ARMY PUBLIC SCHOOL GOPALPUR

CLASS -11 SCIENCE SUBJECT - BIOLOGY

SESSION-2020-2021

CHAPTER-13 PHOTOSYNTHESIS IN HIGHER PLANTS

NOTES
 Photosynthesis, a physico-chemical process by which the green plants use
light energy to drive the synthesis of organic compounds.
 Photosynthesis is the process by which the green plants absorb sunlight to
produce glucose from carbon dioxide and water.
 Photosynthesis is important due to
 it is the primary source of all food on earth
 it is also responsible for the release of oxygen into the atmosphere by green
plants.
EARLY EXPERIMENTS ON PHOTOSYNTHESIS
 Joseph Priestley (1733-1804) in 1770 performed a series of experiments that
revealed the essential role of air in the growth of green plants.
 Priestley observed that a candle burning in a bell jar, soon gets extinguished
and a mouse would soon suffocate in the bell jar because a burning candle or
an animal that breathe the air, both somehow, damage the air.
 When he placed a mint plant in the same bell jar, he found that the mouse
stayed alive and the candle continued to burn.
 Jan Ingenhousz, using a similar setup as the one used by Priestley, but by
placing it once in the dark and once in the sunlight, showed that sunlight is
essential to the plant process that somehow purifies the air fouled by burning
candles or breathing animals.
 Ingenhousz with an aquatic plant showed that in bright sunlight, small bubbles
were formed around the green parts while in the dark they did not, and later
he discovered that these bubbles to be of oxygen.
 Small bubbeles which formed were to be of oxygen; hence, he showed that it
is only the green part of the plants that could release oxygen.
 Julius von Sachs provided evidence for production of glucose in chlorophyll
located in chloroplasts within plant cells and glucose is usually stored as
starch.
 W Engelmann used a prism splitted light into its spectral components and then
illuminated a green alga, Cladophora, placed in a suspension of aerobic
bacteria, which were used to detect the sites of O2 evolution.
 Engelmann observed that the bacteria accumulated mainly in the regions of
blue and red light of the split spectrum, which resembles roughly the
absorption spectra of chlorophyll a and b.
 By the middle of the nineteenth century, the key feature of photosynthesis,
i.e., plants could use light energy to make carbohydrates from CO2 and water,
was known.
 Cornelius van Niel demonstrated that photosynthesis is essentially a light-
dependent reaction in which hydrogen from a suitable oxidisable compound
reduces carbon dioxide to carbohydrates.
 In green plants, H2O is the hydrogen donor and is oxidised to O2, when H2S,
instead is the hydrogen donor for purple and green sulphur bacteria, the
‘oxidation’ product is sulphur or sulphate.
 Hence, it was inferred that the O2 evolved by the green plant comes from H2O,
not from carbon dioxide, and the equation of photosynthesis is

SITE OF PHOTOSYNTHESIS
 The mesophyll cells in the leaves have a large number of chloroplasts, which
align themselves along the walls of the mesophyll cells, such that they get the
optimum quantity of the incident light.
 Within the chloroplast, there is the membranous system consisting of grana,
the stroma lamellae, and the fluid stroma.
 The membrane system is responsible for trapping the light energy and also for
the synthesis of ATP and NADPH.
 In stroma, enzymatic reactions incorporate CO2 into the plant leading to the
synthesis of sugar, which in turn forms starch.
 The set of reactions directly driven by light is called light reaction.
 The reactions which are not directly light driven but are dependent on the
products of light reactions (ATP and NADPH) are known as dark reactions.

Chloroplast

PIGMENTS IN PHOTOSYNTHESIS
 Pigments are substances that have an ability to absorb light, at specific
wavelengths.
 The colour of the leaves is due to four pigments such as
 Chlorophyll a (bright or blue green in the chromatogram),
 chlorophyll b (yellow green),
 xanthophylls (yellow) and
 carotenoids (yellow to yellow-orange)
 The wavelengths at which there is maximum absorption by chlorophyll a, show
higher rate of photosynthesis; hence, we can conclude that chlorophyll a is the
chief pigment associated with photosynthesis.
 Other thylakoid pigments, like chlorophyll b, xanthophylls and carotenoids,
which are called accessory pigments, also absorb light and transfer the energy
to chlorophyll a, and also protect chlorophyll a from photo-oxidation.

