Chapter 3

PHOTOSYNTHESIS

In Class XI, you have read about the structure Joseph Priestley (1733-1804) showed that
plants have the ability to take up CO2 from the
of chloroplast. In this chapter, you will learn
atmosphere and release O2. Later, in 1780, the
about the function of chloroplasts in detail, that
Dutch physician Jan Ingenhousz (1730-1799)
is, photosynthesis. The chapter covers, in
confirmed Priestley’s work. He further
addition to a brief historical perspective,
discovered that release of O2 by plants was
description of photochemical and biosynthetic
possible only in sunlight and only by the green
phases, photorespiration, C4 pathway, factors
parts of the plants. Shortly thereafter, a Swiss
affecting photosynthesis, translocation of
scholar, Theodore de Saussure, found that
photosynthates, and significance of
water is an essential requirement for
photosynthesis.
photosynthesis to occur.
Photosynthesis, as you know, is a
Thus, it became clear that green plants,
process by which green plants synthesise
apart from taking nutrients from soil, make their
their own food in the presence of light. Plants own food by utilising CO2, water and sunlight.
have the ability of capturing energy coming This became the basis of photosynthesis.
from the sun in the form of light, and During this process, O2 is evolved and released
converting it to chemical energy. This to the atmosphere. A simplified equation of
chemical energy is used for the growth and photosynthesis was given by C.B. van Niel of
sustenance of our biosphere. Photosynthesis Stanford University, USA :
by plants requires carbon dioxide from the
atmosphere and uses light energy from the Light energy
CO2 + 2H2O  → (CH O) + H O + O
sun for converting CO 2 to complex foods, Chlorophyll 2 n 2 2

which are consumed by man and animals.

It also provides energy in the form of fossil In photosynthesis, carbon dioxide is fixed
fuels derived from prehistoric photosynthetic (or chemically reduced) to carbohydrate
(CH2O)n, where n is an integer. In glucose, n is
products. Thus, photosynthesis is the single
6, and thus, the chemical formula of glucose
most important biological process for the
is C 6H12O6. Water molecule is split in the
biosphere.
presence of light (called photolysis) to release
3.1 HISTORICAL PERSPECTIVE O2 . Note that O2 released in the above equation
is from H2O and not from CO2.
Study on photosynthesis originated only about
300 years ago. Before this, Aristotle and other 3.2 SITE FOR PHOTOSYNTHESIS
Greeks thought that plants get their food only
from the soil. Later, the Belgian physician, Jan Photosynthesis takes place only in the green
Baptista van Helmont, did a simple experiment parts of the plant, mostly in leaves, and to a
and concluded that all the substance of the lesser extent in green stems or floral parts.
plant was produced from water and none from Within a leaf, photosynthesis occurs
the soil. By the end of the eighteenth century, particularly in specialised cells, called
 PHOTOSYNTHESIS
23

mesophyll cells. These cells contain the of magnesium. A side chain, called the phytol
chloroplasts, which are the actual sites for chain, extends from one of the pyrrole rings
photosynthesis in green plants. Chloroplasts in the chlorophyll molecule (Fig. 3.2). This
are located at the outer margins with their long side chain composed of hydro-carbons
broad surfaces parallel to the cell wall of the helps to anchor the chlorophyll molecules
mesophyll cells. This helps in easy diffusion of within the thylakoid membranes in the
CO2 required for photosynthesis from the chloroplasts. There are different kinds of
atmosphere to the inside of chloroplasts. chlorophyll molecules present in
The thylakoids in the chloroplasts contain photosynthetic organisms. In higher plants,
most of the machinery for the photochemical mostly there are two kinds : chlorophyll a and
reactions of photosynthesis. They contain chlorophyll b. Both chlorophyll a and b are very
pigments required for capturing solar energy similar in their molecular structure, except that
to initiate photosynthesis. The major form of the –CH3 group present in chlorophyll a and
pigment present in thylakoids is chlorophyll. –CHO group in chlorophyll b.
A pigment is a substance that absorbs light of Chlorophyll a is the major pigment involved
in trapping and converting light energy
into chemical energy (Fig. 3.3). Chlorophyll b
Relative light absorption

