(Ch-3, Physology2018

CHAPTER 3:
PHOTOSYNTHESIS
 Life on earth ultimately depends on energy derived from the sun.
 Photosynthesis is the only process of biological importance that
can harvest this energy.
 The name photosynthesis derived from the Greek words phōs,
"light", and synthesis, "putting together"; means literally
“synthesis using light.”
 It is a process used by plants and other organisms to convert light
energy into chemical energy.
 Photosynthesis represents the transfer of matter:
 CO2 from the atmosphere, water from the soil or atmosphere,
 into sugar in the plant and O2 back into the atmosphere.
 It involves oxidation and reduction process.
 The overall process is an oxidation of H2O (i.e. removal of H/e-
to form O2) and
 reduction of CO2 to form organic cpds such as carbohydrates.

Photosynthesis
6CO2 + 6H2O + Energy → C6H12O6 + 6O2
Carbon Water Carbohydrate Oxygen

 The reverse of this process – the oxidation of carbohydrates to form
dioxide
CO2 and H2O - is a spontaneous process that releases energy.
Respiration
C6H12O6 + 6O2 → 6CO2 + 6H2O + Energy
Carbohydrate Oxygen Carbon Water
dioxide
3.1. The Nature of Light
 Light is a form of radiant energy, a narrow band of energy within the
continuous electromagnetic spectrum (EMS) of radiation omitted by the
sun.
 In actual practice light is a mixture of different colors and the EMS

comprises radiations of different wavelengths.
 Light has both wave and particle nature.

 Wave properties of light include the bending of the wave path when
passing from one material (medium) into another.
 The particle nature of light is usually expressed in terms of quanta or

photons
 i.e. discrete packets of energy each having a specific associate

The Nature of Light Cont’d…
 Wavelength (λ) is defined as the distance from peak to peak (or
trough to trough).
 Energy level in each photon is inversely related with λ (E α 1/λ):
longer λ have less energy than do shorter ones.
 The order of colors is determined by the λ of light.
 Visible light is one small part of the EMS.
 The longer the λ of visible light, the more red the color.
 Likewise, the shorter λ are towards the violet side of the spectrum.
 λ longer than red are referred to as infrared (IR), while those shorter
than violet are ultraviolet (UV).
 Solar radiation in the visible range of spectrum from 400 nm (violet) -

700 nm (red) λ known as photosynthetically active radiation (PAR) is
utilized in the photosynthetic process.
 In fact, the λ within this range are the ones absorbed by the
chlorophylls and other photosynthetic pigments in green plants and
algae.
Figure: The electromagnetic spectrum (EMS).

FIGURE: EMS. Wavelength () and frequency () are inversely related. Our
eyes are sensitive to only a narrow range of  of radiation, the visible region,
which extends from about 400 nm (violet) to about 700 nm (red). Short 
(high ) light has a high energy content; long  (low ) light has a low energy
content.
3.2. Photosynthetic Apparatus and Pigments
 When light strikes an object, the light can
 be transmitted (pass through the object),

 be reflected from the surface, or
 be absorbed.
 For light to be absorbed, pigments must be present.
 Pigments (light-absorbing molecules) absorb light selectively,

with different pigments absorbing different s and reflecting
others.
 Each pigment has a characteristic absorption spectrum, which

depicts the absorption at each .
Photosynthetic Apparatus & Pigments Cont’d…
 If all visible s are absorbed, the object appears black;
 However, if all s are reflected, the object appears white.
Green leaves appear green because these λs are

reflected.
 In higher plants, the major organ of photosynthesis is the leaf, and

The green chloroplasts
within the mesophyll are the actual sites of this process.
Chloroplast
 The chloroplast of higher plants is surrounded by the inner and
outer membranes.
 The region of the chloroplast that is inside the inner membrane and
surrounds the thylakoid membranes is known as the stroma
 Stroma contains the enzymes that catalyze C-fixation and

other biosynthetic pathways.
 Embedded throughout stroma are the pigments containing

thylakoids (the site of photosynthesis) wherein light is used to
oxidize the H2O and form ATP and NADPH.
 Stacks of thylakoids are called grana.

