S.

6 notes-Nutrition in plants | LEK

S6 BIOLOGY NOTES
NUTRITION IN PLANTS AND ANIMALS
Nutrition is the process by which organisms obtain energy to maintain life functions, and matter
to create and maintain structure. Both energy and matter are obtained from nutrients.
Modes of nutrition
Organisms are categorized into two groups basing on their source of carbon i.e.
1. Heterotrophic Nutrition (heteros, other; trophos, nourishment): Where organisms
depend on organic nutrients obtained from other organisms due to their inability to
manufacture their own. Such organisms have an organic source of carbon and are referred
to as heterotrophs. This is further categorized into saprophytism, mutualism,
commensalism, parasitism and holozoic nutrition
2. Autotrophic nutrition (autos, self; trophos, nourishment): where organisms make their
own organic nutrients from an external supply of relatively simple inorganic raw
materials and energy. Such organisms have an inorganic source of carbon, namely carbon
dioxide and are referred to as autotrophs.
Types of autotrophic nutrition
This is categorized into two groups basing on the source of energy
(i) Photosynthesis: This is the form of nutrition that occurs in all green plants, algae
some protists and photosynthetic bacteria (cyanobacteria). It is the process by
which organisms synthesize organic compounds sugars, protein and lipids from
carbon dioxide and water using sunlight as source of energy and chlorophyll or
some other closely related pigment for trapping the light energy.
(ii) Chemosynthesis: this is form of nutrition that occurs in certain bacteria see table
below. This is the synthesis of organic compounds from carbon dioxide and water
using energy supplied by special methods of respiration involving the oxidation of
various inorganic materials such as hydrogen sulphide, ammonia and iron (ii).
Table 1: Examples of chemosynthetic bacteria
Bacteria Inorganic material Product Habitat
+ -
1. Nitrosomonas and Ammonium (NH4 ) Nitrite (NO2 ) Soil
Nitrococcus
2. Nitrobacter Nitrite (NO2-) Nitrate ((NO3-) Soil
3. Ferrobacillus / Iron Ferrous (Fe2+) Ferric (Fe3+) Streams flowing over
bacteria iron containing rocks
4. Hydrogen bacteria Hydrogen (H2) Water (H2O) Soil
5. Colourless Sulphur Hydrogen sulphide Water and Decaying organic matter
bacteria (H2S) sulphur
Importance of Photosynthesis
• It is the means by which the sun's energy is captured by plants for use by all organisms.
• It provides a source of complex organic molecules for heterotrophic organisms.
• It releases oxygen for use by aerobic organisms.

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THE LEAF AS AN ORGAN OF PHOTOSYNTHESIS

Although stems, sepals and other parts of the plant photosynthesize, the leaf is the main organ for
photosynthesis in plants.
Adaptations of plants for photosynthesis (a) Adaptations for obtaining sunlight
 Phototropism causes shoots to grow towards the light in order to allow the attached leaves
to receive maximum illumination.
 Etiolation causes rapid elongation of shoots which are in the dark, to bring leaves into
light to capture light.
 Mosaic leaf arrangement minimizes leaf overlap and reduces shading of one leaf by
another.
 Broad lamina to provide Larger leaf surface area enabling capturing maximum sunlight.
 Leaves held at an angle perpendicular to the sun during the day to expose the maximum
area to light.
 Thinness of leaves enables easy penetration of light to lower layers.
 The cuticle and epidermis are transparent to allow light penetration into the
photosynthetic mesophyll beneath.
 The palisade mesophyll cells are densely packed with chloroplasts and arranged with their
long axes perpendicular to the surface to form a continuous layer which traps most of the
incoming light.
 Chloroplasts within the mesophyll cells are capable of moving (Cyclosis) allowing them
to arrange themselves into the best positions within the cells for efficient absorption of
light.
 The chloroplasts hold chlorophyll in an ordered / structured way on the sides of the grana
to present maximum chlorophyll to the light and also bring it close to other pigments /
substances necessary for functioning.
 In leaves of sun plants the palisade layer, whose cells are densely packed with
chloroplasts is more than one cell thick to increase on photosynthetic efficiency.
 In leaves of shade plants, the cells of palisade and spongy mesophylls are densely packed
with chloroplasts to increase on light trapping hence photosynthetic efficiency.
(b) Adaptations for obtaining and removing gases
 Numerous stomata in the epidermis of leaves providing a larger surface area for diffusion
of gases.
 Guard cells bordering stomata pores that open and close stomata to regulate the uptake of
carbon dioxide and the loss of water.
 Numerous airspaces in spongy mesophyll for faster and uninterrupted diffusion of gases
between the atmosphere and the palisade mesophyll.
(c) Adaptations for obtaining and removing liquids
 A large central midrib containing a large vascular bundle comprising xylem and phloem
tissue in most dicotyledonous leaves, the xylem for the entry and transport of water and
mineral salts, and the phloem for carrying away sugar solution, usually in the form of
sucrose.
 A network of small veins is found throughout the leaf to ensure that every cell is close to
xylem vessel or phloem sieve tube for constant supply of water for photosynthesis and a
means of removing the sugars they produce.

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Assignment
a) Outline ways in which the structure of a green leaf facilitates photosynthesis
b) What are the principle functions of leaf veins?

Structure of chloroplast
Is biconvex disc-shaped/ oval shaped, 3 – 10 µm long and 2 – 4 µm wide, enclosed by an envelope
of two membranes; the smooth and continuous outer membrane, the inner membrane gives rise
to strands of branching membranes called the lamellae extending throughout the organelle. The
interior is divided into grana which are surrounded by a, gelatinous semi-fluid called stroma. In
the grana the lamellae are stacked in piles of flat, circular sacs called thylakoids, which contain
photosynthetic pigments. In the stroma the thylakoids are criss-cross loosely, suspended in an
aqueous matrix containing circular DNA, ribosomes, ribosomes, enzymes used in photosynthesis.

Adaptations of chloroplast for photosynthesis

 Biconvex shape which increases surface area for exposure of photosynthetic pigments for
maximum light absorption.
 Surrounded by a double membrane to prevent photosynthetic reactions from mixing with
those in the cell cytoplasm.
 The surface membrane is permeable to allow exchange of materials like carbon dioxide
which is a raw material for photosynthesis with the cell cytoplasm.
 The inner membrane is folded inwards to form a system of layers called lamellae to
provide a large surface area for attachment of photosynthetic pigments.
 The internal membrane also contains electron transport systems for synthesis of ATP to
drive cell metabolism.
 It has thylakoids that increase the surface area for holding chlorophyll molecules.
 The thylakoid granum is connected by intergrana membranes thus maintaining the
thylakoids and chlorophyll stationary in position.
 The stroma contains circular DNA and ribosomes for protein synthesis.
 The stroma contains a high concentration of the necessary enzymes for catalyzing
metabolic reactions occurring within the chloroplast.
 Thylakoids are flattened discs to provide a small internal volume to maximize hydrogen
gradient upon proton accumulation.
 Thylakoids stacked in piles forming grana to increase the surface area to volume ratio of
the thylakoid membrane.
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 Pigments organized into photosystems in thylakoid membranes to maximize light

absorption.

