NUTRITION IN PLANTS

(28 periods)
Competency Learning outcome Knowledge and understanding
The learner Evaluate the relationship between  Introduction to photosynthesis
evaluates the the structure of chloroplast and (definition, requirements, sites)
nutritional photosynthesis in C3 and C4 plants  Structure of the chloroplast and
strategies of (u, s, gs) function of its parts
carbon three (C3) (Details of biochemistry  Adaptations of the chloroplast
and carbon four are not required.)  Process of photosynthesis in C3 plants
(C4) plants by  Process of photosynthesis in C4 plants
analysing their  Evidence of photosynthesis as a two-
photosynthetic stage process
pathways and  Kranz anatomy in C4 plants and
adaptation to photorespiration
environmental  The efficiency of PEP carboxylase
conditions so as versus Rubisco.
to optimise  Advantages of C4 plants over C3
agricultural plants
productivity and  Synthesis of lipids and proteins
food security Assess the influence of  Structure and adaptations of guard
under varying environmental factors on the cells
climatic photosynthetic efficiency of plants to  Photosynthetic theory of stomatal
conditions. optmise photosynthetic rates and opening and closure
crop yields. (u, s,  Effects of different solutions on
gs, v/a) stomatal opening and closure
 Concepts of limiting factors and
compensation points
 Effect of environmental factors on the
rate of photosynthesis (light, carbon
dioxide, water, temperature, pollution,
mineral salts)
 The distribution and abundance of C3
and C4 plants at different altitudes,
temperatures, and oxygen
concentrations
 Greenhouse (design and principle of
operation)

GENERAL OVERVIEW
Nutrition is the process by which an organism obtains food nutrients and prepares them
for its use.
It’s divided into two
1. Autotrophic nutrition; this involves organisms making their food.
2. Heterotrophic nutrition; this involves organisms feeding on already-made food.

Autotrophic nutrition (Greek: auto – ‘self’; trophic - 'feeding')

Autotrophic organisms use an inorganic form of carbon, such as carbon dioxide, and
energy to form complex organic compounds. Therefore, it is the mode of nutrition in
which an organism makes its food from simple inorganic raw materials, using energy
from the sun or chemical reactions.

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Types of autotrophic nutrition;
(a) Photosynthesis: This uses light energy to initiate reactions that lead to the
formation of organic nutrients. It occurs in all green plants, algae, cyanobacteria,
blue-green bacteria, green sulfur bacteria, purple sulfur bacteria, and colorless sulfur
bacteria. These organisms are known as photoautotrophs.
(b) Chemosynthesis. This uses energy from the oxidation of chemical compounds to
initiate reactions to form organic nutrients. It occurs in Nitrosomonas and
Nitrobacter. These organisms are called chemoautotrophs

Photosynthesis
This is the process by which organisms containing chlorophyll make complex organic
substances or food from simple inorganic molecules of carbon dioxide and water using
light energy trapped by chlorophyll, producing oxygen and water as by-products.

Sunlight
6CO2 + 12 H2O C 6H12O6 + 6O2
Chlorophyll

Importance of photosynthesis
 It provides a source of complex organic molecules for heterotrophic organisms. It
makes both carbon and energy available to organisms. All organisms directly or
indirectly depend on photosynthesis.
 It releases oxygen into the atmosphere that is used by aerobic organisms for
respiration.
 It is how plants capture the sun's energy for use by all organisms
 It avails men of fossil fuels
 The process of photosynthesis reduces the amount of carbon dioxide in the
atmosphere, thus controlling global warming
 Photosynthesis, together with respiration, creates a cycling of carbon dioxide (C0 2)
and oxygen in the atmosphere

Conditions necessary for photosynthesis to take place

(a) Carbon dioxide. This is obtained from the atmosphere and is the source of carbon
atoms present in glucose.
(b) Water. It provides the hydrogen ions and electrons for the reduction of carbon
dioxide. It is also the source of the by-product oxygen.
(c) Light. It provides the energy required to initiate the reactions of photosynthesis.
The three properties of light that are of importance to organisms are spectral
quality/colour, intensity/brightness, and duration/time.
(d) Photosynthetic Pigments. These pigments are of two categories:
i. Chlorophyll. This includes mainly chlorophyll molecules a and b.
ii. Carotenoids. These are accessory pigments that increase the amount and
wavelength of light absorbed for photosynthesis.

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Structure of chloroplasts

They belong to the large group of organelles known as plastids. Chloroplasts are
bounded by a double membrane known as the chloroplast membrane or envelope. The
inner and outer membranes are separated by an intermembrane space
The inner membrane encloses the stroma. The stroma is a colourless, watery, and gel-
like matrix in which outer structures are embedded. Such structures involve a network of
grana interconnected by tubular extensions called integranal lamellae. Each granum is
made up of 2 to 100 closed, flattened sacs called thylakoids stacked on each other.
Within the thylakoids are photosynthetic pigments arranged in funnel-shaped structures
called photosystems.
Also within the stroma are numerous starch grains that temporarily store the products of
photosynthesis. It also contains a small amount of circular DNA, oil droplets, and
phosphate granules.
Adaptations of chloroplasts for photosynthesis
 Membrane system composed of flattened fluid sacs called thylakoids, i.e., the grana
and the intergrana, to increase surface area for maximum absorption of sunlight by
chlorophyll molecules.
 The outer membrane is thin, transparent, and permeable towards respiratory
gases/CO2 and O2
 The matrix, known as stroma, contains enzymes that catalyze the reactions of the
dark stage of photosynthesis.
 It has a system of different electron carriers existing at different energy levels via
which electrons emitted by excited chlorophyll molecules are transferred to release
energy used to form ATP from ADP and phosphate.
 It has a double membrane that allows an uninterrupted photosynthesis as it isolates
photosynthetic reactions from interference by other cellular activities.
 Chloroplasts have their DNA and ribosomes, allowing them to make some of their
proteins (electron carriers, enzymes) quickly for photosynthesis.