Fig. Structure of chloroplast

 Pigments present in thylakoid membranes are: :
a) Largely two kinds of green molecules, chlorophyll (Chl) a and

b and
b) Yellow to orange coloured pigments classified as carotenoids
They are of two types —
1) Pure hydrocarbon carotenes (orange pigment) and
2) Oxygen containing xanthophylls (yellow pigment).
The carotenes include α-carotene, β-carotene, lycopene,

phytoene, etc.
Xanthophylls include lutein, violaxanthin, zeaxanthin,

 Although both chl a and chl b absorb light, chl a plays a unique
and crucial role in converting light energy to chemical energy.
 All photosynthetic plants, algae, and cyanobacteria contain

chl a,
 Whereas only plants and green algae contain chl b, along with a
few types of cyanobacteria.
 Because of the central role of chl a in photosynthesis, chl a is

called principal pigment.
 While, all pigments used in addition to chl a are known

as accessory pigments - including other chls, as well as other
classes of pigments like the carotenoids.
Photosynthetic pigments absorb the
light that powers photosynthesis
 Chl appears green to our eyes because it absorbs light mainly

in the:
 red (about 660 nm) and
 blue (about 430 nm) parts of the spectrum,
 So only some of the light enriched in green wavelengths

(about 550 nm) is reflected into our eyes.
 Fundamental principle of light absorption states that any
molecule can absorb only one photon of light at any given time
Absorption Spectra of Photosynthetic Pigments

 It depicts the proportion of light (with different wavelength)
absorbed by pigments.
 Chl a and b absorb light of violet, blue, orange and red

wavelength (> 90%) and reflect/ transmit green wavelength.
 Carotenoids (β-carotene and lutein) absorb blue and violet

wavelength and
 transmit/reflect green, yellow, orange and red wavelengths

and hence give orange or yellow colour appearance.
FIGURE: Absorption spectra of some photosynthetic pigments. Curve

1, bacteriochlorophyll a; curve 2, Chl a; curve 3, Chl b; curve 4,
phycoerythrobilin (pigment in red algae); curve 5, β-carotene.
Redox Reactions
 Many energy transformations in cells involve the transfer of e-s or
hydrogen (H+) atoms.
 When a molecule gains an e- or a H+ atom, the molecule is said
to be reduced, and the molecule that gives up the e- is said to be
oxidized.
 A molecule that has been reduced has gained energy; likewise the
oxidized molecule has lost energy.
 Oxidation and reduction reactions are usually coupled
(sometimes called redox reactions); as one molecule is oxidized,
the other is simultaneously reduced.
AH2 + B → A + BH2
(A-reduced) (B-oxidized) (A-oxidized) (B-reduced)
Redox Reactions Cont’d…
 In many oxidation-reduction reactions, an intermediate is used to
transport e-s from one reactant to another.
 One such e- intermediate is NAD, which can exist in both oxidized

and reduced states (NAD+ = oxidized form and NADH = reduced
form).
 Similarly, NADP and FAD also can exist as NADP+/NADPH and

FAD/FADH2, respectively.
 NAD and FAD are common e- carriers in respiration; NADP

serves the same function in photosynthesis.
3.3. Light Reaction and Dark Reaction
 Photosynthesis consists of two major phases,
 the light reactions (light dependent reactions) and
 the dark reactions (light independent reactions) or

Carbon-fixation reactions:
C3 Cycle/Calvin Cycle or the Calvin-Benson cycle and
C4 Cycle/C4 dicarboxylic acid/C4 CO2 fixation/ Hatch and