THE REQUIREMENTS FOR THE PROCESS OF PHOTOSYNTHESIS

1. Carbon dioxide:
Carbon dioxide is a raw material for photosynthesis and is a source of carbon for the organic
compounds produced in the process. Terrestrial plants obtain carbon dioxide: -
 from the atmosphere (where it’s about 0.03%) via the stomata
 By absorbing carbonates from the soil through the roots, aquatic plants absorb dissolved
bicarbonates through their general surface to carbon dioxide.
2. Water
Water is a raw material for photosynthesis. It is a source of hydrogen for reduction of carbon
dioxide and also an essential donor of electrons to chlorophyll during non-cyclic phosphorylation
which results in production of ATP and NADPH2 both essential for carbon fixation in the light
independent stage.
Water is also essential for the general metabolism of the plant and lack of it to the extent of
causing wilting of leaves results in physiological stress on metabolism which in turn affects
photosynthesis either directly or indirectly.
Qn . Why is it difficult to demonstrate the importance of water to photosynthesis?

3. Temperature
Photosynthesis proceeds by a series of chemical reactions controlled by enzymes. Suitable
temperature is required for activation of enzymes that catalyse photosynthetic reactions.
4. Light
There are three features of light which make it biologically important
1. Spectral quality (color)
2. Intensity (brightness)
3. Duration (time)
To be of use as an energy source for organisms, light must first be converted to chemical energy.
Radiant energy comes in discrete packets called quanta. A single quantum of light is called a
photon. Light also has a wave nature and so forms a part of the electromagnetic spectrum.
Visible light represents that part of this spectrum which has a wavelength between 400nm
(violet) and 700 nm (red) see figure 2 below.

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Figure2: The section of the electromagnetic spectrum

As sunlight falls on a plant, some wavelengths are absorbed, while others are reflected or
transmitted. The absorbed light provides energy that excites electrons in the photosystems whose
transition releases energy for synthesis of ATP, also light is necessary for photolysis of water to
release electrons stabilizing photosystems and hydrogen atoms forming NADPH2 for reduction
of carbon dioxide.
5. Pigments
Photosynthetic pigments of higher plants are categorized into two groups i.e. chlorophylls and
carotenoids. The algae in addition have phycobiliproteins like phycocyanin (blue) and
phycoerythrin (red). The role of the pigments is to absorb light energy, thereby converting it into
chemical energy. They are located on chloroplast membranes (thylakoids) and the chloroplasts
are usually arranged within the cells so that the membranes are at right angles to the light source
for maximum light absorption (a) Chlorophylls
There are several types of chlorophyll, all containing a ring structure called porphyrin ring with
magnesium at the center linked to a long hydrocarbon chain (figure 3)

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The most common chlorophylls

include chlorophyll a, chlorophyll b,
some photosynthetic bacteria have
another type of chlorophyll called
bacteriochlorophyll, which contains
manganese instead of magnesium.
Chlorophylls absorb light in the
blueviolet and the red region of the
visible spectrum. The remaining light,
in the green region of the spectrum, is
reflected and gives chlorophyll its
characteristic color.

Figure 3: Structure of chlorophyll

molecule
(b) Carotenoids
Carotenoids are yellow, orange, red or brown pigments they absorb strongly in the blue-violet
range of the spectrum.
Roles of carotenoids
 They are accessory pigments; they absorb light energy and pass the light they absorb on
to chlorophyll.
 They protect chlorophylls from excess light and from oxidation by oxygen produced in
photosynthesis.
Carotenoids are of two types, carotenes and xanthophylls, these are usually masked by the green
chlorophylls but can be seen in leaves before leaf-fall because chlorophylls break down first. A
common example of a carotene is β-carotene which gives carrots their familiar orange color. It is
easily formed into two molecules of vitamin A.
Absorption and action spectra
If a pigment such as chlorophyll is subjected to different wavelength of light, it absorbs some
more than others. If the degree of absorption at each wavelength is plotted, an absorption
spectrum of that pigment is obtained (figure 4).

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Figure 4: Absorption spectrum of chlorophylls and carotenoids

An absorption spectrum is a graph of the relative amounts of light absorbed at different
wavelengths of light by a pigment.

An action spectrum is a graph showing the effectiveness of different wavelengths of light in

stimulating the process being investigated. Figure 5 below shows an action spectrum for
photosynthesis together with an absorption spectrum for combined photosynthetic pigments.

Figure 5: Action spectrum for photosynthesis compared with absorption spectrum of

photosynthetic pigments.
From figure 5 above: -
 the red and blue ends are the most effective wavelengths in photosynthesis;  green is
only used to a slight extent.
 There is a close correlation between the absorption and action spectrum, indicating that
the pigments, chlorophylls in particular, are responsible for the absorption of light used in
photosynthesis.

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 The non-correspondence in the two spectra between about 450nm and 470nm, is because
this wavelength is absorbed by carotenes which are not used in photosynthesis.

Assignment: Explain the similarities and differences between the absorption and action
spectra in figure 5 above.
Discovery of the role of red and blue light in photosynthesis
In an investigation Engelman, used a species of
motile aquatic aerobic bacterium. Where these
bacteria are found to accumulate, he knew that
oxygen was also present.
Engelman split sunlight into its constituent
colors by means of a prism, and projected them
onto cells so that the different colors of light
were received by different parts of the filament.
The aerobic bacteria in the water collected
around the filaments in areas where the
chloroplast was receiving red and blue light.
Conclusion
Light in the violet-blue and red portions of the
spectrum is most effective in driving
photosynthesis.

Figure 6: results of Engelmann’s experiment

MECHANISM OF PHOTOSYNTHESIS
The overall equation for photosynthesis is:
Light
6CO2 + 12 H2O C6H12O6 + 6H2O + 6O2
Chlorophyll
Photosynthesis is essentially a process of energy transduction. Light energy is first converted into
electrical energy and then into chemical energy in three main phases i.e.
1. Light harvesting. Light energy is captured by the plant using a mixture of pigments
including chlorophyll.
2. Light dependent stage (photolysis) in which a flow of electrons results from the effect
of light on chlorophyll and so causes the splitting of water into hydrogen ions and oxygen
3. The light independent stage during which these hydrogen ions are used in the reduction
of carbon dioxide and hence the manufacture of sugars.
LIGHT HARVESTING
The photosynthetic pigment molecules are clustered in the thylakoid membranes. Each cluster is
called an antenna complex

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Special proteins associated with these pigments channel light energy entering the chloroplast on
to special molecules of chlorophyll a, known as the reaction center chlorophyll molecule. The
reaction Centre and all the other light-gathering molecules combine to form a photosystem.
When light strikes this molecule, an electron in its orbit is raised to a higher energy level, thus
initiating a flow of electrons.
There are two types of photosystems; photosystem I and photosystem II.
In photosystem I, the reaction Centre is called P700 because its chlorophyll a has a maximum
absorption at a wavelength of 700nm (red light).
Photosystem II has a reaction Centre called P680 because its chlorophyll a has a maximum
absorption at 680nm (orange-red).