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 The internal structures create separate environments for different stages of
photosynthesis, making the process more efficient.
Mechanism of photosynthesis
Photosynthesis is an oxidation-reduction process in which water is oxidized to release
oxygen, and carbon dioxide is reduced to form carbohydrates. Photosynthesis occurs in
two stages
 Light-dependent stage. This involves photochemical reactions and is also called
the light or Hill reaction.
 Light-independent stage. This involves biochemical reactions and is also called
dark or Blackman's reaction.

Light-dependent stage
This is a stage of photosynthesis that requires light energy, which is absorbed by the
photosynthetic pigments (chlorophyll) found in the thylakoids of the chloroplast. These
pigments are located in special reaction centres called photosystems. These systems
convert the absorbed light energy (photons) into chemical energy (in the form of ATP)
The light-dependent stage involves 3 major events, namely photolysis of water,
formation of NADPH+, and photophosphorylation

The main functions of the light-dependent stage are:

o Photophosphorylation. This is the formation of Adenosine triphosphate (ATP) by the
addition of an inorganic phosphate to Adenosine diphosphate (ADP) using light
energy.
o Formation of NADPH+, which is the reduced form of Nicotinamide adenine
dinucleotide phosphate.

Types of photosystems.
i. Photosystem 1 (PSI) with a chlorophyll a molecule called P-700, which absorbs
light of wavelength 700 nm, which functions as the reaction centre. In the P700
photochemical reaction takes place. The pigment system I is located on both the
non-appressed part of grana thylakoids and stromal thylakoids.
ii. Photosystem II (PSII) with chlorophyll a absorbing light of wavelength 680nm (p
680) light energy absorbed excites electrons, which are raised to higher energy
levels and then are accepted by electron carriers (coenzymes). The lost
electrons are replaced by the ones from the photolysis of water or photosystem
II.

Differences between PSI and PSII

PSI PSII
Restabilised by electrons from PSII stabilized by electrons from water
Involved in both cyclic and non-cyclic Involved in non-cyclic
photophosphorylation photophosphorylation
It is not associated with the photolysis of It is associated with the photolysis of
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 water water
PSI is at a higher energy level PSII is at a lower energy level
Its reaction centre is P700 Its reaction centre is P680
Mechanism of photophosphorylation
Light energy in the form of photons is absorbed by the photosynthetic pigments in the
reaction centres called photosystems. In each photosystem, there are several
chlorophyll molecules and accessory pigments, i.e., carotenes and xanthophyll, which
all harvest light energy and pass it onto chlorophyll a in the reaction centres. Each
pigment absorbs light of a different wavelength. There are 2 types of
photophosphorylation, namely cyclic photophosphorylation and non-cyclic
photophosphorylation.

Cyclic photophosphorylation
It is the formation of ATP from ADP + Pi using energy emitted by photo-excited electrons
of P-700 as they flow along photosynthetic electron carriers and back to photosystem I
(P-700).
In this photophosphorylation, light energy absorbed by PSI boosts electrons to a higher
energy level, and the excited electrons are accepted by a ferredoxin (electron acceptor).
From ferrodoxin, electrons are recycled back into PSI directly via a series of electron
carriers, arranged in descending energy levels.
The energy lost by the electrons as they are returned to PSI is captured and used in the
synthesis of ATP from ADP and an inorganic phosphate
By so doing, light energy is converted to chemical energy. The only product of cyclic
photophosphorylation is ATP and involves only PSI
NOTE: Cyclic photophosphorylation takes place most efficiently when the dark reaction
is prevented by either the absence of hydrogen acceptor NADP or the lack of carbon
dioxide.

Non-cyclic photophosphorylation
This is the formation of ATP from ADP + Pi using energy emitted by photo-excited
electrons as they flow unidirectionally through electron carriers from PSII/P680 to
PSI/P700.
The light stage reactions are triggered by light energy, exciting photosystems I and II
inside the thylakoid membranes at the same time, not one after the other.