Slack of CO2 pathway
Light Reaction and Dark Reaction Cont’d…
 The light rxns constitute the photochemical phase of
photosynthesis, during which radiant energy is converted into
chemical energy.
 During the light rxns, H2O molecules are split, releasing O2 and
providing e-s for the reduction of NADP+ to NADPH.
 The light rxs also provide the energy for the synthesis of ATP.
 The reaction happens in the thylakoids of the chloroplasts.
 The dark rxn constitutes the biochemical phase and involves the
fixation and reduction of CO2 to form sugars using the ATP and
NADPH produced in the light reactions.
 The "light-independent" or dark reactions happen in

the stroma of the chloroplasts.
Light Reaction and Dark Reaction Cont’d…
3.3.1. Role of Photosystem I and II in Light Reaction
 Photosystems are arrangements of

chlorophyll and other pigments packed into thylakoids.
 They are functional and structural units of protein complexes

involved in photosynthesis
 that together carry out the primary photochemistry of

photosynthesis:
 the absorption of light, & the transfer of energy & e-s.
 Photosystems are found in the chloroplast thylakoid membranes

of plants, algae and cyanobacteria.
Role of PSI and II in Light Reaction Cont’d…
 There are two kinds of photosystems:
Photosystem II (PSII) and Photosystem I
(PSI), respectively.
 Many Prokaryotes have only one photosystem, PSII.
 Eukaryotes have PSII plus PSI.
PSI uses Chl a, in the form referred to as P700.

PSII uses a form of Chl a known as P680.
 Both "active" forms of Chl a function in photosynthesis due to their
association with proteins in the thylakoid membrane.
 PSI preferentially absorbs far-red light of wavelengths > 680 nm;
 PSII preferentially absorbs red light of 680 nm.
 Another difference between them is that PSI produces a strong
reductant, capable of reducing NADP+, and a weak oxidant.
 PSII produces a very strong oxidant, capable of oxidizing water,
and a weaker reductant.
 These properties of the two PSs are shown schematically in Figure

below.
 The scheme of photosynthesis depicted in Figure, called the Z (for
zigzag) scheme, has become the basis for understanding O2-evolving
(oxygenic) photosynthetic organisms.
FIGURE: Z scheme of photosynthesis. Red light absorbed by PSII produces a

strong oxidant and a weak reductant. Far-red light absorbed by PSI produces a weak
oxidant and a strong reductant. The strong oxidant generated by PSII oxidizes water,
while the strong reductant produced by PSI reduces NADP+. This scheme is basic
to an understanding of photosynthetic electron transport. P680 and P700 refer to
the λs of maximum absorption of the rxn center Chls in PSII and PSI, respectively.
THE LIGHT REACTIONS
 The light rxns are composed of two cooperating PSs, PSI and II, and
take place on the thylakoid membranes within the chloroplasts.
 Each PS is a complex of several hundred Chl and carotenoid

molecules (known as light-harvesting antennae) and associated
membrane proteins.
 When light strikes a pigment molecule in either PS, the energy is

channeled into a RC, which consists of a chl a molecule bound to a
membrane protein.
 The RC for PSI is known as P700, which indicates the λ of maximum

light absorption in the far-red region of the spectrum;
 The RC for PSII is P680, again indicating the peak absorbance in the red
region of the spectrum.
 Associated with the PSs are various enzymes and coenzymes that
function as e- carriers and are components of the thylakoid membranes.
Light Reactions Cont’d…
 All the chemical processes that make up the light rxns of photosynthesis
are carried out by 4 major protein complexes,
 which are vectorially oriented in the thylakoid membrane to
function as follows:
i. Photosystem II:
 Oxidizes water to O2 in the thylakoid lumen and in the
process releases protons (H+) into the lumen.
2H2O + 4Photons + 2PQ + 4H+ → O2 + 4H+ + 2PQH2
2H2O → O2 + 4H + 4e + -
ii. Cytochrome b6 — Cytochrome f complex

Cytochrome is a cpd containing protein and Fe that plays a
role in photosynthesis and respiration
It is an e- transport chain that connects the two PSs.
Receives e-s from PSII and deliver them to PSI;
it also transports additional H+s into the lumen from the stroma
2PQH2 + 4PC (Cu2+) → 2PQ + 4PC (Cu+) + 4H+

iii. Photosystem I:
Receives e-s taken from water by PSII,
Oxidizes reduced PC and transfer e-s to ferredoxin (Fd).
Light + 4PC(Cu+) + 4Fd(Fe3+) → 4PC(Cu2+) + 4Fd(Fe2+)