Figure 7: A photosystem: a light-harvesting cluster of photosynthetic pigments in chloroplast

thylakoid membrane
Exercise 1
1. (a) Explain how a photosystem increases the light harvesting ability of a chloroplast?
(b) Explain the relationship between the action spectrum and the absorption spectrum of
photosynthetic pigments in green plants.
Evidence that photosynthesis is a two-stage process
(i) The overall process is influenced by increase in temperature which would not be the case
if it was dependent on light alone.
(ii) The amount of carbohydrate produced in a given quantity of light is greater if the light is
supplied intermittently/in flashes rather than continuously suggesting that some part of
photosynthesis is independent of light.
(iii) By use of radioactive tracers like C14 in C14O2 which have subsequently been detected in
the organic compounds produced. It has been shown that the reduction of carbon dioxide
can occur in the absence of light.

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THE LIGHT-DEPENDENT STAGE

How light trapped by chlorophyll is used
1. Provides energy to convert ADP and an inorganic phosphate (Pi) to ATP a process called
photophosphorylation
2. Necessary for the splitting of water molecules to release electrons and hydrogen ions a
process known as photolysis.
Importance of light dependent stage of photosynthesis
1. produces Adenosine triphosphate (ATP) which is a source of energy for subsequent
synthesis of carbohydrates.
2. Photolysis of water produces hydrogen atoms for the reduction of carbon dioxide during
the dark stage.
The light dependent reactions of photosynthesis
The light-dependent reactions occur in the thylakoid membranes of a chloroplast’s grana. It
involves the splitting of water by light (photolysis of water) to give hydrogen ions (protons) and
the synthesis of ATP in photophosphorylation. The hydrogen ions combine with a carrier
molecule NADP to make reduced NADP. ATP and reduced NADP are passed from the light
dependent to the light independent reactions.
Photophosphorylation of ADP to ATP can be cyclic or non-cyclic, depending on the pattern
of electron flow in one or both types of photosystem. (i) Cyclic photophosphorylation
Cyclic photophosphorylation involves only photosystem I. Light of wave length 700nm is
absorbed by photosystem I (P700) and is passed to the primary pigment. An electron in the
chlorophyll molecule is excited to a higher energy level and is emitted from the chlorophyll
molecule. This is called photoactivation. Instead of falling back into the photosystem and losing
its energy as thermal energy or as fluorescence, the excited electron is captured by an electron
acceptor and passed back to a chlorophyll molecule via a chain of electron carriers i.e.
ironprotein complex, to cytochromes b, to plastoquinone, to cytochrome-f, to plastocyanin and
again back to P-700.
The flow of electrons through carriers in the thylakoid membrane releases energy for active
pumping of hydrogen ions (H+) from the stroma to the thylakoid space.
The highly concentrated H+ inside the thylakoid space diffuse along the steep electrochemical
gradient from the thylakoid space via the stalked particles into the stroma, thereby providing
energy to form ATP in the presence of ATPase enzyme. this process is called chemiosmosis. The
ATP then passes to the light independent reactions.

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Figure 8: Summary of events that occur during cyclic photophosphorylation

(ii) Non-cyclic photophosphorylation
Non-cyclic photophosphorylation involves both photosystems I and II in the so-called ‘Z
scheme’ of electron flow (unidirectional electron flow) (Figure 9).
Light strikes both photosystems I and II simultaneously, excited electrons are emitted from the
primary pigments of both reaction centres. These electrons are absorbed by electron acceptors
and pass along chains of electron carriers, leaving the photosystems positively charged. The
electrons from photosystem II are passed from the electron acceptor along a series of electron
carriers to photosystem I. The primary pigment at photosystem II receives replacement
electrons from the splitting (photolysis) of water.
Photosystem II includes a water-splitting enzyme that catalyses’ the breakdown of water:
H2 O 2H+ + 2e- + 1/2 O2
Oxygen is a waste product of this process. The hydrogen ions combine with electrons from
photosystem I and the carrier molecule NADP to give reduced NADP.
2H+ +2e- +NADP reduced NADP
Reduced NADP passes to the light independent reactions and is used in the synthesis of
carbohydrate.

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Figure 9: The Z-Scheme

As in cyclic photophosphorylation, ATP is synthesized as the electrons lose energy while passing
along the carrier chain.
The movement of electrons in the thylakoid membranes releases energy which enables active
pumping of hydrogen ions (H+) from the stroma to the thylakoid space. At the same time,
photolysis of water
(i) causes accumulation of H+ inside the thylakoid space (ii)
provides electrons to replace those lost from PSII.
The high accumulation of H+ photolysis and active pumping of proton creates a steep
electrochemical gradient between the thylakoid space and stroma, resulting in diffusion of H+ via
the stalked particles into the stroma this provides
(i) energy to form ATP in the presence of ATPase enzyme (ii)
H+ for reducing NADP to form NADPH.
The NADPH and ATP formed then enter the dark stage.

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Figure 10: Summary of events that occur during non-cyclic photophosphorylation

COMPARISON BETWEEN CYCLIC AND NON-CYCLIC PHOTOPHORYLATION

Similarities
In both
▪ there is flow of electrons through electron carriers ▪ there are pigment systems which
accept and lose electrons.
▪ ATP is formed.
▪ 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 stroma into thylakoid space.
▪ there is photo-excitation of electrons in the pigment systems.
Differences
Non-cyclic photophosphorylation Cyclic photophosphorylation
▪ Electrons flow unidirectionally (noncyclically) ▪ Electrons flow in a cyclic pattern

▪ First electron donor is (source of electrons) ▪ First electron donor is photosystem I

water
▪ Last electron acceptor is NADP ▪ Last electron acceptor is photosystem I
▪ The products are ATP, NADPH and Oxygen ▪ The product is only ATP.