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Chlorophyll molecules of PSII and PSI are excited by light of wavelength 680 nm and
700 nm, respectively, causing the loss of electrons to a chain of electron carriers in a
series of reduction-oxidation reactions.
This pathway consists of pigment system I (photosystem I) and pigment system II
(photosystem II), which are at different energy levels
Photosystem II is at the lower energy level compared to PSI. When both photosystems
are struck by sunlight of appropriate wavelength, their electrons get excited, and
electrons from PSII are raised to a higher energy level and accepted by the electron
carrier known as plastoquinone (PQ).
Electrons lost from a photosystem II (PSII) are replaced by the ones from the photolysis
of water to restore its neutrality, i.e.
2H2O light energy O 2 + 4H+ + 4e- (free electrons used to restore the neutrality
of PSII). Photolysis is the splitting of water molecules into hydrogen and oxygen using
sunlight energy trapped by chlorophyll.
The electrons from PSII flow from PQ down through a chain of electron carriers to PSI
to replace the lost electrons. As the electrons are carried from a carrier at a higher
energy level to one at a lower energy level, energy is lost that is used in phosphorylation
(ATP synthesis from ADP and Pi). The electrons are received by PSI/p700, which
becomes restabilized.
The electrons from PSI/p700 are then passed to another electron acceptor, Ferrodoxin,
and combine with hydrogen from the photolysis of water.
The electrons and hydrogen reduce Nicotinamide adenine dinucleotide phosphate
(NADP) to form reduced Nicotinamide adenine dinucleotide phosphate /NADPH.
The main Products of non–cyclic photophosphorylation, therefore, are ATP, reduced
NADP/ NADPH2, while the by-product is Oxygen
Non-cyclic photophosphorylation is also called the Z scheme because it involves the
flow of electrons from light-excited chlorophylls in a zig-zag pattern via chains of
electron carriers to form ATP and reduced NADP/NADPH

Comparison between cyclic and non-cyclic photophosphorylation

Similarities
In both:
 There is a flow of electrons through several electron carriers
 Photosystems accept and lose electrons.
 ATP is formed.

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  Pigment system I is involved
 Electron movement is located in the thylakoid membranes
 Protons are moved outwards of the thylakoids.
 Protons (H+) are actively pumped from the stroma into the thylakoid space.
 There is photo-excitation of electrons in the pigment systems.
Differences
Non-cyclic photophosphorylation Cyclic photophosphorylation
Electrons flow unidirectionally (non-cyclically) Electrons flow cyclically
The first electron donor is (source of electrons) water The first electron donor is pigment system I (PSI)
The last electron acceptor is NADP The last electron acceptor is pigment system I (PSI)
The products are ATP, NADPH, and Oxygen The product is ATP only
Involves both pigment systems I and II Involves only the pigment system I
Photolysis of water occurs No photolysis of water

Light-independent reaction or dark stage

The light-independent reactions are called the dark stage because they occur in the
absence of light, provided the products of the light-dependent stage are present. It takes
place in the stroma of the chloroplasts of C3-plants.
The major purpose of the dark reaction is to reduce the CO 2 absorbed from the
atmosphere and water to carbohydrates.
This requires ATP and reduced NADP from the light stage of photosynthesis. ATP
provides energy for the endergonic with action reaction of the dark stage, and reduced
NADP provides hydrogen atoms required to reduce CO2 to carbohydrates.
The main pathways for the dark reaction are Calvin-Benson cycle / C 3 pathway and
Hatch-Slack pathway / C4 pathway

Calvin cycle
It involves a cycle of reactions named after Melvin Calvin. The C3 cycle is a series of
reactions in plants to form glycerate-3-phosphate as the first stable organic substance
during photosynthesis.
C3 Plants are plants that fix CO2 directly in glycerate-3-phosphate/G.P., which is a 3-
carbon organic compound as the first stable product during photosynthesis. OR. These
are plants whose first stable compound of carbon dioxide fixation is a 3-carbon organic
compound called PGA/G.P./Phosphoglycerate.
Main stages of the Calvin-Benson cycle (C3 cycle)
1. Carboxylation stage
This occurs in the stroma of the chloroplast of the mesophyll cells. CO 2, which has
diffused into the stroma of the chloroplast, reacts with a 5-carbon compound, ribulose
Bisphosphate, under the catalysis of ribulose Bisphosphate carboxylase
enzyme/RUBISCO to produce an unstable 6-carbon compound.
The 6-carbon compound splits into two molecules of 3 carbon compound, the first stable
product of photosynthesis called phosphoglyceric acid (PGA) or glycerate -3- phosphate
/phosphoglycerate.

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Some of the Phosphoglycerate is used for the synthesis of amino acids and fatty acids
needed for the synthesis of proteins and lipids, respectively.
The fixing of CO2 by RUBP is called carboxylation of RUBP or carbon dioxide fixation,
i.e.
CO2 + RUBP (5C) RUBP unstable 6C organic compound
Carboxylase
2. Reduction stage
The remaining portion of phosphoglycerate is reduced by hydrogen atoms donated by
reduced NADP, using energy from hydrolysis of ATP to form phosphoglyceraldehyde
(PGAL)/triose phosphate/glyceraldehyde-3-phosphate/GALP/3-phosphoglyceraldehyde.
Water molecules are released as PGA is reduced to the Aldehyde PGAL.
ATP ADP
PGA/G. P PGAL + H2O
NADPH2 NADP+
Part of PGAL is used for the synthesis of glycerol. The triose phosphate is the end
product of photosynthesis.
3. Isomerisation and condensation
The remaining PGAL passes via a series of reactions to form monosaccharide sugars,
mainly hexose sugars, which condense into sucrose and starch. Two of the three 3-
phosphoglyceraldehyde molecules undergo isomerisation and several reactions to form
fructose-1-phosphate and glucose-1-phosphate, both of which may condense to form
sucrose or starch.