Reduces NADP + to NADPH in the stroma by the action of
Fd and the flavoprotein ferredoxin—NADP reductase (FNR).
4Fd(Fe2+) + 2NADP+ + 4H+ → 4Fd(Fe3+) + 2NADPH2

iv. ATP Synthase or Coupling Factor:
Produces ATP as protons (H+) diffuse back through it from
the lumen into the stroma.
H+s due to water oxidation move from lumen to stroma

through ATP Synthase,
 creating a gradient in electrochemical potential, which

is driving ATP synthesis.
This synthesis of ATP is known as photophosphorylation,

since the energy that drives the whole process is from
sunlight.
FIGURE: The transfer of e-s and H+s in the thylakoid membrane is carried out vectorially by 4
protein complexes. Water is oxidized and H+s are released in the lumen by PSII. PSI reduces
NADP+ to NADPH in the stroma, via the action of Fd and the FNR. H +s are also transported
into the lumen by the action of the cytochrome b6 f complex and contribute to the
electrochemical H+ gradient. These H+s must then diffuse to the ATP synthase enzyme, where
their diffusion down the electrochemical potential gradient is used to synthesize ATP in the
stroma. Reduced PQH2 and plastocyanin transfer e-s to cytochrome b6 f and to PSI, respectively.
Dashed lines represent e- transfer; solid lines represent H+ movement.
 The overall light rxns proceed with breathtaking speed as a constant flow
of e-s moves from H2O to NADPH, powered by the vast energy of the
sun.
 The ATP and NADPH that result from the light rxs are needed to drive
the biochemical rxs in the dark rxn.
 The overall equation for the light-dependent rxs under the conditions of
non-cyclic e- flow in green plants is:
2H2O + 2NADP+ + 3ADP + 3Pi + light

→
2NADPH + 2H+ + 3ATP + O2
Electron transport
 There are two methods of e- transport:
 Cyclic and
 Non-cyclic e- transport.
1. Non-cyclic electron transport

 It begins with the oxidation of water and
 Completes by reducing NADP to NADPH using flavoprotein

ferredoxin-NADP reductase (FNR).
 Both PSII and PSI are involved.

2. Cyclic electron transport
 Cyclic e- flow generates ATP but no NADPH
 Occurs inside PSI
 Some of the cytochrome b6-f complexes are found in the stroma
region of the membrane, where PSI is located
 Under certain conditions cyclic e- flow from the reducing side of PSI,
through the cytochrome b6-f complex and back to PSI, is known to
occur.
 This cyclic e- flow is coupled to H+ pumping into the lumen, which
can be utilized for ATP synthesis but does not oxidize water or
reduce NADP+
Photophosphorylation
 Phosphorylation: a chemical/biochemical rxn producing an
organic phosphate by adding a phosphate group to an organic
molecule,
 E.g. ATP synthesis in respiration
ADP (adenosine diphosphate) + Pi → ATP (adenosine triphosphate)
 Photophosphorylation: ATP synthesis with the aid of light.
ADP + Pi + energy → ATP

3.4. Path of Carbon in Photosynthesis
 There are three metabolic pathways for
carbon fixation in photosynthesis:
C3,
C4 and
CAM
3.4.1. C3 Photosynthesis Cycle
 C3 photosynthetic cycle is known as:
 C3 - CO2 fixation pathway in plants/

 Photosynthetic Carbon-di-oxide Reduction (PCR) pathway/
 Calvin Cycle or the Calvin-Benson cycle.
 These reactions utilize the ATP and NADPH produced in the light
reactions
 But do not involve the direct participation of light and are hence
sometimes referred to as light-independent reactions, or dark reactions.
 The actual fixation of CO2, which diffuses into the leaf from the atm.,
occurs by a cyclic series of reactions called the Calvin cycle (named
after one of the pioneer researchers in this area).
C3 photosynthesis cycle Cont’d…
Calvin of the University of California,
Berkeley
Melvin Calvin works in his photosynthesis laboratory