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▪ Involves both photosystems I and II ▪ Involves only photosystems I

▪ Photolysis of water occurs ▪ No photolysis of water
▪ Two electron acceptors involved ▪ Only one electron acceptor involved
Note
During cyclic photophosphorylation no oxygen and NADPH produced as photolysis of
water does occur,
Relatively less energy is produced in cyclic photophosphorylation than in non-cyclic
photophosphorylation since non-cyclic photophosphorylation involves two photosystems
of which each takes up a quantum of light.
When carbon dioxide concentration is limiting, both photosystems cannot operate at the
same time hence only photosystem I operates and photophosphorylation is mainly cyclic.
THE LIGHT-INDEPENDENT STAGE
This also referred to as the dark stage because the reactions can take place in the dark if sufficient
ATP and NADPH are available. It occurs in the stroma of the chloroplast and takes place
whether or not light is present. The reactions are controlled by enzymes and their sequence was
determined by Melvin Calvin, Benson and Bassham during a period of 1946-53. The process is
often called the Calvin cycle.
The dark reactions involve main pathways which include
▪ Calvin-Benson cycle / C3 pathway ▪
Hatch-Slack pathway / C4 pathway
1. Calvin-Benson cycle / C3 pathway
This is the series of reactions in plants involving formation of glycerate-3-phosphate which has 3
carbon atoms as first stable organic substance during photosynthesis.
MAIN STAGES OF C3 PATHWAY
1. Carboxylation
Carbon dioxide diffuse into the stroma and then combines with a 5-carbon sugar, Ribulose
biphosphate (RUBP), in a reaction catalysed by enzyme RuBP carboxylase (Rubisco), the
resulting 6 carbon compound is unstable and immediately breaks down to form two molecules of
3-carbon compound known as 3-phosphoglyceric acid (PGA)/ glycerate-3-phosphate (GP),
which is the first stable organic compound in C3 plants.
2. Reduction phase
3-phosphoglyceric acid (PGA) molecules are phosphorylated by ATP from the light stage ADP,
and then reduced by NADPH (formed in light stage) to form a triose phosphate (TP) called
3phosphoglyceraldehyde (PGAL) or glyceraldehyde-3-phosphate(GALP), which is a 3-carbon
sugar, NADP+, ADP and an inorganic phosphate (Pi).
Note:
Triose phosphate is the first stable carbohydrate formed in the Calvin cycle.
NADP+ is regenerated and this returns to the light dependent stage to accept more hydrogen

3. Regeneration phase
Five-sixth of the triose phosphates are converted through a series of reactions into RUBP which
then fixes more carbon dioxide. This reaction requires both ATP and NADPH from the light
stage.
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4. Product synthesis phase

One-sixth of the triose phosphate molecules are used to produce other molecules needed by the
plant. Some of these triose phosphates condense to become hexose phosphates which, in turn,
are used to produce starch for storage, sucrose for translocation around the plant, or cellulose for
making cell walls. Others are converted to glycerol and fatty acids to produce lipids for cellular
membranes or to acetyl coenzyme A for use in respiration or in the production of amino acids
for protein synthesis.

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Figure 11: the Calvin cycle

Figure 12: Summary of process of photosynthesis

Exercise 2
1. (a) Outline the light-independent reactions of photosynthesis.
(b) Explain:
(i) why the light-independent reactions of photosynthesis can only continue for a
short time in darkness.
(ii) how the light-independent reactions of photosynthesis rely on light-dependent
reactions.
2. (a) Outline the formation of carbohydrate molecules in photosynthesis starting from the
absorption of light energy
(b) Compare the structure of a chloroplast and a mitochondrion in relation to function.
3. (a) Explain photophosphorylation in terms of chemiosmosis in chloroplast.
(b) Explain the reactions involving the use of light energy that occur in the thylakoids of the
chloroplast.

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4. Experiments on cultures of a unicellular protist to investigate the effect of light and

carbon dioxide on certain metabolites. In the first experiment, the levels of PGA, RuBP
and sucrose in the protest were determined at different time intervals in the presence of
light. At the 35th minute, light was switched off, suddenly putting the protists in darkness;
the results are shown in the table below
Time (minutes) 0 20 35 40 50 60 70
Amount of RuBP 35 35 35 30 15 10 10
metabolite PGA 45 45 45 50 65 70 70
Sucrose 10 54 72 66 52 35 20
(a) Represent the data provided graphically
(b) Using the graph obtained in (a) above, explain the variation in the
levels of the metabolites with time
5. In an experiment, samples of algae were collected at 1-minute intervals over a period of 5
minutes. The quantities of glycerate-3-phosphate (GP) and ribulose bisphosphate (RuBP)
were measured. At the beginning of the experiment, the concentration of carbon dioxide
supplied was high. After 2 minutes, the concentration of carbon dioxide was reduced. The
graph in the figure below shows the results of this experiment.
(a) Describe the effects of the
decrease in carbon dioxide after 2
minutes on:
(i) Glycerate 3-phosphate
(GP)
6. (ii) Ribulose bisphosphate (RuBP)
(b) Suggest explanation for these
changes to the levels of glycerate
3phosphate (GP) and RuBP.

Metabolism of Glycerate phosphate (GP) and Glyceraldehyde phosphate (TP/ PGAL)

(a) Synthesis of carbohydrates
Glyceraldehyde-phosphate molecules are converted to form monosaccharides e.g. glucose.
Glucose may combine with fructose to form sucrose, transported in phloem sieve tubes or can be
polymerized into starch for storage or cellulose; a component of plant cell walls.
(b) Synthesis of lipids
• Glycerate-phosphate enters glycolysis pathway and is converted to pyruvate, which can be
converted into acetyl group, which combines with coenzyme A to form acetyl coenzyme A.
This can be used to form a variety of fatty acids in the cytoplasm and chloroplast.
• Glycerate-phosphate can also be converted to glycerol
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Lipids such as triglycerides are esters of fatty acids and glycerol, which are important components
of cell membranes. (c) Synthesis of proteins
Glycerate-phosphate is converted into acetyl coenzyme A and enters into the Krebs cycle. Some
of its intermediates can produce different amino acids by transamination reactions. The amino
acids are then polymerized into proteins which are required for growth and development, synthesis
of enzymes and structural components of the cell.
NB:
The nitrogen, Sulphur and phosphorus required for protein synthesis are absorbed from the soil.
Nitrogen is taken up as nitrates or ammonia, Sulphur as sulphates and phosphorus as phosphates.

Figure 13: Summary of metabolism of intermediates of dark stage

Assignment: Compare light dependent and light independent stages of photosynthesis
NOTE
 The enzyme Ribulose biphosphate carboxylase that catalyses the reaction of carbon dioxide
with RuBP unfortunately, it can also catalyse the reaction of oxygen with RuBP. When this
happens, less photosynthesis takes place, because some of the RuBP is being ‘wasted’ and
less is available to combine with carbon dioxide. This unwanted reaction is known as
photorespiration. It happens most readily in high temperatures and high light intensity –
that is, conditions that are found at low altitudes in tropical parts of the world.