 Light reactions or the ‘photochemical’ phase include light absorption,

water splitting, oxygen release, and the formation of high-energy chemical
intermediates, ATP and NADPH.
 The pigments are organised into two discrete photochemical light
harvesting complexes (LHC) within the Photosystem I (PS I) and
Photosystem II (PS II).
 The LHC are made up of hundreds of pigment molecules bound to
proteins.
 Each photosystem has all the pigments (except one molecule of
chlorophyll a) forming a light harvesting system also called antennae.
 The single chlorophyll a molecule forms the reaction centre.
 In PS I, the reaction centre chlorophyll a has an absorption peak at 700
nm, hence is called P700, while in PS II it has absorption maxima at 680
nm, and is called P680.

Light harvesting complex

 THE ELECTRON TRANSPORT
 In photosystem II, the reaction centre chlorophyll a absorbs 680 nm
wavelength of red light causing electrons to become excited and jump, which
are picked up by an electron acceptor which passes them to an electrons
transport system consisting of cytochromes.
 The electrons are passed on to the pigments of photosystem PS I, and the
movement of electrons is downhill.
 Electrons in the reaction centre of PS I are also excited when they receive red
light of wavelength 700 nm and are transferred to another accepter molecule
that has a greater redox potential.
 The electrons then are moved downhill to a molecule of energy-rich
NADP+ and the addition of these electrons reduces NADP+ to NADPH + H+.
 The whole scheme of transfer of electrons is called the z-scheme, due to its
characteristic shape

Z scheme of light reaction

Splitting of Water

 The electrons that were moved from photosystem II must be replaced.

 This is achieved by electrons available due to splitting of water.
 The splitting of water is associated with the PS II; water is split into H+, [O] and
electrons.
 This creates oxygen, one of the net products of photosynthesis.
 The electrons needed to replace those removed from photosystem I are provided
by photosystem II.
 2H2O ——-> 4H+ + O2 + 4e–
 water splitting complex is associated with the PS II, which itself is physically
located on the inner side of the membrane of the thylakoid.

CYCLIC AND NON-CYCLIC PHOTO-PHOSPHORYLATION

 Living organisms have the capability of extracting energy from oxidisable

substances and store this in the form of bond energy.
 Special substances like ATP, carry this energy in their chemical bonds.
 The process of which ATP is synthesised by cells (in mitochondria and
chloroplasts) is named phosphorylation.
 Photo- phosphorylation is the synthesis of ATP from ADP and inorganic
phosphate in the presence of light.
 When the two photosystems work in a series, first PS II and then the PS I, a
process called non-cyclic photo-phosphorylation occurs.
 The two photosystems are connected through an electron transport chain, as
seen earlier – in the Z scheme.
 Both ATP and NADPH + H+ are synthesised by this kind of electron flow.
 When only PS I is functional, the electron is circulated within the photosystem
and the phosphorylation occurs due to cyclic flow of electrons.
 A possible location where this could be happening is in the stroma lamellae.
 While the membrane or lamellae of the grana have both PS I and PS II the
stroma lamellae membranes lack PS II as well as NADP reductase enzyme.
 The excited electron does not pass on to NADP+ but is cycled back to the PS I
complex through the electron transport chain. The cyclic flow hence, results only
in the synthesis of ATP, but not of NADPH + H+.
 Cyclic photophosphorylation also occurs when only light of wavelengths beyond
680 nm are available for excitation.
 Cyclic photophosphorylation