 The Cycle occurs in the stroma of chloroplasts (mesophyll cell).
 Plants in which the first detectable product of photosynthesis is a 3-

C cpd About 85% of the plant species on the planet are C3
plants, including rice, wheat, soybeans and all trees.
 C3 - CO2 fixation pathway is the first identified and principal

pathways.
 CO2 fixation in chloroplasts utilizes NADPH produced during non-cyclic

e- transport, whereas ATP produced during photophosphorylation serves
as source of energy facilitating the process of reduction.
 the C of the first stable intermediate, 3-phosphoglycerate, and

 it is further reduced in the glyceraldehyde-3-phosphate product
– so chemically prone to reduction.
 The first stable product formed in a shortest time of 2 seconds is a 3 C
cpd 3-phosphoglyceric acid (3-PGA) and the substrates are CO2 and 5
C sugar cpd (RuBP).
 An enzyme, ribulose-1,5-bisphosphate carboxylase (Rubisco)
irreversibly catalyses the reaction of combining CO2 with RuBP.
5 C cpd
Unstable 6 Cpd
A stable 3 C cpd
 Two molecules of 3-PGA are further phosphorylated by
phosphogycerokinase enzyme using 2 molecules of ATP to yield
two molecules of 1,3-bisPGA.
 Enzyme 3-phosphoglyceraldehyde dehydrogenase further

catalyzes the reaction and
 utilizes 2 molecules of NADPH as reductant to reduce two

molecules of 1,3-bisPGA into two molecules of 3-
phosphoglyceraldehyde (glyceraldehyde-3-Phosphate) (3-PGAld)
 3-PGAld – a sugar and a precursor for other sugars.

 These reactions are given as:
The Calvin Cycle proceeds in 3 stages
1. Carboxylation of RuBP (CO2 acceptor), forming two molecules
of 3-PGA, the first stable intermediate of the Calvin Cycle.
2. Reduction of 3-PGA, forming glyceraldehyde-3-phosphate (3-

PGAld), a carbohydrate (sugar).
3. Regeneration of the CO2 acceptor ribulose-1,5-bisphosphate

(RuBP) from 3-PGAld.
 Summary of Calvin Cycle:
3.4.2. C4 Photosynthesis Cycle
 This photosynthetic pathway is known as:
 C4 dicarboxylic acid/
 C4 CO2 fixation/
 Hatch and Slack of CO2 pathway.
 The naming 'Hatch-Slack pathway is in honor of

Marshall Davidson Hatch and Roger Slack, who elucidated it in Australia
in 1966.
 The C4 pathway is used in about 3% of all vascular plants and occurs in

several thousand species of tropical and subtropical plants,
 including the economically important crops of corn,
Maize (or corn) and Sugarcane, a common C4 plants

 There are differences in leaf anatomy between plants that have a
C4 C-cycle (called C4 plants) and those that photosynthesize solely
via the Calvin photosynthetic cycle (C3 plants).
 A cross-section of a typical C3 leaf reveals one major cell type

that has chloroplasts, the mesophyll cell.
 In contrast, a typical C4 leaf has two distinct chloroplast-

containing cell types:
 mesophyll (MS) and

 bundle sheath (BS) cells.
 There is considerable anatomic variation in the arrangement of the

BS cells with respect to the MS and vascular tissue.
 The C4 plants often possess a characteristic leaf anatomy

called Kranz anatomy, from the German word for “wreath/ring”.
 Their vascular bundles are surrounded by two rings of cells; the

inner ring, called BS cells,
 BS cells contain starch-rich chloroplasts lacking grana, which differ

from those in MS cells present as the outer ring.
Mesophyll cells forming the outer ring
Bundle sheath cells (surrounding vascular

bundles) forming the inner ring
 In all cases, however, operation of the C4 cycle requires the cooperative

effort of both cell types (MS and BS).
 In addition, an extensive network of plasmodesmata connects MS and