40 acts as an accessory pigment and helps

absorb light form a broader spectrum during
30
photosynthesis. Chlorophyll b constitutes
20 about one-fourth of the total chlorophyll
content and absorbs light of different
10
wavelength than the chlorophyll a. On
0 0 absorbing light, the chlorophyll b molecule is
10 400 450
500 550 600 650 700 750 105
Wavelength, nm
excited and transfers its energy to the
X-rays Ultra- Violet Blue Green Yellow Orange Red Infrared Radio chlorophyll a molecule. Thus, chlorophyll a
violet waves
Visible spectrum molecules act as reaction centres.
Pigments other than chlorophyll are also
involved in the process of photosynthesis. These
pigments, called carotenoids, are of red,
Fig. 3.1 Absorption spectrum of chlorophyll orange and yellow colours. Like chlorophyll
showing maximum absorption in the
blue and red regions of the visible
they are also embedded in the thylakoid
specturm of light membranes of chloroplasts. These accessory
pigments act as antenna complexes and
different wavelengths. Chlorophyll, however, is harvest light from different wavelengths of the
a pigment that gives green colour to the leaves. spectrum other than that of the chlorophyll.
It absorbs light in the violet and blue The light captured by these pigments
wavelengths, and also in the red region of the is funnelled into the reaction centres
visible spectrum of light. This portion of the (chlorophyll a) for conversion into the
spectrum between 400nm and 700nm is also electrical energy (Fig. 3.4). The accessory
referred to as photosynthetically active pigments and the reaction centre, together form
radiation (PAR) (Fig. 3.1). But, the chlorophylls photosystem.
reflect the green light, and hence, they impart 3.3 THE PHOTOCHEMICAL AND
green colour to the leaves. BIOSYNTHETIC PHASES
Chlorophyll is the principal pigment
involved in photosynthesis. It is a large Photochemical Phase (Light reactions)
molecule composed of four 5-membered By now, we have learnt that different pigments
rings, called pyrrole rings, and a central core – chlorophyll a, chlorophyll b – and carotenoids
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Molecular Model of Chlorophyll

Fig. 3.2 The chemical structure of chlorophyll. X*: – CH3 in Chlorophyll a, – CHO in Chlorophyll b
 PHOTOSYNTHESIS
25

Oxygen-using bacteria
Relative Absorption

Filament of
Green Alga

Violet Blue Green Yellow Orange Red

400 450 550 600 650 700

Wavelength (nm)

Fig. 3.3 Action spectrum of photosynthesis determined by T.W. Englemann in 1882 using green
alga. The scientist measured rate of photosynthesis as the amount of O2 released, which he
detected by using bacteria that are attracted by O2. Note that the above action spectrum
parallels the absorption spectrum of chlorophyll a, meaning thereby that photosynthesis
depends on the light absorbed by chlorophyll

Light Light

hv hv

Antenna
(Accessory
pigments)

Photosystem

Reaction
A Q centre
e–
Electron (Chlorophyll a)
Electron +
A – acceptor
donor Q

Fig. 3.4 Conversion of light into electrical energy. Accessory pigment molecules absorb light and
funnel it to the reaction centre for conversion to electrical energy
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participate in photosynthesis. Chlorophyll a is acceptor molecule before) is able to restore its

present in different forms, which have electrons from the water molecule. After
maximum absorption at different wavelengths accepting electrons from the excited P680, the
of visible light. For instance, one of the forms primary electron acceptor (pheophytin in
shows an absorption peak at a wavelength of plants) is reduced. The reduced acceptor (a
670 nanometers (nm) and is designated as Chl strong reducing agent) now donates its
a 670. Similarly, two other forms of electrons to the downstream components of the
chlorophyll a are Chl a 680 (P680) and Chl a electron transport chain.
700 (P700), with peak absorption at 680 and Similar to the photosystem II (P 680 ),
700 nm, respectively. These pigments are photosystem I (P700) is excited on absorbing light
anchored in thylakoids in discrete units of and gets oxidised. It transfers its electrons to the
organisation called photosystems (Fig. 3.4). primary electron acceptor, which, in turn, gets
About 250 to 400 pigment molecules reduced. While the oxidised P700 draws electrons
constitute a single photosystem. Two different from photosystem II, the reduced electron acceptor
photosystems exist with different forms of of photosystem I, transfers electrons to ferredoxin
chlorophyll a as the reaction centre. In and ferredoxin-NADP reductase to reduce NADP+
photosystem I (PS I), chlorophyll a with to NADP. The NADPH+, a powerful reducing agent,
maximum absorption at 700 nm (P700), and in is then utilised in the reduction of CO2 to
photosystem II (PS II), chlorophyll a with peak carbohydrates in the carbon reaction of
absorption at 680 nm (P680), acts as reaction photosynthesis. The reduction of CO 2 to
centres. Here, P stands for pigments. The PS II carbohydrates also requires energy in the form of
is located in the appressed regions of grana ATP, produced during light reactions. This process
thylakoids, and the PS I in the stroma of ATP formation from ADP in the presence of light
thylakoids and non-appressed regions of in chloroplasts is called photophosphorylation.
grana. The primary function of the two
photosystems which interact with each other, Photophosphorylation
is to trap light energy and convert it to the Photophosphorylation occurs in chloroplasts
chemical energy (ATP). This chemical energy in two ways :
stored in the form of ATP is used by living cells. (i) Non-cyclic, and
Electron Transport Chain (ii) Cyclic.
Non-cyclic photophosphorylation : Non-
The light-driven reactions of photosynthesis,
cyclic photophosphorylation is a result of an
referred to as electron transport chain, were
interaction of photosystem I and photosystem
first formulated by Robert Hill in 1939. The II. As discussed above, there is a continuous
electron transport chain of photosynthesis is flow of electrons from water to photosystem II
initiated by the absorbance of light by the to photosystem I, and finally, to NADP. As the
photosystem II (P 680) (Fig. 3.5). When P680 electrons pass downhill in an electron transport
absorbs light, it is excited and its electrons are chain, ATP is formed from ADP (Fig. 3.5).
transferred to an electron acceptor molecule. Because the electron flow from water to NADP
By doing so, P680 becomes a strong oxidising is unidirectional, the process of ATP formation
agent and splits a molecule of water to release is termed as non-cyclic photophosphorylation.
oxygen. This light-dependent splitting of the Cyclic photophosphorylation : When
water molecule is called photolysis. non-cyclic photophosphorylation is stopped
Manganese, calcium and chloride ions play under certain conditions, cyclic
prominent roles in the photolysis of water. With photophosphorylation occurs. The non-cyclic
the breakdown of water, electrons are photophosphorylation can be stopped by
generated, which are then passed to the illuminating isolated chloroplasts with light of
oxidised P680. Thus, the electron-deficient P680 wavelengths greater than 680nm. By this way,
(because it had transferred its electrons to an only photosystem I is activated, as it has a
 PHOTOSYNTHESIS
27