4. Regeneration of ribulose Bisphosphate (RUBP)

Another portion of PGAL is used for the regeneration of RUBP via several enzyme-
catalyzed reactions, using energy from the hydrolysis of ATP into ADP.
In the regeneration of RUBP, 5 PGALs are used to regenerate 3 RUBPs, this process
requires ATPs, and rearrangement of the carbon atom in the sugar phosphate to
generate 5 carbon compounds from 3 carbon compounds.
3ATP 3ADP+ 3P i
5 PGAL rearrangement of carbon chain 3RUP 3RUBP
5. Product synthesis stage: -
Product of photosynthesis (T.P) is assimilated through different pathways, including;
 Is converted into sucrose, a form in which it’s translocated either in storage organs
or growing points.
 It is fed into the glycolytic pathway (respiration) to produce energy required for
endergonic reactions. T.P/GPAL enters the Glycolytic pathway, where it is converted
into acetyl-CoA, which enters the Krebs cycle

Synthesis of lipids:
Lipids are formed from glycerol, which is formed directly from T.P/PGAL and fatty acids,
which are obtained from phosphoglycerate/PGA/G.P.

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PGA/G. P enters the glycolytic pathway to form Acetyl co-enzyme A, which is then used
to synthesize fatty acids, which finally react with glycerol through a condensation
reaction, forming lipids.

Synthesis of proteins: -
The Triose Phosphate is fed into the Krebs cycle after converting it to acetyl-CoA.
Proteins are formed from amino acids, which are also formed from
phosphoglycerate/glycerate-3-phosphate.
Phosphoglycerate via the acetyl-CoA enzyme A forms an intermediate organic acid of
the Krebs cycle.
The intermediate Krebs cycle acid reacts with ammonia from the reduction of nitrates
obtained from the soil to form amino acids, which are used in protein synthesis.

SUMMARY OF THE CALVIN CYCLE

Light-independent mechanisms of photosynthesis in C 4-plants.

C4 Plants are the ones whose first formed stable compound of carbon dioxide fixation is
a 4-carbon compound known as oxaloacetate.
C4 plants include maize, sugar cane, millet, sorghum, and many tropical grasses. These
are plants that are mainly monocots that produce a 4-carbon compound called
oxaloacetic acid (OAA) as the first stable product of carbon dioxide fixation.
They undergo two pathways of photosynthetic reactions, which include the Hatch-Slack
pathway and the Calvin cycle.

Hatch-slack pathway
A type of photosynthetic pathway in which CO2 is first fixed by phosphoenol pyruvate,
catalyzed by PEP carboxylase (PEP), into Oxaloacetate (OAA) inside mesophyll cells.

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Oxalo-acetate (OAA) is reduced by hydrogen from Nicotinamide adenine dinucleotide
phosphate, Hydrogen /reduced NADP to form a 4-carbon compound called Malate.
Fixation of CO2 by PEP to form Malate occurs in the mesophyll cells.
OAA (4C) Malate (4C)

NADPH2 NADP+
The malate produced in the mesophyll cells diffuses through the plasmodesmata and
then diffuses into the chloroplasts of the bundle sheath cells, a process called the
malate shunt.
Within the chloroplasts of the bundle sheath cells, malate is dehydrogenated to give a
large amount of H. + ions and decarboxylated to form CO2 and pyruvate. The C4
pathway pumps CO2 and H+ ions into the bundle sheath cells where they are used by
the normal Calvin’s cycle.
CO2 Calvin cycle
Malate (4C) pyruvate (3C)

NADP+ NADP H2
The H+ ions produced are used to reduce NADP to form reduced NADP/NADPH +,
whose synthesis is limited to the bundle sheath cells.
The formed CO2 in the bundle Sheath cells is fixed by RUBP under the catalysis of
RUBP carboxylase to form organic food substances via a series of reactions.
The pyruvate diffuses back into the mesophyll cells where it is phosphorylated using 2
molecules of ATP to regenerate the carbon dioxide acceptor, PEP.
Pyruvate
(3C)
PEP (3C)

ATP
ADP +Pi

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Note.
1. Carbon dioxide fixation in the mesophyll cells does not occur inside their
chloroplasts because they lack the RUBP carboxylase enzyme.
2. PEP carboxylase has a much higher affinity for CO 2 than RUBP carboxylase, and
therefore, a higher level of carbon dioxide is fixed into the carbohydrate metabolism,
forming a larger amount of food energy than in C3 plants.
3. Because of the high concentration of CO2 fixed by PEP under the initial catalysis of
PEP carboxylase, RUBP carboxylase only catalyses the fixation of CO2 rather than
oxygen. The high CO2 concentration in the chloroplasts of the bundle sheath cells
outcompetes O2 for the RUBP carboxylase active site. Hence, it prevents
photorespiration in C4 plants, ensuring efficient CO2 fixation by RUBP carboxylase.
4. PEP’s high affinity for CO2 also makes it unable to fix oxygen instead of CO2.
5. Most C4 plants have high photosynthetic yield in the tropics and subtropics regions
with high temperatures and high light intensity due to their ability to fix a high
concentration of CO2.
6. C4 plants yield more food materials than C3 plants because they don’t photorespire.