BS cells,
 Thus providing a pathway for the flow of metabolites between the

cell types.
 4 C compounds (aspartic and malic acids) are produced as first stable
product.
 Carboxylation reaction in C4 plants is catalyzed by the cytosolic

enzyme phoshpo-enol-pyruvate carboxylase (PEPCase).
 In C4 plants, there is a division of labour between two types of

photosynthetic cells - MS cells and BS cells -
 the former are involved in initial carboxylation reaction and
 the later in decarboxylation and refixation of CO2.
 All PEPCase is present in MS cells and Rubisco in BS cells enabling C4

plants to use both kinds of CO2 fixing mechanism.
 The basic C4 cycle consists of four stages:
1. Fixation of CO2 by the carboxylation of phosphoenolpyruvate

(PEP) in the MS cells to form a 4-C acid (malate and/or aspartate).
2. Transport of the 4-C acids to the BS cells.
3. Decarboxylation of the 4-C acids within the BS cells and generation

of CO2, which is then reduced to carbohydrate via the Calvin cycle.
4. Transport of the 3-C acid (pyruvate or alanine) that is formed by

the decarboxylation step back to the MS cell and regeneration of the
CO2 acceptor, PEP.
FIGURE: The basic C4

photosynthetic carbon cycle
involves 4 stages in two d/t
cell types: (1) Fixation of CO2
into a 4-C acid in a MS; (2)
Transport of the 4-C acid from
the MS to a BS cell; (3)
Decarboxylation of the 4-C
acid, and the generation of a
high CO2 concentration in the
BS cell. The CO2 released is
fixed by Rubisco and
converted to carbohydrate by
the Calvin cycle. (4) Transport
of the residual 3-C acid back
to the MS cell, where the
original CO2 acceptor, PEP,
is regenerated.
3.4.3. CAM Photosynthesis Cycle
 This cycle is also known as:
 CO2 Fixation in Succulent Plant Species/ Crassulacean Acid
Metabolism (CAM).
 First discovered in succulent plants of the Crassulaceae family,
therefore, called Crassulacean Acid Metabolism (CAM) pathway.
 CAM plants (plants in Crassulaceae family) are adapted to desert/arid

ecosystem with peculiar morphology, anatomy and physiology to suit
this ecosystem.
 Initial carboxylation takes place in the dark,
 Followed by decarboxylation and refixation of CO2 in light to

economize the water loss through transpiration.
 E.g. Pine apple, vanilla, cacti, etc..
CAM photosynthesis cycle Cont’d…
Cacti
(CAM plant)
in desert
 The CAM mechanism is similar in many respects to the C4 cycle.
 In C4 plants, formation of the 4-C acids in the MS is spatially

separated from decarboxylation of the 4-C acids and from refixation of
the resulting CO2 by the Calvin Cycle in the BS.
 In CAM plants, formation of the C4 acids and its decarboxylation is

both temporally and spatially separated.
 At night, CO2 is captured by PEPCase in the cytosol, and the

malate that forms from the oxaloacetate product is stored in the
vacuole.
 During the day, the stored malate is transported to the chloroplast
and decarboxylated by NADP-malic enzyme,
 The released CO2 is fixed by the Calvin Cycle.

 The adaptive advantage of CAM is:
 the reduction of water loss by transpiration (increased efficiency in

the use of water),
 achieved by the stomatal closing during the day, but open at night to
collect CO2.
 Being able to keep stomata closed during the hottest and driest part of
the day
 reduces the loss of water through evapotranspiration,
 allowing such plants to grow in environments that would otherwise be

far too dry.
 Plants using only C3 carbon fixation, for example, lose 97% of the water
they uptake through the roots to transpiration
 a high cost avoided by plants able to employ CAM.
 Typically, a CAM plant loses 50-100 g of water for every gram of CO 2

gained,
 compared with values of 250-300 g and
 400-500 g for C4 and C3 plants, respectively.
 Thus, CAM plants have a competitive advantage in dry environments.