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 Photorespiration is a wasteful process in which carbon fixation in C3 plants is prevented due

to the light dependent uptake of oxygen by RuBP carboxylase (RUBISCO enzyme) and
release of carbon dioxide
 Tropical grasses such as maize, sorghum and sugar cane which are C4 plants have evolved a
method of avoiding photorespiration. They keep RuBP and rubisco well away from high
oxygen concentrations. The cells that contain RuBP and rubisco are arranged around the
vascular bundles, and are called bundle sheath cells the arrangement known as the Kranz
anatomy. They have no direct contact with the air inside the leaf.
2. HATCH-SLACK PATHWAY OR C4 METABOLISM
This is a type of photosynthesis in which CO2 is first, fixed by phosphoenolpyruvate carboxylase
(PEPCO) into Oxaloacetate (OAA) inside mesophyll cells, stored as organic acid (mainly malate)
which is later decarboxylated, refixed and CO2 is assimilated in the Calvin‐cycle inside bundle
sheath cells.
Examples of C4 plants: maize, sorghum, Amaranthus, Sugar cane, paspalums (Paspalum
notatum), Bermuda grass, blue grama, Rhodes grass, troublesome weeds like nut grass, crabgrass
and barnyard. They are found mainly in hot / arid / saline tropical habitats.
WHAT IS KRANZ LEAF ANATOMY?
A condition in which bundle sheath cells and palisade cells of the mesophyll form two concentric
layers (rings) around each vascular bundle of leaves.
COMPARISON OF LEAF ANATOMY IN C3 AND KRANZ ANATOMY IN C4 PLANTS

QN. How do the chloroplasts of mesophyll cells differ from those of bundle sheath in C4
plants?

THE HATCH- SLACK PATHWAY

Leaf anatomy in C3 plants Kranz anatomy in C4 plants This is a pathway for transporting
carbon dioxide and hydrogen from mesophyll cells to bundle sheath cells. Once in the bundle sheath
cells, the carbon dioxide is released again and normal C3 photosynthesis occurs. Stages in C4
pathway
1. acceptance of carbon dioxide (carbon dioxide fixation) in mesophyll cells In the
presence of phosphenol pyruvate carboxylase (PEPCO) enzyme, the carbon dioxide acceptor
with 3 carbon atoms, phosphenol pyruvate (PEP) combines with carbon dioxide inside the
chloroplasts of mesophyll cells to form oxaloacetate (OAA) a 4-carbon compound. This is the

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first stable compound formed in C4 plants. Oxaloacetate is reduced by NADPH from the
light stage to malate a 4-carbon acid. This occurs in the presence of malate dehydrogenase
enzyme.
2. Malate shunt
From chloroplasts of mesophyll cells, the malate is translocated (shunted) to the chloroplasts of
bundle sheath cells where it is decarboxylated and dehydrogenated by NADP to form pyruvate a
3-carbon acid and carbon dioxide. The pyruvate produced returns to mesophyll cells for
phosphorylation by ATP to regenerate PEP; the CO2 acceptor.
Now the second carboxylation occurs in the chloroplasts of bundle sheath cells through Calvin
cycle.
3. Regeneration of the carbon dioxide acceptor
Pyruvate is returned to the mesophyll cells and is used to regenerate PEP by the addition of
phosphate from ATP. This requires the energy from two high energy phosphate bonds.

Figure 14: Summary of Hatch-slack pathway

ADVANTAGES OF HATCH_SLACK PATHWAY

 C4 plants ably photosynthesize at very low CO2 concentration (e.g. in dense tropical
vegetation) because PEP carboxylase enzyme has a very high affinity for carbon dioxide.
 Concentric arrangement of mesophyll cell produces a smaller area in relation to volume for
better utilization of available water and reduce, the intensity of solar radiations.
 Photorespiration, which inhibits growth in C3 plants is avoided / reduced in C4 because
▪ the CO2 fixing enzyme PEP carboxylase has no affinity for oxygen
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▪ RUBISCO enzyme inside the bundle sheath cells is shielded from high oxygen
concentration by the ring of palisade cells.
 The CO2 fixing enzymes in C4 plants are more active at hot temperature and high
illumination, therefore photosynthesis occurs rapidly at low altitude, hot and brightly lit
tropical conditions than in C3 plants.
 The productivity of C4 almost four times greater than in C3 because:
(i) of the increased rate of CO2 uptake caused by
▪ large internal leaf surface area
▪ short CO2 diffusion distance
(ii) CO2 steep diffusion gradients in the bundle sheath cells in which dark reactions occur
have ▪ a large photosynthetic surface area enabled by un-usually large chloroplasts
▪ lack of grana on which O2 would be produced, so no photorespiration.
▪ the Palisade cells in which light reactions occur have large grana to increase the
photosynthetic surface area.
Disadvantages of hatch-slack pathway
 The CO2 fixing enzymes in C4 plants are less active at cool temperature and low
illumination, therefore photosynthesis occurs slowly at high altitude with cool temperature
and in low light intensity of temperate conditions.
 Since every carbon dioxide molecule has had to be fixed twice, the energy requirements for
C4 photosynthesis is roughly double that for C3 photosynthesis.
COMPARISON BETWEEN C3 AND C4 PLANTS
Similarities Both:
▪ contain RUBISCO enzyme
▪ depend on light for their reactions
▪ show CO2 fixation
▪ have RuBP
▪ form several same organic products e.g. PG, PGA, sucrose
▪ have the Calvin cycle

Differences
C3 Plants C4 plants
 Lack Kranz anatomy  Exhibit Kranz anatomy
 All chloroplasts have identical  Chloroplasts are dimorphic (are in two forms) e.g.
structure those of palisade cells have grana yet are lacking
bundle sheath cells.
 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
 Less photosynthetically efficient  More photosynthetically efficient

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 GP is the first organic product  OAA is the first organic product

 Enzymes are more efficient at lower  Enzymes are more efficient at high temperatures
temperatures
 RUBISCO enzyme is used  PEP carboxylase enzyme is used
 Compensation point is attained at  Compensation point is attained at lower CO2
higher CO2 concentration concentration

CRASSULACEAN ACID METABOLISM (CAM) PHOTOSYNTHESIS

This is a type of photosynthesis in which CO2 is taken in at night via open stomata, fixed by
phosphoenolpyruvate carboxylase (PEPC) into OAA, stored as organic acid (mainly malate) which
is later decarboxylated during daytime, refixed and CO2 is assimilated in the Calvin‐cycle when
stomata are closed.
Examples of CAM plants
Cacti, agaves (sisal), opuntia, Kalanchoe (Bryophyllum), Vanilla (family: Orchidaceae),
pineapples (Family: Bromeliaceae), Mesembryanthemum crystallinum (Common ice plant), and
Euphorbia milii (Crown of Thorns plant). These are mainly succulent plants that live in hot arid
climates.
When the stomata open at night, carbon dioxide enters the leaves and combines with PEP to form
OAA in the presence of an enzyme PEPCO found in their cytoplasm. The OAA is then reduced to
malate a reaction catalysed by an enzyme malic dehydrogenase which accumulates in the leaf
vacuoles.
During the day when the stomata are closed the malate is transported to the cytoplasm where it is
decarboxylated to pyruvate and carbon dioxide, the carbon dioxide released enters the chloroplast
where it is fixed to sugars in the Calvin cycle.