Kranz anatomy in the leaves of C4 plants

C4 plants have a characteristic leaf Anatomy which is described as Kranz anatomy due
to arrangement of 2 distinct rings of leaf cells around the vascular bundles each with a
different type or form of chloroplasts, where by the inner ring of cells are called the
bundle sheath surrounded by the outer ring referred to as the mesophyll cells.
Chloroplasts in the C4 plants show some Dimorphism, i.e., they exist in two forms.
Those of the bundle sheath cells have rudimentary grana, whereas the grana are
prominent in the mesophyll cells.

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 Advantages and significance of the C4 pathways
 It makes C4 plants more adapted to high light intensities and temperatures of the
tropics due to higher optimum temperatures of PEP carboxylase for CO2 fixation.
 The dimorphic nature of the chloroplast limits photorespiration greatly, leading to
efficient fixation of CO2 by RUBP carboxylase and higher yield, i.e., higher
productivity in C4 plants
 PEP carboxylase does not accept O2, thus limiting photorespiration
 Closing of stomata during the day by desert plants does not affect the rate of
photosynthesis due to large stores of CO2 at night, as malate
 The high affinity of PEP carboxylase for CO2 leads to high productivity of
photosynthesis.
 Because they can reduce the aperture of their stomata during high light intensity and
hot conditions, they lose less water by transpiration.

Disadvantages
1. Consumes a lot of energy compared to C3 pathways.

Comparison of C3 and C4 plants

Similarities
Both:
 contain RUBISCO enzyme
 depend on light for their reactions
 show CO2 fixation
 have RuBP
 form several of the same organic products, e.g., PG, PGA, sucrose
 have the Calvin cycle

Diff C3 PLANTS C4 PLANTS

structural Lack Kranz anatomy Exhibit Kranz anatomy
All chloroplasts have the identical Chloroplasts are dimorphic (have two types of
structure (have one type of chloroplasts), e.g., those of palisade cells have
chloroplast) grana yet lack bundle sheath cells.
Physiological CO2 acceptor is a 5-carbon RuBP CO2 acceptor is a 3-Carbon PEP
CO2 fixation occurs once CO2 fixation occurs twice
Photorespiration occurs No photorespiration

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 Less photosynthetically efficient More photosynthetically efficient
G.P. is the first stable organic product
OAA is the first stable organic product
Enzymes are more efficient at lower Enzymes are more efficient at high
temperatures(20-250C) temperatures (30- 35 °C)
Use only the RUBISCO enzyme for PEP carboxylase and RUBP carboxylase
CO2 fixation enzymes are used
The compensation point is attained when
The compensation point is attained at Oxygen is not an inhibitor of the
a higher CO2 concentration photosynthetic process, at a lower CO2
concentration
Oxygen is an inhibitor of Oxygen is not an inhibitor of photosynthesis.
photosynthesis.
Grows at a low rate Grows at a high rate

Note
The C3 plants can survive best in an environment of low temperature, low light intensity,
low oxygen level, and high CO2 level.
The C4 plants can survive best in an environment of high temperature, high light
intensity, high oxygen level, and high CO2 levels.
Photorespiration
This is the light-dependent uptake of oxygen by RUBP carboxylase and the output of
CO2, which mainly occurs in C3 plants.
It's a wasteful process in which carbon fixation in C 3 plants is prevented due to the light-
dependent uptake of oxygen by RuBP carboxylase (RUBISCO enzyme) and release of
carbon dioxide.

Effects of photorespiration on plants

When C3 plants are exposed to low carbon dioxide concentration (or high oxygen
concentration), e.g., when stomata close to reduce water loss, RuBP carboxylase
catalyses the reaction between RuBP and oxygen to form a 2-carbon compound,
phosphoglycolate, which is oxidized to release carbon dioxide.

This means that carbon dioxide and oxygen compete for the active site of the enzyme.
Hence, oxygen is a competitive inhibitor during carbon dioxide fixation. Because RUBP
carboxylase enzyme catalyses fixation of both CO 2 and O2 due to the affinity it has for
both of them, it can be referred to as RUBISCO/Ribulose bisphosphate carboxylase-
oxygenase enzyme.

When C3 plants are exposed to low carbon dioxide or high oxygen concentrations,
instead of RUBP carboxylase accepts O2, leading to the production of one PGA/G.P.
molecule and a phosphoglycolate. This reduces PGA yields by ½.
Plants recover the lost carbon in the phosphoglycolate by converting it into PGA/G.P.
through a series of reactions that occur in 3 cell organelles, i.e., chloroplasts,
peroxisomes, and mitochondria. Phosphoglycolate is immediately dephosphorylated
into glycolate.

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To recover another molecule of G.P., two molecules of glycolate via a series of reactions
are used to form one molecule of G.P., and one remaining carbon atom is lost in the
form of CO2 without net production of ATP.
Conditions for photorespiration
 Low CO2  High light intensity
 High O2 concentration  High temperatures

Disadvantages of photorespiration: -
1. It cuts the yield of photosynthesis by half
2. There is a loss of carbon, which reduces the productivity of the plant.

Evidence of photosynthesis as a two-stage process

The following evidences indicate that the overall reaction in photosynthesis takes place
in two steps: one is light-dependent and the other is light-independent.