Fig. 3.5 Electron transport flow and non-cyclic photophosphorylation : Light energy absorbed by
the reaction centre, P680 of Photosystem II is used in taking electrons to a higher energy
level, which are then replaced by electrons drawn from water molecules. The electrons
enter electron transport chain and are passed to a lower energy level, that is, the reaction
centre, P700 of Photosystem I. During this process, energy is released and stored in the form
of ATP. From Photosystem I, the electrons are passed to another primary electron acceptor
and via other electron carriers to NADP + to form NADPH. ATP and NADPH+H+ thus formed
are used in carbon reactions to fix CO2 to (CH2O)n
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maximum absorption at 700nm, and photophosphorylation (Fig. 3.6). It may be

photosystem II, which absorbs at 680nm, noted that in cyclic photophosphorylation,
remains inactivated. Due to the inactivation of oxygen is not released (as there is no photolysis
photosystem II, the electron flow from water to of water) and NADPH is also not produced.
NADP+ is stopped, and also CO2 fixation is Biosynthetic Phase (Dark reactions)
retarded. When CO 2 fixation is stopped, We have learnt in the previous section how ATP
electrons will not be removed from the reduced and NADPH are formed during the
NADPH. This means that NADPH will not be photochemical reactions in chloroplasts. Both
oxidised and NADP+ will no longer be available ATP and NADPH2 are essential requirements
as an electron acceptor. Under these for the assimilation of CO2 to carbohydrates.
circumstances, cyclic photophosphorylation The reactions catalysing the assimilation of CO2
occurs. During cyclic photophosphorylation, to carbohydrates take place in the stroma,
electrons from photosystem I are not passed where all the necessary enzymes are localised.
to NADP from the electron acceptor (as NADP These reactions are referred to as dark
is no longer available in oxidised state to receive reactions, leading to the photosynthetic
the electrons). Instead, the electrons are reduction of carbon to carbohydrates.
transferred back to P 700 . This downhill In the first phase of carbon reactions,
movement of electrons from an electron CO 2 is accepted by a 5-carbon molecule,
acceptor to P700 results in the formation of ATP ribulose-1, 5-bisphosphate (RuBP) and two
from ADP, and is ter med as cyclic molecules of 3-carbon compound, i.e.,
3-phosphoglycerate (PGA) are formed (Fig. 3.7).
This 3-carbon molecule is the first stable
Electron product of this pathway, and hence it is called
-1.0
acceptor C3 pathway . Such plants which fix Co2 using
the C3 pathway are called C3 plants. The
formation of PGA is called as carboxylation.
This reaction is catalysed by an enzyme called
-0.8
ribulose bisphosphate carboxylase
in

oxygenase (Rubisco). In addition to the

Transport Cha

carboxylase activity, this enzyme also possesses

-0.6 oxygenase activity, and hence abbreviated as
2e –

Rubisco (ribulose bisphosphate carboxylase

Redox potential (ev)