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Figure 15: A representation of the CAM cycle Assignment: State the differences between
metabolism in C4 plants and CAM plants Exercise 3
1. (a) Distinguish between light compensation point and compensation period (02 marks)
(b) (i) Explain why C3 plants have a higher carbon dioxide compensation point compared to
the C4 plants. (02 marks)
(ii) Suggest the physiological advantages of C4 having low carbon dioxide
compensation point compared to C3 plants. (03 marks)
(c) State any three differences between the mesophyll and bundle sheath chloroplasts in C4
plants. (03 marks)

2. (a) What is meant by the term C4 plant? (02 marks)

(b) (i) explain the significance of C4 plants for being more efficient at carbon dioxide
fixation than C3 plants. (06 marks)
(ii) Explain how carbon dioxide is fixed by C4 plants. (06 marks)
(c) How are leaves of C4 plants modified to suit it for carbon dioxide fixation? (06 marks)

Significance of CAM photosynthesis

For terrestrial CAM plants, there is increased water use efficiency in which nocturnal stomatal
opening greatly reduces stomatal loss of water as it would in day light.
Note
 CAM is an adaptation for hot and dry conditions. It enables the plant to conserve water by
keeping stomata closed in the heat of the day when transpiration would be most rapid
 CAM plants are extremely efficient at conserving water but unfortunately their rate of net
photosynthesis per unit area of plant or ground is very low and correspondingly their growth
rates are also very low.

MEASUREMENT OF THE RATE OF PHOTOSYNTHESIS

Rate of photosynthesis can be measured by measuring the rate of:- 
uptake of CO2
 production of O2
 production of carbohydrates
 increase in dry mass
(a) Measuring the rate of Uptake of CO2
Uptake of CO2 can be measured with the means of an IRGA (Infra-Red Gas Analyser) which can
compare the CO2 concentration in gas passing into a chamber surrounding a leaf / plant and the
CO2 leaving the chamber. The soil and roots must NOT be in the bag to avoid CO2 production
from respiration
NOTE: CO2 uptake can also be measured by following the uptake of carbon dioxide labelled
with 14C
(b) Rate of Production of carbohydrates
This is a crude method where a disc is cut out of one side of a leaf (using a cork borer against a
rubber bung) and weighed after drying. Some weeks later, a disk is cut out of the other half of the

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leaf, dried and weighed. Increase in mass of the disc is an indication of the extra mass that has
been stored in the leaf.
Explain why this method is inaccurate.

(c) Measuring the increase in dry mass-

Dry mass is often monitored by the technique of 'serial harvests' where several plants are
harvested, dried to constant weight and weighed - this is repeated over the duration of the
experiment so as to have an accurate measure of the surplus photosynthesis over and above the
respiration that has taken place. As with most methods, several plants are needed to have replicate
measurements which are used to calculate the average and a standard deviation if necessary.
(d) Measuring the rate of production of O2
Oxygen can be measured by (a) counting bubbles evolved from pond weed with the Audus
apparatus Requirements
Test tube plastic tube Previously well
Watch connector illuminated aquatic plant
Water at room graduated scale e.g. Elodea or Cabomba
temperature retort stand 0.2 % sodium bicarbonate
bench lamp to provide soft board solution
light thermometer plastic Syringe
Knife capillary tube
3
500 cm glass beaker Ruler
Procedure:
 Set up the apparatus as below in TOTAL DARKNESS

 A light source is placed 50 cm away facing the test tube and is powered on, a 5 minutes lapse
is allowed to enable the plant adjust to the light intensity.
 The length of gas bubble evolved in 10 second, 30 second, and 1minute intervals is measured
by pulling the syringe plunger to draw the bubble slowly along the capillary tube.
 Steps above are repeated with the light source placed at 40 cm from the test tube with the
plant, then 30 cm, 20 cm, and finally 10 cm.
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 a control experiment is set up using natural room lighting and repeating the above steps.
Observation / results Explanation
A colorless gas which relights a glowing splint The gas is oxygen released from
evolves from the cut end of the plant. Photosynthetic reactions.
 The rate of gas evolution is directly  This is because of the increased light
proportional to light intensity up to a certain intensity which provides more energy for
illumination i.e. the closer the light source is to photo-activation of electron flow.
the plant; the more oxygen bubbles evolve up  Increased illumination may not cause any
to a certain light intensity then remains further evolution of oxygen because
relatively constant and may decrease.  of light saturation
Determination of amount of gas released a)  other factors limit the process
if scale is marked in mm3 or cm3: read volume  Increased illumination may cause a
directly decrease in bubble evolution because
b) if scale is marked in mm: calculate volume chlorophyll gets bleached with increased
from πr2h illumination.
π=3.14, r=capillary tube radius, h=distance
bubble covers

Precautions to avoid experimental

inaccuracies / errors Explanation / Remedy

 Thermostatically controlled bath should be

 Temperature fluctuation of the water in the used to maintain temperature constant since
beaker it affects photosynthetic activity.
 The experiment must be conducted in total  To avoid effects of external light fluctuations on
darkness photosynthesis
 There must be periodical refilling of HCO3-
solution  To avoid depletion of carbon dioxide

 To saturate the water with oxygen such that the

 The water should be aerated first. oxygen evolved does not dissolve into water.
 Each time the light position is adjusted, a 5-
minute lapse must be allowed before  To allow the plant equilibrate (adjust) to the new
bubble counting light intensity.

 Use voltage that gives constant light for a long

 Light intensity fluctuation time
 Trapped gas bubbles  Swirl the water weed to release them

NOTE:

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 Instead of measuring the length of bubble, bubbles can be counted, but this has several
disadvantages
(i) Some bubbles may not be seen due to variations in size, which can be avoided by adding
a little detergent to lower the surface tension
(ii) Bubbles may evolve very fast to be counted, especially in much illumination.
 The percentage of oxygen in the evolved gas is only about 40% because of dilution by
(i) dissolved N2 or other gases released from solution
(ii) CO2 which had accumulated from respiration, and is first displaced into the capillary
tubing, especially if the plant had been kept in the dark

FACTORS INFLUENCING THE RATE OF PHOTOSYNTHESIS

The rate of photosynthesis is affected by a number of factors which are both internal and external
(environmental)
Environmental factors
 Carbon dioxide concentration  Some air pollutants e.g. Sulphur
 Light intensity dioxide
 Temperature  Altitude
 oxygen concentration  Salinity
Internal factors include
 Chlorophyll concentration
 Water and dissolved nutrients
 Enzyme inhibitors e.g. cyanide, dichlorophenol dimethyl urea – DCMU
The level of each factor determines the yield of material by a plant; therefore, it is necessary to
first consider the interaction of factors controlling photosynthesis
THE PRINCIPLE OF LIMITING FACTORS
It states that:
‘At any given moment, the rate of a chemical process is limited by the one factor which is
nearest its minimum value, and by that factor alone’
A limiting factor is a factor which is nearest to its minimum value in a chemical process that is
affected by more than one factor.