CHEMIOSMOTIC HYPOTHESIS
 The theory which explains how ATP is synthesized in the chloroplast is
chemiosmotic hypothesis.
 Like in respiration, in photosynthesis too, ATP synthesis is linked to
development of a proton gradient across the membranes of the thylakoid.
 In phototsynthesis, proton accumulation is towards the lumen, whereas in
respiration, protons accumulate in the intermembrane space of the
mitochondria when electrons move through the ETS.
 The processes that take place during the activation of electrons and their
transport to determine the steps that cause a proton gradient to develop are
the following:
 Since splitting of the water molecule takes place on the inner side of the
membrane, the hydrogen ions that are produced accumulate within the lumen
of the thylakoids.
 As electrons move through the photosystems, protons are transported across
the membrane because the primary accepter of electron transfers its electron
to an H carrier; hence, this molecule removes a proton from the stroma while
transporting an electron.
 Protons are necessary for the reduction of NADP+ to NADPH+ H+, these
protons are also removed from the stroma.
 Within the chloroplast, protons in the stroma decrease in number, whereas in
the lumen there is accumulation of protons, which creates a proton gradient
across the thylakoid membrane as well as a decrease in pH in the lumen.
 The gradient is broken down due to the movement of protons across the
membrane to the stroma through the transmembrane channel of the F0 of the
ATPase.
 The ATPase enzyme consists of two parts
 one is F0 embedded in the membrane and forms a transmembrane channel
that carries out facilitated diffusion of protons across the membrane
 other portion is called F1 and protrudes on the outer surface of the thylakoid
membrane on the side that faces the stroma.
 The break down of the gradient provides enough energy to cause a
conformational change in the F1 particle of the ATPase, which makes the
enzyme synthesise several molecules of energy-packed ATP.
 Chemiosmosis requires a membrane, a proton pump, a proton gradient and
ATPase.
USE OF ATP AND NADPH
 The phase in which O2 diffuses out of the chloroplast while ATP and NADPH
are used to drive the processes leading to the synthesis of sugar is called
biosynthetic phase of photosynthesis.
 Biosynthetic phase does not directly depend on the presence of light but is
dependent on the products of the light reaction, i.e., ATP and NADPH,
besides CO2 and H2
 CO2 is combined with H2O to produce sugars.
 The use of radioactive 14C by Melvin Calvin in algal photosynthesis studies led
to the discovery that the first CO2 fixation product was a 3-carbon organic acid.
 Calvin worked out the complete biosynthetic pathway; hence, it was called
Calvin cycle.
 The first product identified was 3-phosphoglyceric acid (PGA).
 Another experiment on another plant discovered oxaloacetic acid or OAA as
the first stable product of photosynthesis.
 CO2 assimilation during photosynthesis was said to be of two main types:
 those plants in which the first product of CO2 fixation is a C3 acid (PGA), i.e.,
the C3 pathway,
 the plants in which the first product was a C4 acid (OAA), i.e., the C4

C3 pathway

THE PRIMARY ACCEPTOR OF CO2

 The accepter molecule is a 5-carbon ketose sugar, which is ribulose
bisphosphate (RuBP).
 Since the first product was a C3 acid, the primary acceptor would be a 2-
carbon compound.
 Scientists spent many years trying to identify a 2-carbon compound before
they discovered the 5-carbon RuBP.
THE CALVIN CYCLE
 The Calvin pathway occurs in all photosynthetic plants.
 The Calvin cycle can be described under three stages
 Carboxylation
 Reduction and
 Regeneration

 Carboxylation
 Carboxylation is the fixation of CO2 into a stable organic intermediate, where
CO2 is utilised for the carboxylation of RuBP.
 Carboxylation is catalysed by the enzyme RuBP carboxylase, which results in
the formation of two molecules of 3-PGA.
 Since this enzyme also has an oxygenation activity it is called as RuBP
carboxylase-oxygenase or RuBisCO.
 Reduction
 The steps involve utilisation of 2 molecules of ATP for phosphorylation and 2
of NADPH for reduction per CO2 molecule fixed.
 The fixation of six molecules of CO2 and 6 turns of the cycle are required for
the removal of one molecule of glucose from the pathway.
 Regeneration
 The regeneration steps require one ATP for phosphorylation to form RuBP.
 Regeneration of the CO2 acceptor molecule RuBP is crucial if the cycle is to
continue uninterrupted.
 For every CO2 molecule entering the Calvin cycle, 3 molecules of ATP and 2
of NADPH are required.
 To make one molecule of glucose, 6 turns of the cycle are required.