2e –
cyt b6 oxygenase). The oxygenase activity of the
ADP + P i
Electron

-0.4 enzyme allows O2 to compete with CO2 for

combining with RuBP (photorespiration).
After the carboxylation reaction, reduction
Reaction centre
0.2 f (P700) of PGA occurs, where ATP and NADPH2 formed
ATP yt
c during the photochemical reactions are
utilised. With the reduction of PGA,
0 Antenna glyceraldehyde-3-phosphate is formed. These
molecules 3-carbon molecules, also called triose
phosphates, are diverted from the Calvin cycle
0.2 Photosystem I and act as precursors for the synthesis of
sucrose and starch. To complete the cycle, and
for the cycle to continue on its own,
Light
regeneration of the initial 5-carbon
acceptor molecule, that is RuBP, takes place.
The regeneration of RuBP from glyceraldehyde-
Fig. 3.6 Cyclic photophosphorylation 3-phosphate requires another ATP molecule
 PHOTOSYNTHESIS
29

called oxygenation (Fig. 3.8). Respiration that

Atmosphere is initiated in chloroplasts under high light
conditions, is called photorespiration. This
C02 + H2o
occurs essentially because of the fact that the
Ribulose-1,5-
bisphosphate active site of enzyme Rubisco is the same for
both carboxylation and oxygenation. The
Carboxylation
oxygenation of RuBP in the presence of O2 is
ADP
the first reaction of photorespiration, which
leads to the formation of one molecule of
phosphoglycolate, a two-carbon compound
Regeneration and one molecule of PGA. While the PGA is
3-phosphoglycerate used up in the Calvin cycle, the
phosphoglycolate is dephosphorylated to form
ATP glycolate in the chloroplast (Fig. 3.9). From the
+
ATP NADPH chloroplast, the glycolate is diffused to
peroxisome, where it is oxidised to glyoxylate.
Reduction
In the peroxisome, the glyoxylate is used to
Triose form the amino acid, glycine. Glycine enters
phosphate
ATP
+
Pi NADP+

Sucrose, starch
RuBP

CO 2
Fig. 3.7 The Calvin cycle proceeds in three O2
stages : (1) carboxylation, during which
CO 2 combines with ribulose-1,5- O2
bisphosphate; (2) reduction, during Rubisco
Carboxylation Oxygenation
which carbohydrate is formed at the (Calvin cycle)
expense of the photochemically made
ATP and NADPH; and (3) regeneration
during which the CO2 acceptor ribulose-
1,5-bisphosphate is formed again so
3-Phosphoglycerate
that the cycle continues 2-Phosphoglycolate

Photorespiration
formed as a result of photophosphorylation
during the light reactions.
The C3 type of carbon reactions occur in CO 2
the stroma of chloroplast. This C3 pathway is
also called Calvin cycle after its discoverer,
Melvin Calvin, who received Nobel Prize for Fig. 3.8 Photorespiration involves oxygenation
discovering this pathway. of ribulose-bisphosphate catalysed
by the enzyme Rubisco. The reactions
3.4 PHOTORESPIRATION lead to the formation of
2-phosphoglycolate, oxidation of
We know by now that the enzyme Rubisco
which results in release of CO2. In
catalyses the carboxylation reaction, where CO2 contrast, carboxylation, also
combines with RuBP for the Calvin cycle to catalysed by Rubisco, results in
initiate. But, this enzyme also has the ability fixation of CO2 for the production of
to catalyse the combination of O2 with RuBP, carbohydrate
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photorespiration has a useful role in protecting

the plants from photooxidative damage. This
means that if enough CO2 is not available to
utilise light energy for carboxylation to
proceed, the excess energy causes damage to
plants. However, photorespiration, that is
oxygenation of RuBP, utilises part of the light
energy and saves the plant from the
photooxidative damage.
The relative levels of O2 and CO2 determine
the occurrence of photorespiration, since both
gases compete for the same active site of
Rubisco. Increased O 2 level increases
photorespiration, whereas increased CO2 level
decreases photorespiration (and increases C3
photosynthesis).
3.5 C4 PATHWAY
Photorespiration, which primarily occurs in C3
plants, leads, as discussed above, to a 25 per
cent loss of the fixed CO2. It takes place in C3
plants because of the fact that the enzyme
Rubisco catalyses both carboxylation and
oxygenation of the initial acceptor molecule,
that is RuBP. As we know now, photosynthesis
has two types of reactions : light reactions and
Fig. 3.9 The photorespiratory pathway carbon reactions. In the light reactions, ATP
and NADPH2 are produced, and O2 is released
mitochondria where two glycine molecules as a by-product, as a result of photolysis of
(4 carbons) give rise to one molecule of serine H 2 O. During carbon reactions, CO 2 is
(3 carbons) and one CO2 (one carbon). The assimilated to produce carbohydrates. Since
serine is taken up by the peroxisome, and both light reactions and carbon reactions occur
through a series of reactions, is converted to in mesophyll cells in C3 plants, it becomes
glycerate. The glycerate leaves the peroxisome inevitable for the enzyme Rubisco to catalyse
and enters the chloroplast, where it is simultaneously, both oxygenation and
phosphorylated to form PGA. The PGA carboxylation of RuBP.
molecule enters the Calvin cycle to make In certain category of plants called C4 plants
carbohydrates, but one CO2 molecule released such as maize, sugarcane, pearl millet,
in mitochondria during photorespiration has amaranth, etc., nature has evolved a wonderful
to be re-fixed. In other words, 75 per cent of mechanism to avoid the occurrence of
the carbon lost by the oxygenation of RuBP is photorespiration, which is considered to be a
recovered, and 25 per cent is lost as release of wasteful process. In these, plants the first stable
one molecule of CO2. compound formed after carboxylation is a
Photorespiration is also called 4-carbon compound. This mechanism requires
photosynthetic carbon oxidation cycle, the presence of two types of photosynthetic
and, as discussed above, involves an cells, that is, mesophyll cells and bundle-
interaction of three organelles – chloroplast, sheath cells (Fig. 3.10). The bundle-sheath cells
peroxisome and mitochondria. Under are arranged in a wreath-like manner. This
conditions of high light and limited CO2 supply, kind of arrangement is called Kranz anatomy
 PHOTOSYNTHESIS
31