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Graph illustrating the

concept of limiting factors on
the rate of photosynthesis
From the graph above, the rate
of photosynthesis increases
with increase in light intensity
and then rate remains constant
as the process reaches its
maximum rate due to;
(i) The photosynthesis
process is going at the fastest
possible pace, and no amount
of additional light will make it
go any faster.
(ii) There is insufficient
carbon dioxide available to
allow the process to speed up
any further
(iii) The temperature is too
low for the chemical reactions
to go any faster. Therefore, the
rate of photosynthesis can be
increased further by increasing either temperature of carbon dioxide concentration
which are limiting factors.

(i) Temperature
Changes in temperature have little effect on the reactions of the light-dependent stage because
these are driven by light, not heat. However, the reactions of the Calvin cycle are catalysed by
enzymes which, like all enzymes are sensitive to temperature.
Note:
 the effect of temperature on these reactions is similar to its effect on other enzymes
 The optimum temperature varies for each species, but many temperate plants have an
optimum temperature ranging from 25oC to 35oC.

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Observation / description Explanation

 Below 10 C, C3 rate of photosynthesis is  C4 photosynthetic enzymes are less active
0

higher than in C4 above 100C. in the cold but become more active with
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 450C, the rate of photosynthesis  Enzymes controlling photosynthesis are
decreases denatured by very high temperatures
 There is an initial increase in  Light intensity becomes a limiting factor in
photosynthetic rate to a maximum at each of the three cases
about 40-420C, in spite of further increase
in temperature
 There is increase in the rate of  Increase in temperature activates enzymes
photosynthesis with increase in to a level beyond which enzyme
temperature until up to at about 400C denaturation occurs.

(ii) Carbon dioxide concentration

In the atmosphere, the concentration of carbon dioxide ranges from 0.03 to 0.04 %. However, it is
found that 0.1% of carbon dioxide in the atmosphere increases the rate of photosynthesis
significantly. As long as there is no other facto limiting photosynthesis, an increase in carbon
dioxide concentration up to 0.5% usually results in an increase in the rate of photosynthesis.
However, concentrations above 0.1% can damage leaves see the graph below.

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On a warm sunny day, the concentration of carbon

dioxide in the air is probably the factor that limits
photosynthesis more than any other.
Enriching air with carbon dioxide has a significant
effect on crop plants, this is achieved in the
greenhouses which are enclosed chambers where
plants are grown under controlled conditions. Where
the concentration of carbon dioxide is increased by
installing gas burners which liberate carbon dioxide as
the gas burns.

Qn. Mention other ways the carbon dioxide

concentration of the environment can be increased

Observation / description Explanation

 the rate of photosynthesis increases  Rubisco fixes carbon dioxide instead of oxygen,
rapidly with increasing carbon because the carbon dioxide concentration is very
dioxide concentration to a maximum high out competing oxygen for occupation of
at 30 Pa in C4 plants and 90 Pa in C3 active site on RUBISCO.
plants.

 The rate of photosynthesis increases  PEPCO of C4 has a higher affinity for carbon
faster in C4 than C3. dioxide than Rubisco of C3.
 The overall photosynthetic products are  C4 needs more ATP than C3 which generally
greater in C3 than in C4 reduces photosynthetic out put

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 The C4 plants are more efficient at  At lower CO2 concentration in C3

lower CO2 concentration while C3 photorespiration reduces the photosynthesis
more efficient at higher CO2 efficiency yet C4 plants are not affected by
photorespiration as PEPCO has no affinity for
oxygen even at very low carbon dioxide
concentration.

 C3 plant has a higher compensation  PEPC has a high affinity for carbon dioxide
point than C4
 After attaining the maximum, the rate  It is because other factors limit the process e.g.
of photosynthesis remains constant in temperature, light intensity etc.
both

(iii) Light intensity

Increase in light intensity results in an increased in the rate of photosynthesis.
With a continuing increase in light intensity a point is reached where carbon dioxide is neither
evolved nor absorbed this point is the Light compensation point.
Light compensation point is the light intensity at which the photosynthetic intake of carbon
dioxide is equal to the respiratory output of carbon dioxide.
The time taken for a plant which has been in darkness to reach the compensation point is called
the compensation period. It occurs during early morning or late evenings. This varies for different
plants.
However, after reaching a certain
light intensity further increase in
light intensity has no effect on the
rate because photosynthetic pigments
have become saturated with light,
and some other factor either
availability of carbon dioxide,
amount of chlorophyll or
temperature stops the reaction from
going faster. Very high intensities
may actually damage some plants in
fact, it bleaches the chlorophyll
reducing their ability to
photosynthesize.

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SUN AND SHADE PLANTS

Sun plant are those with leaves growing on branches exposed to direct sunlight while shade
plants are those with leaves growing on branches exposed to light that has passed through leaves.
 In low light, plants need to maximise light absorption for photosynthesis to exceed
respiration if they are to survive.
 In high light environment, plants maximise their capacity for utilising abundant light
energy, while at the same time dealing with excess sunlight which can bleach chlorophyll.

• At very low light intensity, shade plants have higher

CO2 uptake; these photosynthesise best at low light
intensities which reduces with illumination.
• sun plants have a higher light compensation point
than shade plants;
• shade plants have relatively low light compensation
point this is a physiological adaptation which enables
shade plants to make efficient use of light of low
intensity.
• P represents compensation point at which CO2
uptake equals CO2 output.
• At Y biomass decreases because the rate of
respiration exceeds that of photosynthesis.

• ADAPTATIONS TO PHOTOSYNTHESISE IN SUN AND SHADEN SUN

AHADE
Adaptation: a genetically determined capability to acclimate to environmental condition.
Shade plant Sun plants

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▪ Abundant chlorophyll b (low chlorophyll a ▪ Abundant chlorophyll a (high chlorophyll

to chlorophyll b ratio) which gives leaves a to chlorophyll b ratio) to increase light
dark green colour to increase light absorption;
absorption in the dark; ▪ Palisade/ spongy mesophyll ratio high to
▪ Palisade/ spongy mesophyll ratio low to minimise light penetration;
allow maximum light penetration; ▪ Mesophyll cell surface / leaf area ratio
▪ Mesophyll cell surface / leaf area ratio low high to minimize excessive light and
to maximise light trapping; transpiration;
▪ Leaf orientation horizontal to maximise ▪ Leaf orientation erect to minimise light
light trapping; trapping;
▪ Reddish leaf undersides to enhance ▪ Stomatal density high to avoid over
reflectance back up heating;
▪ through the photosynthetic tissue; giving ▪ Much carotenoids to prevent damage to
the plant a second chance to utilize the chlorophyll from very bright light.
light. ▪ Thick leaves to minimise light
▪ Stomatal density low to avoid over penetration;
cooling; ▪ Stomatal size small to minimise water
▪ Thin leaves to maximise light penetration; loss;
▪ Stomatal size large to allow loss of excess
water; Other features
▪ Elongated internodes for increased access (i) RuBISCO and soluble protein
to light; content
▪ Chloroplast size large to increase the /mass higher
surface area for storage of photosynthetic (ii) Chlorophyll / soluble protein ratio
pigments. high
(iii) Chloroplast size small