(a) Temperature coefficient studies

The rate of photosynthesis of two groups of plants was compared. Both were supplied
with an adequate concentration of carbon dioxide. But one group was kept under a light
of high intensity, and the other group under a light of low intensity. When the rate of
photosynthesis was measured at different temperatures, it was found that the high light
group had a Q10 = 2, but the low light group Q10 = 1. Strictly, chemical reactions
characteristically have a Q10 from 2 to 3. This fact indicates that at least one of the
reactions involved in photosynthesis is of a purely chemical type. This reaction is called
the dark reaction. The other reaction of Q 10 indicates that one of the reactions proceeds
only at the expense of absorbed light; it is called the dark reaction. The Q10 of the light
reaction is 1

(b) Flashing light experiments (effect of intermittent light)

Photosynthesis involves photochemical and biochemical reactions, is also shown by the
results of investigations in which plants are exposed to intermittent light. Warburg
(1919) obtained higher rates of photosynthesis in Chlorella when it was exposed to
rapid and alternate periods of light and darkness instead of continuous illumination.
Emerson and Arnold (1932) found that at 250 °C, the maximum photosynthesis took
place when light and dark periods were respectively 10 -5 seconds and 0.055 seconds.
At 1.10C, maximum photosynthesis could be obtained with the same light period, but the
dark period has to be increased to 0.4 seconds. It means that temperature influences
reactions of the dark period, and reactions of the light phase are photochemical.

Mechanisms of Stomatal opening and closure

A stoma is a pore in the epidermis of a plant surrounded by two guard cells through
which gaseous exchange and transpiration occur.

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Structure of guard cells
It is sausage-shaped with an uneven thickened cell wall, i.e. thicker inelastic inner
cellulose cell wall (wall lining the pore) but a thinner elastic outer cellulose cell wall (wall
furthest from the pore); the microfibrils of the cellulose cell wall are arranged in hoops.
Its cytoplasm has a peripheral nucleus, a central large vacuole, and chloroplasts.

Photosynthetic theory of stomatal movement

It states that:
During the day, guard cells carry out photosynthesis and manufacture sugars that
accumulate and increase the osmotic pressure of the guard cells beyond that of the
surrounding epidermal cells, resulting in osmotic movement of water from the
neighboring epidermal cells to the guard cells, which become turgid, and the stomata
open
However, at night, there is little sugar is manufactured due to the absence of light, thus
increasing the osmotic pressure of the guard cells. Consequently, they lose water by
osmosis to the neighboring epidermal cells, the guard cells become flaccid, and the
stomata close.
Effect of different concentrations of solutions on stomatal opening and closure
(a) Hypotonic Solution (Low solute concentration)
Water enters the guard cells by osmosis. The guard cells become turgid (swollen) and
the outer, thin, and elastic wall bulges outwards while the thick inner wall invaginates.
This causes the stomata to open, hence more gaseous exchange and transpiration.
(b) Hypertonic Solution (High solute concentration)
Water leaves the guard cells by osmosis. The guard cells become flaccid (shrink),
causing the stomata to close, reducing water loss.
Effect of environmental factors on the rate of photosynthesis
These include
 Quantity and quality of light incident on leaves
 Suitable temperature
 Concentration of carbon dioxide in the surrounding atmosphere
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  Concentration of oxygen in the surrounding atmosphere
 Availability of water
 Inorganic ions; absence of ions like Mg, N, and Fe, chlorophyll can’t be
synthesized
 Level of pollution

The principle of limiting factors

It states that when a chemical process depends on more than one essential condition
being favourable, its rate is limited by that factor nearest its minimum value/in its short
supply. I.e., it’s this factor that directly affects the process if the quantity is changed.
For example, photosynthesis can’t proceed in the dark because the absence of light
limits the process.

When one factor is favourable, e.g., when light is increased, the rate of photosynthesis
increases until it levels off because another factor, other than light intensity, limits the
rate of photosynthesis. But when the limiting factor, such as CO 2, is increased, the rate
of photosynthesis further increases until yet another factor, like temperature, tends
towards its minimum and limits the rate of photosynthesis, resulting in its leveling off.

Effect of carbon dioxide

Observation/description Explanation
Generally, the rate of photosynthesis  RuBISCO attaches carbon dioxide instead of
increases rapidly with increasing carbon oxygen, because the carbon dioxide
dioxide concentration to a maximum at 30 Pa concentration is higher than the oxygen

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in C4 plants and 90 Pa in C 3 plants, and concentration.
thereafter remains constant. More cells photosynthesize because of the
increased carbon dioxide molecules available.
The rate of photosynthesis is faster in C4 than PEP Carboxylase of C4 has a higher affinity for
in C3. carbon dioxide than RuBISCO of C3 and hence
acts faster.
The overall photosynthetic products are C4 needs more ATP than C3, which generally
greater in C3 than in C4 reduces photosynthetic output
The C4 plants are more efficient at lower CO2 At lower CO2 concentration in C3
concentration, while C3 plants are more photorespiration reduces the photosynthesis
efficient at higher CO2 efficiency, yet PEP Carboxylase has a high
affinity for carbon dioxide.
The C3 plant has a higher compensation point PEP Carboxylase has a high affinity for carbon
than the C4 plant dioxide
After attaining the maximum, the rate of It is because other factors limit the process, e.g.,
photosynthesis remains constant in both. temperature, light intensity, etc.
At the CO2 concentration of about 70 Pa, the
rate of photosynthesis is equal in both plants.