Calvin cycle

THE C 4 PATHWAY
  Though these plants have the C4 oxaloacetic acid as the first CO2 fixation
product, plants use the C3 pathway or the Calvin cycle as the main
biosynthetic pathway.
 C4 plants are special because
1. they have a special type of leaf anatomy,
2. they tolerate higher temperatures,
3. they show a response to highlight intensities,
4. they lack a process called photorespiration and have greater productivity of
biomass.
5. The particularly large cells around the vascular bundles of the C4 pathway
plants are called bundle sheath cells, and the leaves which have such anatomy
are said to have ‘Kranz’ anatomy.
6. Kranz’ means ‘wreath’ and is a reflection of the arrangement of cells.
7. The bundle sheath cells may form several layers around the vascular bundles.
8. Bundle sheath cells are characterised by having a large number of
chloroplasts, thick walls impervious to gaseous exchange and no intercellular
spaces.
9. The presence of the bundle sheath would help to identify the C4
10. The pathway discovered by Hatch and Slack is called Hatch and Slack
Pathway, which is again a cyclic process.
11. The primary CO2 acceptor is a 3-carbon molecule phosphoenol pyruvate
(PEP) and is present in the mesophyll cells and the enzyme responsible for
this fixation is PEP carboxylase or PEPcase.
12. The mesophyll cells lack RuBisCO enzyme and the C4 acid OAA is formed
in the mesophyll cells.
13. It then forms other 4-carbon compounds in the mesophyll cells itself,
which are transported to the bundle sheath cells.
14. In the bundle sheath cells, these C4 acids are broken down to release
CO2 and a 3-carbon molecule.
15. The 3-carbon molecule is transported back to the mesophyll where it is
converted to PEP again, thus, completing the cycle.
16. The CO2 released in the bundle sheath cells enters the C3 or the Calvin
pathway.
17. The bundle sheath cells are rich in an enzyme ribulose bisphosphate
carboxylase-oxygenase (RuBisCO), but lack PEPcase.
18. The Calvin pathway is common to the C3 and C4
19. In the C4 plants, it does not take place in the mesophyll cells but does so
only in the bundle sheath cells.
 Hatch-Slack pathway

PHOTORESPIRATION
 The first step of the Calvin pathway is the first CO2 fixation step.
 Respiration is the reaction where RuBP combines with CO2 to form 2
molecules of 3PGA, that is catalysed by RuBisCO.
RuBP + CO2 -> 2 X 3PGA
 RuBisCO has a much greater affinity for CO2 than for O2.
 In C3 plants, some O2 do bind to RuBisCO, and hence CO2 fixation is
decreased.
 The RuBP instead of being converted to 2 molecules of PGA binds with O2 to
form one molecule and phosphoglycolate in a pathway called
photorespiration.
 In the photorespiratory pathway, there is neither synthesis of sugars, nor of
ATP.
 In C3 plants, some O2 do bind to RuBisCO, and hence CO2 fixation is
decreased.
 In the photorespiratory pathway, there is no synthesis of ATP or NADPH;
therefore, photorespiration is a wasteful process.
 In C4 plants, photorespiration does not occur because they have a mechanism
that increases the concentration of CO2 at the enzyme site.
 This takes place when the C4 acid from the mesophyll is broken down in the
bundle cells to release CO2, which results in increasing the intracellular
concentration of CO2.

Photorespiration

FACTORS AFFECTING PHOTOSYNTHESIS

 Photosynthesis is under the influence of several factors, both internal and
external.
 The plant factors include the number, size, age and orientation of leaves,
mesophyll cells and chloroplasts, internal CO2 concentration and the amount
of chlorophyll.
 The external factors would include the availability of sunlight, temperature,
CO2 concentration and water.
 When several factors affect any biochemical process, Blackman’s law of
limiting factors comes into effect, which states that “If a chemical process is
affected by more than one factor, then its rate will be determined by the factor
which is nearest to its minimal value: it is the factor which directly affects the
process if its quantity is changed.”
External factors
 Light
 There is a linear relationship between incident light and CO2 fixation rates at
low light intensities.
 At higher light intensities, gradually the rate does not show further increase as
other factors become limiting.
 Increase in incident light beyond a point causes the breakdown of chlorophyll
and a decrease in photosynthesis.
 Carbon dioxide concentration
 The concentration of CO2 is very low in the atmosphere.
 The C3 and C4 plants respond differently to CO2
 At low light conditions, neither C3 nor C4 group responds to high
CO2 conditions, whereas at high light intensities, both C3 and C4 plants show
increase in the rates of photosynthesis.
 C4 plants show saturation at about 360 μlL-1, whereas C3 responds to
increased CO2 concentration and saturation is seen only beyond 450 μlL-1.
 Temperature
 The dark reactions being enzymatic are more sensitive to temperature than
light reaction.
 The C4 plants respond to higher temperatures and show higher rate of
photosynthesis, whereas C3 plants have a much lower temperature optimum.
 Tropical plants have a higher temperature optimum than the plants adapted to
temperate climates.
 Water
 Water stress causes the stomata to close hence reducing the CO2
 Besides, water stress also makes leaves wilt, thus, reducing the surface area
of the leaves and their metabolic activity as well.

Factors affecting photosynthesis

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