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Exercise 4
1. The table 1 below shows the rate at which carbon dioxide is taken up (+) and released (-) from
stem of an herbaceous plant and from a single leaf of the same species at different light
intensities.
Light intensity (arbitrary UPTAKE (+) AND RELEASE (-) OF CARBONDIOXIDE
units) /mg50cm-2h-1
STEM LEAF

0.0 -0.5 -0.5

1.0 -0.2 +0.6

2.5 +0.3 +2.8

4.0 +0.7 +4.6

5.0 +1.0 +5.3

7.0 +1.6 +6.0

11.0 +2.5 +6.3

(a) Present the data provided in the table above in a suitable graph. (06 marks)
(b) Calculate the rate at which carbon dioxide is used in photosynthesis by
50cm2 of the plant organ at light intensity of 3 arbitrary units. (03 marks) (c)
Explain,
(i) the rate of uptake and release of carbon dioxide of the leaf of a plant as
light intensity increases. (14 marks)
(ii) the difference in the rate of uptake of carbon dioxide of leaf and stem of
plants. (06 marks)
(iii) From your graph, the difference in the light compensation points of the leaf
and stem of plants. (04 marks)
(d) Suggest any three practical difficulties you would meet in conducting an
experiment to obtain data of the kind given in the table. (03 marks)
(e) State physiological problems likely to be faced by a plant beyond light intensity of 3
arbitrary units. (04 marks)

(iv) Salinity
Increase in salinity brings about osmotic stress, leading to drought stress or ‘water stress’. This
results in stomata closure in an effort to avoid desiccation, which reduces photosynthesis because
uptake of CO2 reduces.

(v) Chlorophyll Concentration

The concentration of chlorophyll affects the rate of reaction as they absorb the light energy without
which the reactions cannot proceed. Lack of chlorophyll or deficiency of chlorophyll results in
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chlorosis or yellowing of leaves. It can occur due to disease, mineral deficiency or the natural
process of aging (senescence). Lack of iron, magnesium, nitrogen and light affect the formation of
chlorophyll and thereby causes chlorosis. (vi) Water
The effect of water can be understood by studying the yield of crops which is the direct result of
photosynthetic activity. It is found that even slight deficiency of water results in 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 in
order to conserve water being lost as water vapour through them. (vii) 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. In fact, 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.
(viii) altitude and oxygen

Observation / description Explanation

 The decrease in atmospheric pressure at higher
 C3 plants are more abundant at altitude decreases the partial pressure of oxygen
high altitude/elevation enables more productivity since photorespiration
reduces
 Even when temperature is high, nocturnal stomatal
opening and closure in day light enables them to
reduce transpiration.
 CAM plants are more abundant
 CAM plants that store a lot of malate and due to its
at low altitude
high osmotic value conserve a lot of water, are
usually less frost resistant than C3 plants.

 The enzymes are tolerant to these high temperatures

 C4 plants are widely distributed at and the Kranz mesophyll anatomy shields Rubisco in
low altitude and slight elevation bundle sheath cells from much oxygen to avoid
photorespiration.

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Exercise 5
1. a) Explain the significance of pigments and light in photosynthesis. (12 marks)
(b) How does altitude affect distribution of C3 and C4 plants? (08marks)

PRODUCTIVITY OF PLANTS AND PLANT COMMUNITIES

The entire plant is potential food for consumer organisms. Plants normally grow with others of the
same species or of different species in plant communities e.g. a field of wheat, natural forest and
a woodland. The efficiency with which whole plants and plant communities produce dry matter
determines how much food is available for the higher trophic levels in an ecosystem.
Factors of fundamental importance to crop yield
 Leaf area index
 Unit leaf rate
1. Leaf area index
Plants with a large surface area of leaves and other parts which can photosynthesise may be
expected to produce more dry matter than plants having shoot systems with small surface area.
The area of leaves available for photosynthesis can be expressed as the leaf area index (LAI)
𝑇𝑜𝑡𝑎𝑙 𝑙𝑒𝑎𝑓 𝑎𝑟𝑒𝑎 𝑜𝑓 𝑝𝑙𝑎𝑛𝑡
𝐿𝐴𝐼 =
𝑎𝑟𝑒𝑎 𝑜𝑓 𝑔𝑟𝑜𝑢𝑛𝑑 𝑐𝑜𝑣𝑒𝑟𝑒𝑑 𝑏𝑦 𝑝𝑙𝑎𝑛𝑡
It determines the amount of light intercepted by the shoot system of a plant.
During the early stages of growth, crop plants have small LAI values because each plant has only
a few small leaves and is surrounded by a patch of bare ground, as growth proceeds and the
shoot system enlarges, the LAI increases
Note
The shape of the shoot system is particularly important in determining the leaf area index of a plant
where plants which can be grown close to each other and which have leaves held vertically have
higher LAI than those with horizontally held or drooping leaves.
2. Unit leaf rate
Whatever the LAI value, increases in organic matter occur efficiently only if most of the
photosynthetic products are converted to plant tissue or storage materials. If most of the products
of photosynthesis are respired dry matter accumulates slowly.
Unit leaf rate expresses the efficiency of dry matter accumulation by green plants.
ULR of a plant can be calculated from measurements of the leaf area and dry mass of a
representative sample of plants at different stages of growth.
Note
 Some species have higher ULR values than others because they do not photorespire and
have very short compensation periods. E.g. C4 plants such as sugar cane and maize have a
much grater unit leaf rate than most C3 plants.
Synthesis of dry matter by green plants is called primary production, the total amount of dry matter
produced per unit area of ground per year is called gross primary productivity. Some dry matter is
used by green plants in respiration. What is left is called net primary productivity (NPP) and it is
which is available for consumer organisms including man NPP = LAI x ULR
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Therefore, plants which quickly achieve high LAI values and which sustain an efficient ULR
over a long growing period are highly productive.
Exercise 6
1. The figure below shows the changes in leaf area index (ratio of leaf surface to soil surface
(m2cm-2) of two species of clover, Triforium ripens and Triforium fragiferum, growing in a pure
and mixed stand.

Table 1: shows the characteristics of the petioles and leaf size of the two species of clover.

Characteristics
T. fragiferum T.repens
Petiole length Long Short
Leaf size Large Small

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Use
( the information in the figure and table to answer the questions that follow,
a) Compare the leaf area index of Trifolium repens and T.fragiferum in the,
( (i) pure stands.(06 marks) (ii) mixed stands. (06 marks)
b) Explain the trend in leaf area index for Trifolium repens in pure stands.
( (10 marks)
c) Explain the differences in growth rate of the two species in mixed stands.
( (07 marks)
d) Explain why Trifolium fragiferum continues to grow after the peak of Trifolium
( repens?
(f (04 marks)
e) What conclusion can you draw from the results in a mixed stand? (04 marks)
) What other factors are likely to have caused the difference in growth rate of the two
species in mixed stand? (03 marks)

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