Bundle-sheath cells

Mesophyll cells

Fig. 3.10 T ransverse section of maize leaf

showing the arrangement of mesophyll
and bundle-sheath cells. The C 4
pathway takes place in the mesophyll
cells, and the C3 pathway (Calvin cycle)
operates in the bundle-sheath cells.
Both types of cells contain chloroplasts.
Note that maize is a C4 plant

(Kranz : wreath). C3 plants lack bundle sheath.

The C4 plants contain dimorphic chloroplasts,
that means, chloroplasts in mesophyll cells are
granal, whereas in bundle-sheath cells they are
agranal. The granal chloroplasts contain
thylakoids that are stacked to form grana (as
in C3 plants), but in agranal chloroplasts, grana
Fig. 3.11 The C4 photosynthetic carbon cycle
are absent and the thylakoids are present only involves two cell types and proceeds
as stroma lamelle. The presence of two types in four stages : (a) fixation of CO2 into
of cells allows the occurrence of light reactions a 4-carbon acid in the mesophyll
and carbon reactions separately in each type. cells; (b) transport of the 4-carbon
As a result, release of O2 takes place in the acid from the mesophyll cells to the
bundle-sheath cells; (c) decarboxylation
mesophyll, while the CO2 fixation catalysed by
of the 4-carbon acid, generating a
Rubisco occurs in the bundle sheath. In C4 high concentration of CO 2 in the
plants, light reactions occur in mesophyll cells, bundle-sheath cells; and cells, and
whereas the CO2 assimilation is carried out in regeneration of the CO 2 acceptor,
bundle-sheath cells. This type of cellular phosphoenol pyruvate, to continue
the cycle (d) transport of the residual
arrangement does not allow O2 released in
3-carbon acid back to the mesophyll
mesophyll cells to escape to bundle-sheath
cells. Thus, Rubisco, which is present only in CO 2 concentrating mechanism. This CO 2
bundle-sheath cells, does not come into contact concentrating mechanism is called C4 pathway.
with O2, and as a result, oxygenation of RuBP Operation of the C4 pathway, however, requires
is completely avoided. To further reduce the the cooperation of both cell types, that is,
occurrence of photorespiration (oxygenation of mesophyll and bundle-sheath cells. The
RuBP), C4 plants have been endowed with a objective of this pathway is to build up high
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concentration of CO2 in the vicinity of Rubisco transported back to the mesophyll cells, where
in the bundle-sheath cells. High concentration it is used in regenerating phosphoenol pyruvate
of CO2 near Rubisco favours carboxylation and for the pathway to continue on its own.
suppresses photorespiration. The C4 photosynthetic pathway is more
In C4 pathway, CO2 from the atmosphere efficient than the C3 pathway due to the absence
enters through open stomata into the mesophyll of photorespiration in C 4 plants. Major
cells, where it combines with phosphoenol differences in the C3 and C4 pathways are
pyruvate (3-carbon compound) in a reaction summarised in Table 3.1.
catalysed by the enzyme phosphoenol pyruvate 3.6 CRASSULACEAN ACID METABOLISM
carboxylase, (PEPCase) to form a C4 acid,
oxaloacetic acid (OAA) as shown in Figure 3.11. Crassulacean acid metabolism (CAM) refers to
a mechanism of photosynthesis that is different
This reaction occurs in cytosol of the mesophyll
than already discussed C3 and C4 pathways.
cells and is called fixation of CO 2 or
This occurs only in succulents and other plants
carboxylation, to give rise to the first stable
that normally grow in dry conditions. In CAM
product of the pathway, which is a C 4
plants, CO2 is taken up by the leaves on green
compound, and hence the name C4 pathway. stems through stomata, which remain open in
The OAA is then converted into malate or the night. However, during the day, the stomata
aspartale (both C4 acids) and then transported. remain closed in these plants to conserve
The next step is the transport of the OAA from moisture. The CO2 taken up in the night is fixed
the cytosol of mesophyll cells to bundle-sheath in the same way as it happens in C4 plants to
cell chloroplasts. The C4 acid is decarboxylated form malic acid, which is stored in the vacuole
to release the fixed CO2, thus generating high (Fig. 3.12). The malic acid thus formed during
concentration of CO2 near Rubisco. The other the night, is used during the day as a source
product of the decarboxylation reaction, the 3- of CO2 for photosynthesis to proceed via the C3
carbon compound (pyruvic acid or pyruvate) is pathway. Thus, CAM is a kind of adaptation