Why Rubisco and not PEP Carboxylase fixes CO2 in day light?
 Both Rubisco and PEP Carboxylase are present in CAM plants and both
have about equal affinities for dissolved CO2.
 Furthermore, cytosolic PEPCase should encounter the incoming CO2 first

before CO2 reaches to chloroplasic Rubisco.
 However, it is Rubisco that fixes CO2 in light and not the PEPCase
because:
a. In day light PEPCase in CAM plants is converted into inactive form

that has extremely low affinity to CO2,
b. PEPCase is strongly inhibited by malic acid released from vacuoles in

day light,
c. Changes in the activities of still other enzymes that favour CO2 fixation by
Differences among plant species with 3 types of photosynthetic pathways.
“What drives life is
thus a little electric
current, set up by
the sunshine.”,
American biochemist and Nobel laureate
ALBERT SZENT-GYORGYI
3.6. Photorespiration
 As CO2 []s rise, the rate at which sugars are made by the light-independent
reactions increases until limited by other factors.
 Rubisco, the enzyme that captures CO2 in the light-independent reactions, has a
binding affinity for both CO2 and O2.
 When the [] of CO2 is high, Rubisco will fix CO2.
 However, if the CO2 [] is low, Rubisco will bind O2 instead of CO2.
 This process, called photorespiration, uses energy, but does not produce sugars.
 It is known as the “Wasteful” Oxygenase Reaction of Rubisco/ C2 Oxidative

Photosynthetic Carbon Cycle.
 An important property of Rubisco is its ability to catalyze both the carboxylation

and the oxygenation of RuBP.
 Oxygenation is the primary reaction in a process of photorespiration.

Photorespiration Cont’d…
 Because photosynthesis and photorespiration work in diametrically
opposite directions, photorespiration results in loss of CO2 from cells
that are simultaneously fixing CO2 by the Calvin Cycle.
 The reactions of photorespiration occur in three different compartments

in higher plants:
 the chloroplast,
 the peroxisome and
 the mitochondrion.
 The reactions begin in the chloroplast with the oxygenation of RuBP.
 A highly simplified summary is:
2 Glycolate + 2 RUBP + ATP → 3-PGA + CO 2 +

ADP + NH
 Photorespiration is common in C3 plants, but absent/negligible in C4

plants.
 The affinity of Rubisco for CO2 is much greater than for O2,
 but O2 fixation in C3 plants occurs because cellular concentration of

O2 is much greater than that of CO2.
 Photorespiration is light dependent/mediated
 Photorespiration is active at high temperature, because high

temperature lowers the CO2:O2 ratio.
RuBisCO oxygenase activity is disadvantageous to plants for
several reasons:
1. One product of oxygenase activity is phosphoglycolate (2
carbon) instead of 3-phosphoglycerate (3 carbon).
 Phosphoglycolate cannot be metabolized by the Calvin-Benson
Cycle and represents carbon lost from the cycle.
 A high oxygenase activity, therefore, drains the sugars that
are required to recycle RuBP and for the continuation of
the Calvin-Benson Cycle.
2. Phosphoglycolate is quickly metabolized to glycolate that is
toxic to a plant at a high [];
 it inhibits photosynthesis.
3. Salvaging glycolate is an energetically expensive process that

uses the glycolate pathway, and
 only 75% of the carbon is returned to the Calvin-Benson

Cycle as 3-PGA.
 The reactions also produce ammonia (NH3), which is able to

diffuse out of the plant, leading to a loss of nitrogen.
Importance of Photorespiration
 Under conditions of high light intensity and low intercellular CO 2 []
(e.g., when stomata are closed because of water stress), it dissipates
excess ATP and reducing power from the light reactions,
 Thus preventing damage to the photosynthetic apparatus (from 2

RuBP three 3-PGA enter the Calvin Cycle). .
 There is evidence from work with transgenic plants that

photorespiration protects C3 plants from photooxidation and
photoinhibition.
C-fixation efficiency in C3 and C4
 In the absence of photorespiration, the Calvin cycle produces six 3-
PGAld for three RuBP molecules.
 In the presence of photorespiration, from the same amount of RuBP
molecules, only 4.5 molecules of 3-PGAld are produced.
 C4 plants capture CO2 by PEP, which is mediated by the enzyme

