compound is the
in bundle shean
in mesophyll and agranal
5. Chloroplasts are ofonly one type (granal) Dimorpiic (granal
Photor spirationis absent or negligible
6. Photores piratonylossis high CO2 ')
compensation point (0-10 ul.
7. High CO; compensation point(25-100 ul.CO,t) | Low C
6.
is about 25°C | Abou 3545
Optimum temperature for photosynthesis productiwly hign.
Piulisyntheticallymore efficient and
9. Photos yntheticallyless efficientand producivity low. aie, sugarcane, amaranth,
sorghum
10 Eg. Rice, wheal, bean, potato |Eg.
FACTORS AFFECTING PHOTOSYNTHESIS
Carbon dioxide Concentration
(plant) factors: The number, sze, age and
factor for photosynthes is.
Internal
orientation of leaves, mesophyll cels and chloroplasts, CO is the nmajor limiting
concentration is very low in the
atmosphere (0.03-
internal CO2 concentration and amount of chlorophyll. CO
concentration up to 0.05%% cause
on the genes and growth
of the plant. 0.04%). Increase in
Plant factors depend this the levels can
CO increase in CO^ fixation rates. Beyond
Fxternal factors Sunlight, temperature,
become clamaging uver longer periods.
Concentralion and water CO2.
light, Ca and Ca plants do not respond high
to
"f At low
Law of Limiting Factors (1905):
a
Black man's i iigh light, thoy show
increased rate of photosynthesis.
more than one factor,
biochemical process is ajfected by
saturation at about 360 ulL.
nearest 1o its minimu plants show
its rale is determined by the factor and
directBy affects the process f plants respond to increased CO2 concentration
value: it is the factor which current
aturation is seen only beyond 450 uL. Thus,
its quantinis changed"
a ailabilty of CO% levels is limiting to the Ca plants.
conditions
E.g.a plantwith green leaf, optimal light CO%
&
if the temperature is very low. If - D to response to higher CO: concentration, Ca plants
may not photosynthesize sh increased photosynthesis and higher productivity.
it wl start photosynthesis.
optimal temperature is given,
Th factis used for somegreenhouse crops (tomatoes, bell
Light in enriched atmosphere.
pep er etc). They are grown CO%
Light quality, light intensity
and duration of exposure
Temperature
to light influence photosynthesis.
and - Dark 2ac tions, being enzymatic, are temperature controled
There is a linear relationship
between ncident ight
Infuer e of temperature on Light reactions is very less.
CO fixation rates at
- The C plants respond to higher temper atures and show
low light intensities.
higher "e ofphoiosynthesis.
At higher ight
aw intensities, the rate -Cplants1ave a much lower temperature optimum.
- The temp rature optimum of plants also depends on their
augalst does not show
furnher crease
habitat. T pic al plants have a higher temperature optinum
than the p nts adapted to temperate climates.
because other factrs
become limiting. Water
Light intensity D
Light saturation Water stre s closes the stomata hence reduce the CO
occurs at 10% of the full sunhght. Hence, except for plants
factar
availability.
shade or in dense foresis, ight is rarely a imting - Water stres also wiks leaves, thus reduce the surface area
n nafure. of the leaves and their metabolc activity.
It
High increase in incident ight breaksdown chlorophyll.
decreases photosynthess
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