Table 3.1 : Difference between C3 and C4 Photosynthetic Pathways

Features C3 C4
Cell type One (mesophyll) Two (mesophyll and bundle-sheath)
Kranz anatomy No Yes
Chloroplasts One type (granal only) Two types (granal and agranal)
CO2 acceptor RuBP PEP
First CO2 fixation product 3-PGA (3C compound) Oxaloacetic acid (4C compound)
Carboxylase enzyme Rubisco PEPcase; Rubisco
CO2 fixation rate Low High
O2 inhibition of Yes No
photosynthesis
Photorespiration High Negligible
Productivity Low High
CO2 compensation point High (25-100 µl CO2 .l )
-1
Low (0-10 µl CO2 .l-1)
Temperature optimum 20-25°C 30-45°C
Examples Rice, wheat, potato Maize, pearl millet, amaranth
 PHOTOSYNTHESIS
33

Fig. 3.12 Crassulacean Acid Metabolism (CAM) pathway showing CO2 uptake through open stomata
during night and its utilisation for the formation of malic acid which is stored in the vacuole.
During day, the malic acid is decarboxylated to release CO2 which is re-fixed to produce
starch inside chloroplast via C3 Calvin cycle

that allows certain plants (for example, and include leaf age, leaf angle and leaf
pineapple) to carry out photosynthesis without orientation. According to F.F. Blackman, who
much loss of water, which is inevitable in plants postulated Law of Limiting Factors in 1905,
with C3 and C4 mechanisms. photosynthesis is affected by several
environmental and other factors but the role
3.7 FACTORS AFFECTING PHOTOSYNTHESIS of photosynthesis is limited by the slowest step
Photosynthesis is influenced by both in the pathway, or the most limiting factor. This
environmental and genetic factors. The means that at a given time, only the factor that
environmental factors include light, availability is most limiting among all will determine the
of CO2, temperature, soil, water and nutrient rate of photosynthesis. For example, if CO2 is
supply. Genetic factors are all related with leaf available in plenty but light is limiting due to
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cloudy weather, the rate of photosynthesis temperature, as compared to C3 plants is due

under such a situation will be controlled by to a particular enzyme (pyruvate phosphate
the light. Furthermore, if both CO2 and light dikinase) that is required in the C4 pathway.
are limiting, then the factor which is the most In general, different habitats show different
limiting of the two, will control the rate of response of photosynthesis to a given
photosynthesis. temperature. For example, plants adapted to
Light colder environments show a higher
Both quality and intensity of light influence photosynthetic rate at low temperatures than
photosynthesis. Light between the wavelength plants adapted to higher temperatures.
of 400 nm and 700 nm is most effective for Carbon Dioxide
photosynthesis, and this light is called The current level of atmospheric CO2 is about
photosynthetically active radiation (PAR). 0.036 per cent or 360 µl.l–1 (360 ppm), which is
With regard to light intensity, it has a direct very low as compared to the concentration of
relationship with the rate of photosynthesis. other gases in the atmosphere, such as O2 (about
As the intensity of light (µmol.m–2.s–1) increases, 20 per cent) and nitrogen (nearly 80 per cent).
the rate of photosynthesis (CO 2 uptake,
µmolCO2.m–2.s–1) increases. However, this direct
relationship is not seen at higher light
intensities. At higher light intensities, the rate
of photosynthesis decreases. This is because
of two reasons : C4 species (high light)
(i) other factors required for photosynthesis
become limiting, and C3 species
(high light)
(ii) destruction of chlorophyll occurs.
Rate of CO2 Uptake

When the intensity of light falling on leaf

increases beyond a point, chlorophyll is
destroyed. This phenomenon occurs in the
presence of O2 and is called photooxidation.
Carotenoids play a protective role by absorbing
C3 species (low light)
the excess light and diverting it away from
chlorophyll, preventing photooxidation. The
0 100 200 300 400 500 600 700
carotenoids also act as antioxidants, and help –1
CO2 concentration (µ1.1 )
detoxify the bad effect of activated O2 species
on chlorophyll molecules.
Temperature
Photochemical reactions and carbon reactions Fig. 3.13 Response of C3 and C4 plants to CO2
of photosynthesis respond differently to concentration measured as rate of CO2
temperature. While the photochemical uptake. Thick arrows ( ) show the
reactions in thylakoid membrane remain CO2 concentration at which the rate
largely unharmed by temperature, the of photosynthesis get saturated
under high light in C3 and C4 plants.
enzymatic carbon reactions in stroma get
Thin arrows ( ) show the CO 2
influenced adversely. Overall, the process of
compensation point in C 3 and C 4
photosynthesis is sensitive to higher plants. CO2 compensation point refers
temperatures, primarily because of the fact that to CO2 concentration at which the rate
at higher temperatures enzymes become of photosynthesis just balances the
inactive. Affinity of the enzyme Rubisco for CO2 rate of respiration, i.e., at this CO2
is also reduced at higher temperature. Low concentration, the rate of CO2 release
temperature also inactivates enzymes. The during respiration is equal to the rate
sensitivity of C 4 photosynthesis to low of CO2 uptake in photosynthesis
 PHOTOSYNTHESIS
35