Carbon dioxide is a raw material for the dark stage of photosynthesis in that it’s reduced
by hydrogen donated by reduced NADP via a series of reactions to form carbohydrates.
High Carbon dioxide concentration in the atmosphere increases the rate of
photosynthesis significantly.

In the atmosphere, the concentration of carbon dioxide ranges from 0.03 to 0.04 %.
However, the highest CO2 level needed for photosynthesis is 0.1%, beyond this
optimum CO2 concentration, the rate may reduce due to the inactivation or denaturation
of the photosynthetic enzymes due to the acidic PH as a result of the formation of
carbonic acid from the reaction of excess CO2 with water.

Temperature
Most reactions of photosynthesis are catalyzed by enzymes; hence require an optimum
temperature for optimum enzyme activity for a high photosynthetic rate.
At temperatures below the optimum (around 0°C), the photosynthetic rate is reduced
due to the inactivation of enzymes, and when the temperature is increased beyond the
optimum, the rate of photosynthesis reduces until the reactions stop because of the
denaturation of enzymes until all of them are fully denatured. Different species of plants
have different optimum ranges of temperature; most temperate plants need an optimum
range of temperature between 20-25 oC, while tropical plants need 35- 40 °C.
Since both stages of photosynthesis require enzyme activity, the temperature affects
the rate of photosynthesis.

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 Observation/description Explanation
Below 100C, the C3 rate C4 photosynthetic enzymes are less
of
photosynthesis is higher than in C 4 above active in the cold but become more
100C. active with an increase in temperature.
The maximum rate of photosynthesis The optimum temperature for enzymes
attained in C4 is much higher than in C3. involved in the C4 cycle is higher than in
the C3 cycle.
At about 45°C, the rate of Enzymes controlling photosynthesis
photosynthesis decreases are denatured
There is an initial increase in Light intensity becomes a limiting
photosynthetic rate to a maximum at factor in each of the three cases.
about 400C to 420C, despite a further
increase in temperature.
There is an increase in the rate of An increase in temperature activates
photosynthesis with an increase in enzymes to a level beyond which
temperature until up to about 400 enzyme denaturation occurs.

Water
Water is a metabolite/raw material for photosynthesis. Water is split by light energy to
provide hydrogen ions needed in the dark stage of photosynthesis. It also provides
electrons, which stabilise the PSII/photo system II after it has emitted its electrons.

It is found that even a slight deficiency of water results in a significant reduction in the
crop yield. The lack of water not only limits the amount of water but also the quantity of
carbon dioxide. This is because in response to drying, the leaves close their stomata to
conserve water being lost as water vapour through them.

Pollution
Pollution of the atmosphere with industrial gases has been found to result in as much as
15% loss. Soot can block stomata and reduce the transparency of the leaves. Some of
the other pollutants are ozone and sulphur dioxide. Lichens are very sensitive to sulphur
dioxide in the atmosphere. Pollution of water affects the hydrophytes. The capacity of
water to dissolve gases like carbon dioxide and oxygen is greatly affected.

Mineral salts

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Mineral salts such as nitrates, phosphates, magnesium, and nitrogen affect chlorophyll
formation. In a high concentration of mineral ions, there is a high production of
chlorophyll molecules. Total absence results in chlorosis, hence, no photosynthesis will
take place.
Salinity
One of the major effects of salinity is osmotic stress, and hence there are intimate
relationships to drought stress or ‘water stress’. This results in stomata closure to avoid
desiccation, which reduces photosynthesis because the uptake of CO2 is reduced.

Oxygen
High concentration of oxygen, mainly in C3 Plants reduces the rate of photosynthesis
because oxygen competes with CO2 for the active site of ribulose bisphosphaate
carboxylase enzyme used to catalyze the fixation of CO 2 by RUBP, carbon dioxide
acceptor into an unstable 6C intermediate compound, this is the enzyme has an equally
high affinity for oxygen unlike CO2. Because this enzyme is called RUBISCO (ribulose
bisphosphate carboxylase-oxygenase.

Light Intensity and Compensation Point

When light intensity is Low, the rate of photosynthesis is low. As the intensity is
increased, the rate of photosynthesis also increases. This is because light is used
during the light stage of photosynthesis to provide energy in the form of ATP and
reduced NADPH (Nicotinamide adenine dinucleotide phosphate hydrogen). ATP and
reduced NADP, as products of the light stage, are required as raw materials in the dark
stage of photosynthesis.
Light is also needed during the photolysis of water, resulting into the production
hydrogen. The hydrogen ions produced during photolysis are used to reduce NADP to
NADPH +H+.
Light is used to raise the energy level of electrons of chlorophyll molecules for them to
be emitted and passed via electron carriers at different energy levels to produce energy
needed to combine ADP with inorganic Phosphate to form ATP.