PEPCase.
 PEPCase has a stronger affinity for CO2 than does RuBP.
 When CO2 levels decline below the threshold for Rubisco, RuBP is
catalyzed with O2 instead of CO2.
 C4 plants adjust to decreased levels of CO2 by artificially raising the

CO2 [] in certain cells (BS, where C3 cycle occurs in C4) to prevent
photorespiration.
3.7. Factors Affecting Photosynthesis
 Many external and internal factors influencing the rate of photosynthesis
are the following.
A. External factors B. Internal or Plant factors (Leaf

(Environmental factors): or Genetic factors):
1. CO2 []s 1. Age of the leaf
2. Light 2. Chlorophyll
3. Temperature 3. Hormones (cytokinin and
4. Oxygen gibberllin)
5. Soil water 4. Leaf anatomy
6. Air pollutants 5. Accumulation of end
7. Minerals (Nutrient supply) products/carbohydrates

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CHAPTER 3:

 reduction of CO2 to form organic cpds such as carbohydrates.

Carbon Water Carbohydrate Oxygen

 In actual practice light is a mixture of different colors and the EMS

 Light has both wave and particle nature.

 The particle nature of light is usually expressed in terms of quanta or

 i.e. discrete packets of energy each having a specific associate

 Visible light is one small part of the EMS.

 Solar radiation in the visible range of spectrum from 400 nm (violet) -

Figure: The electromagnetic spectrum (EMS).

 be transmitted (pass through the object),

 Pigments (light-absorbing molecules) absorb light selectively,

 Each pigment has a characteristic absorption spectrum, which

 However, if all s are reflected, the object appears white.

Green leaves appear green because these λs are

 In higher plants, the major organ of photosynthesis is the leaf, and

 Stroma contains the enzymes that catalyze C-fixation and

 Embedded throughout stroma are the pigments containing

 Stacks of thylakoids are called grana.

Fig. Structure of chloroplast

a) Largely two kinds of green molecules, chlorophyll (Chl) a and

b) Yellow to orange coloured pigments classified as carotenoids

They are of two types —

1) Pure hydrocarbon carotenes (orange pigment) and

2) Oxygen containing xanthophylls (yellow pigment).

The carotenes include α-carotene, β-carotene, lycopene,

Xanthophylls include lutein, violaxanthin, zeaxanthin,

 All photosynthetic plants, algae, and cyanobacteria contain

 Because of the central role of chl a in photosynthesis, chl a is

 While, all pigments used in addition to chl a are known

 Chl appears green to our eyes because it absorbs light mainly

 red (about 660 nm) and

 blue (about 430 nm) parts of the spectrum,

 So only some of the light enriched in green wavelengths

Absorption Spectra of Photosynthetic Pigments

 Chl a and b absorb light of violet, blue, orange and red

 Carotenoids (β-carotene and lutein) absorb blue and violet

 transmit/reflect green, yellow, orange and red wavelengths

FIGURE: Absorption spectra of some photosynthetic pigments. Curve

 One such e- intermediate is NAD, which can exist in both oxidized

 Similarly, NADP and FAD also can exist as NADP+/NADPH and

 NAD and FAD are common e- carriers in respiration; NADP

 the light reactions (light dependent reactions) and

 the dark reactions (light independent reactions) or

C3 Cycle/Calvin Cycle or the Calvin-Benson cycle and

C4 Cycle/C4 dicarboxylic acid/C4 CO2 fixation/ Hatch and

 The reaction happens in the thylakoids of the chloroplasts.

 The "light-independent" or dark reactions happen in

 Photosystems are arrangements of

 They are functional and structural units of protein complexes

 that together carry out the primary photochemistry of

 the absorption of light, & the transfer of energy & e-s.

 Photosystems are found in the chloroplast thylakoid membranes

PSI uses Chl a, in the form referred to as P700.

 These properties of the two PSs are shown schematically in Figure

FIGURE: Z scheme of photosynthesis. Red light absorbed by PSII produces a

 Each PS is a complex of several hundred Chl and carotenoid

 When light strikes a pigment molecule in either PS, the energy is

 The RC for PSI is known as P700, which indicates the λ of maximum