The photosynthetic response of C3 and C4 nitrogen is a basic constituent of chlorophyll

plants to the available CO2 concentration is and all enzymes involved in carbon reactions,
variable (Fig. 3.13). In C 3 plants, rate of any reduction in nitrogen supply to plants has
photosynthesis increases with an increase in an adverse effect on photosynthesis. The major
CO2 concentration (up to at least 500 µl.L–1) enzyme of carbon metabolism in plants, that
when other factors are not limiting. This is, Rubisco, alone accounts for more than half
increased rate of photosynthesis in C3 plants of the total leaf nitrogen. In general, all essential
is primarily due to two factors : elements affect the rate of photosynthesis.
(i) high availability of substrate for the Leaf Factors
carboxylation reaction, and Among various leaf factors, such as leaf age,
(ii) reduced photorespiration due to more leaf angle and leaf orientation, leaf age has the
available CO2 to Rubisco. most prominent effect on photosynthesis. If leaf
In C4 plants also, photosynthesis increases as undergoes senescence (that means, it turns
the concentration increases. However, the C4 yellow due to ageing), loss of chlorophyll
plants attain saturation at much lower CO2 occurs. The photosynthetic enzymes also
concentration (around the present level of get deactivated, resulting in reduced
360 µl.L–1) than the C3 plants, which become photosynthesis in a senescent leaf.
saturated at CO2 level of about 500 µl.L–1. This
means that the current availability of CO2 in 3.8 TRANSLOCATION OF PHOTOSYNTHATES
the atmosphere is a limiting factor for C3 plants.
Photosynthates or photoassimilates, i.e. the
Photosynthesis is also influenced by the
energy-rich carbon compounds formed
current increase in the atmospheric CO2. It is
during the process of photosynthesis, are
expected that CO2 concentration could reach
to a level of about 600 µl.L–1 by 2020. In such transported out of the leaf to non-
a case, the C3 plants are likely to be benefitted photosynthetic organs and tissues, such as
more than the C 4 plants. Scientists have roots, stem tissues and developing seeds and
demonstrated that C3 plants can grow faster grains. This long distance transport of
and yield more due to higher rate of photosynthates occurs through phloem and
photosynthesis, when CO2 concentration is is known as translocation. The translocation
raised to 600 µl.L–1. The primary effect of of photosynthates to storage organs plays a
increased atmospheric CO 2 levels on significant role in determining crop yield.
photosynthesis would be through an increase Sucrose is the principal form of carbohydrates
in the intercellular CO2 concentration by an that is translocated from leaf to the non-
increased rate of diffusion of CO2 into the leaf. photosynthetic plant organs. It is a non-
Soil Water reducing sugar, and chemically stable.
Availability of water in soil has a prominent Because of this property, sucrose does not
effect on plant photosynthesis. If soil water react with other substances during
becomes limiting, plants undergo water stress. translocation through phloem.
Under conditions of water stress, the rate of The photosynthates provide energy to
photosynthesis declines because of two factors: tissues through respiration. In storage organs,
(i) stomatal closure and the resultant they are stored in the form of starch or as other
decrease in CO2 supply, and carbohydrates.
(ii) reduced leaf water potential.
3.9 SIGNIFICANCE OF PHOTOSYNTHESIS
Low leaf water potential reduces leaf expansion,
which causes a significant reduction in the Photosynthesis is vital for life on planet
photosynthetic surface area. earth. It helps conversion of the solar
Nutrient Supply energy into organic matter, which makes
Among various nutrients, nitrogen has a direct bulk of the dry matter of any organism.
relationship with photosynthesis. Since The plant biomass or dry matter, derived
 BIOLOGY
36