As the light intensity is increased, the rate of photosynthesis also increases. However,
after reaching an intensity of 10,000 lux (lux is the unit for measuring light intensity),
there is no effect on the rate. Very high intensity may slow down the rate as it bleaches
the chlorophyll. Normal sunlight (usually with an intensity of about 100,000 lux) is quite
sufficient for a normal rate of photosynthesis.
When a plant in a region of low light / in darkness is provided with high light intensity, its
rate of photosynthesis increases until it equals the rate of respiration, whereby there is
no net release of oxygen by the plant to the atmosphere.
When the rate of photosynthesis is equal to the rate of respiration, the plant is said to
have reached its Compensation point. This happens at dawn and dusk.

The light compensation point is the light intensity at which the rate of respiration is equal
to the rate of photosynthesis. At this point, CO 2 is neither evolved nor absorbed, i.e.,
there is no net loss or gain in CO2, and there is no net loss or gain in carbohydrates, and
there is no net exchange of O2 and CO2.

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Compensation point: The light intensity at which the photosynthetic intake of carbon
dioxide is equal to the respiratory output of carbon dioxide. It occurs during early
morning or late evening.
Beyond the compensation point further increase in light intensity results in a
proportional increase in the rate of photosynthesis until light saturation is reached.
Beyond this point, further increases in light intensity do not affect the rate of
photosynthesis unless some other factors, like CO2 has its concentration, are
increased. The period taken for the plant to reach the compensation point is known as
the compensation period.

Shade plants have a shorter

compensation period than those in bright light/light plants for their maximum and
efficient utilization of light.
At very low light intensity, shade plants have higher CO 2 uptake, which reduces with
illumination.
Light plants have a higher compensation point than shade plants.
At a certain light intensity, the rate of CO2 uptake is the same in both.
In both, CO2 increases with an increase in illumination to a maximum and then levels
off.
Shade plants reach maximum CO2 uptake at a lower illumination than light plants.
Increased illumination causes a bigger increase in CO 2 uptake in light plants than in
shade plants.
Letter P represents the compensation point at which CO2 uptake equals CO2 output.
At Y, biomass decreases because the rate of respiration exceeds that of
photosynthesis.

Effect of altitude (and oxygen)

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 Observation/ Explanation
description
C3 plants are more The decrease in atmospheric pressure at higher
abundant at high altitudes decreases the partial pressure of oxygen,
altitudes/elevations enabling more productivity since photorespiration
reduces
CAM plants are more Even when the temperature is high, nocturnal
abundant at low altitudes stomatal opening and closure in daylight enable them to
reduce transpiration.
CAM plants that store a lot of malates and, due to the
thus high osmotic value, also a lot of water, are usually
less frost resistant than C3 plants.
C4 plants are widely The enzymes are tolerant to these high temperatures,
distributed at low and the Kranz mesophyll anatomy shields RuBISCO in
altitudes and slight bundle sheath cells from much oxygen to avoid
elevations photorespiration.

Effect of water stress on the rate of photosynthesis

Water stress significantly affects the rate of photosynthesis by influencing various
physiological and biochemical processes in plants. Here are the key effects:
1. Closure of Stomata – Water stress causes stomata to close to prevent water
loss through transpiration. However, this also reduces CO₂ uptake, which limits
the availability of carbon dioxide for the Calvin cycle, thereby reducing the rate of
photosynthesis.
2. Reduced Chlorophyll Content – Prolonged water stress can degrade
chlorophyll pigments, leading to lower light absorption and a decrease in the
efficiency of photosynthesis.
3. Impairment of Photosynthetic Machinery – Water deficiency can damage the
photosystem complexes (PSI and PSII), reducing the efficiency of light-
dependent reactions and electron transport in the chloroplasts.
4. Oxidative Stress – Water stress increases the production of reactive oxygen
species (ROS), which can damage proteins, lipids, and chloroplast membranes,
further inhibiting photosynthesis.
5. Decline in ATP and NADPH Production – Limited water supply reduces the
photolysis of water (H₂O) in photosystem II, which decreases oxygen evolution
and limits the production of ATP and NADPH, essential for carbon fixation in the
Calvin cycle.

Greenhouse: design and principle of operation

A greenhouse is a structure designed to create a controlled environment for growing
plants. It traps heat from the sun to maintain warm conditions even when it is cold
outside. The greenhouse farming involves manipulation of environmental factors that
affect the rate of photosynthesis to optimize crop yields.

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Design of a Greenhouse
 Structure: Usually made of a metal or wooden frame covered with transparent
materials like glass or plastic.
 Roof: Often sloped to allow rain runoff.
 Walls: Transparent to let sunlight enter.
 Doors and Vents: For air circulation and temperature regulation.
 Optional Additions:
o Heating systems (in colder climates).
o Irrigation systems.
o Shading for very hot days.

Principle of Operation (How it Works)

1. Sunlight enters through the transparent walls and roof.
2. The sunlight is absorbed by plants and soil, converting it into heat.
3. The heat is trapped inside because the glass/plastic prevents it from escaping
easily — this is called the greenhouse effect.
4. The inside stays warmer than the outside, even during cold weather.
5. This creates ideal conditions for plant growth — warm temperatures, protection
from pests, and controlled humidity.
Benefits of a Greenhouse
 Grows crops all year round.
 Protects plants from harsh weather (wind, frost, heavy rain).
 Improves crop yield and quality.
 Allows control of water, temperature, and light.

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