through photosynthesis supports humans this process for further increasing the
and all other heterotrophic organisms living in productivity of agricultural crops.
the biosphere. Presence of oxygen in the
atmosphere is also an outcome of 3.10 CHEMOSYNTHESIS
photosynthesis. This oxygen is helpful to living Another mode of obtaining energy in some
organisms in two ways : species of bacteria is chemosynthesis, which
(i) in efficient utilisation of the energy-rich is different from photosynthesis. The process
molecules (carbohydrates formed during of carbohydrate synthesis, in which the
photosynthesis) through respiration, and organisms use chemical reactions to obtain
(ii) in making ozone (O3) in the outer layer of energy from inorganic compounds, is called
atmosphere, which helps in stopping the chemosynthesis. For example, bacteria of the
highly destructive ultraviolet (UV) rays genus Nitrosomonas oxidise ammonia to
from reaching the earth. nitrite. The energy released during oxidation,
Without oxygen, life of all aerobic organisms is used by the bacteria in the same way as
including humans, is not possible. plants use energy from sunlight during
Agricultural productivity is also totally photosynthesis, for converting carbon dioxide
dependent on photosynthesis. Scientists are to carbohydrates. Such bacteria are called
currently engaged in genetically manipulating chemosynthetic autotrophs.

SUMMARY

Photosynthesis is one of the most important biological processes occurring

in plants, algae and some bacteria. During this process, carbon dioxide from
the atmosphere is taken in by leaves through stomata, and is used for making
carbohydrates, primarily sucrose and starch. The process occurs inside
chloroplasts, where green pigments called chlorophyll, and other accessory
pigments called carotenoids, are located on thylakoids often stacked to form
grana. Chlorophyll, particularly chlorophyll a, which acts as reaction center,
takes part in the photochemical reactions. Light energy is initially absorbed
by various chlorophyll molecules and accessory pigments, and funneled to
the reaction centers. Together, these pigments form two types of photosystems,
called photosystem I and photosystem II. In photosystem I, chlorophyll a
molecule, which absorbs light maximum at 700nm, acts as the reaction centre
and is referred to as P700. Chlorophyll a that has maximum absorption at
680nm, is the reaction centre in photosystem II and is referred to as P680. The
cascade of events during light reactions involves both photosystems I and II.
During these photochemical reactions, splitting of water molecule initially
occurs in the presence of light, and O2 is released at the photosystem II site.
Electrons released due to splitting of water initiate the electron transport
chain. This leads to the formation of ATP and NADPH2, energy required for
converting CO2 to carbohydrates.
The chemical energy, also called assimilatory power, stored in the form of
ATP and NADPH2, is utilised for CO2 assimilation to carbohydrates. The carbon
reactions for the synthesis of carbohydrates from CO2 take place in the stroma
of chloroplasts, where all necessary enzymes are present, the most notable
being ribulose bisphosphate carboxylase oxygenase (Rubisco). The enzyme
Rubisco catalyses the initial carbon reaction, in which CO2 combines with
ribulose bisphosphate (RuBP), a 5-carbon compound. The product of this
 PHOTOSYNTHESIS
37

reaction is a 3-carbon compound, 3-phosphoglycerate. Hence, this pathway

of carbon reactions is called C3 pathway or the Calvin cycle. Similarly, there
is another pathway of CO2 fixation called as C4 pathway, which is different
than the C3 pathway. The first product of the C 4 pathway is a 4-carbon
compound, oxaloacetic acid. Occurrence of C4 pathway in certain plants (called
C4 plants) in nature is an outcome of a necessity. This necessity was to get
rid off an apparently wasteful process of photorespiration. During
photorespiration, there is a loss of fixed carbon (25 per cent loss) and no
energy rich compound is produced.
There are certain environmental factors that affect the rate of
photosynthesis in plants. Such factors include quality and intensity of light,
CO2, water and temperature. Some other factors, like age of leaf, chlorophyll
content and nutritional status of leaves also affect the rate of photosynthesis.
The end product of photosynthesis is sucrose, which is synthesised in
the cytosol of a mesophyll cell. Starch, however, is synthesised inside
chloroplasts only under certain conditions. The sucrose moves from leaf to
other organs of the plant, where it is used either for growth, or for storage in
storage organs.

EXERCISES

1. (a) Tick ( ü ) the correct answer:

O2 evolved during photosynthesis is from
(i) CO 2
(ii) H2O
(b) Describe very briefly the contribution of the following scientists :
(i) Jan Ingenhousz
(ii) C.B. van Niel
(iii) Joseph Priestley
2. Describe in detail how ATP and NADPH are formed during
photochemical reactions?
3. Describe carbon reactions of the C3 pathway. Does this pathway also
operate in C4 plants?
4. What is photorespiration? Describe the process in detail and link it
with the Calvin cycle.
5. Describe briefly the experiment conducted by T.W. Englemann.
6. Give comparison between the following :
(a) C3 and C4 pathways
(b) Cyclic and non-cyclic photophosphorylation
(c) Carboxylation and oxygenation
(d) Anatomy of leaf in C3 and C4 plants
7. What is a photosystem? Which is the pigment that acts as reaction centre?
Describe the interaction of photosystem I and photosystem II.
8. What led to the evolution of C4 pathway of photosynthesis? Describe
in detail.