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Organisms that synthesise their organic

substances especially carbohydrates from
carbon dioxide and water with energy
source in the form of light obtained by
photosynthetic pigments.
Example : plants, algae, cynobacteria
Green sulphur bacteria (Chlorobium)
Purple sulphur bacteria (Chromatium)
H2S + 6CO2
C6H12O6 + 12S + 6H2O

What is Photosynthesis?

Photosynthesis the process by which plants,

some bacteria, and some protistans use the
energy from sunlight to produce sugar, which
cellular respiration converts into ATP, the
"fuel" used by all living things.
The conversion of unusable sunlight energy
into usable chemical energy, is associated
with the actions of the green pigment
Most of the time, the photosynthetic process
uses water and release the oxygen that we
absolutely must have to stay alive.

We can write the overall reaction of this

process as:
6H2O + 6CO2 ----------> C6H12O6+ 6O2
Most of us don't speak chemicalese, so the
above chemical equation translates as:
six molecules of water plus six molecules
of carbon dioxide produce one molecule
of sugar plus six molecules of oxygen

Figure 4.1 Diagram of a typical plant, showing the inputs and

outputs of the photosynthetic process.

Iron bacteria
Iron II to Iron III
Colourless sulphur bacteria
Sulphur to sulphuric acid
Nitrifying bacteria
Ammonia / ammonium salts to nitrite
Nitrite to nitrate

Photosynthetic Pigments
Photosynthetic Pigments


Chorophyll a


Chorophyll b






Phaeophytin a


Phaephytin b


Chlorophyll a

Most abundant photosynthetic pigment.

Absorb light mainly in blue-violet (430nm)
and red (662nm) region.
Has porphyrin ring with a Mg atom at its
Contains a hydrocarbon tail which interacts
with the hydrophobic regions of proteins in
the thylakoid membrane.

Figure 4.9 The molecular structure of chlorophylls.

Figure 4.8a Molecular model of chlorophyll.

Figure 4.8b Molecular model of carotene.


A cluster of photosynthetic pigments (200300 molecule of chorophyll) grouped together

with a primary electron accepter and a series
of electron carriers.
Function : to capture light energy for
When a photon of light energy strikes a
pigment molecule, the energy passed from
one pigment molecule to another until it
reaches a particular chlorophyll molecule
(P700 or P680) at the reaction centre where it
can release electron.

Photosystem I

System of pigments in smaller particles

that usually found in larger numbers in the
intergranal lamella than the thylakoid
The wavelength of light most strongly
absorbed is 700nm.
Has three kinds of pigments : chlorophyll a,
chlorophyll b and carotenoids.
Chlorophyll a is more abundant

Photosystem I

Photosystem II

cluster of pigments in the form of bigger

particles that usually found in the thylakoid
membrane of the grana.
The wavelength of light most strongly
absorbed is 680nm.
Has three kinds of pigments : chlorophyll a,
chlorophyll b (or c or d, depending on the
species of plant) and carotenoids.
Chlorophyll b (or c or d) is more abundant
than chlorophyll a

Photosystem II

The Nature of Light

White light is separated into the different

colors (=wavelengths) of light by passing it
through a prism. Wavelength is defined as
the distance from peak to peak (or trough to
The energy of is inversely proportional to the
wavelength: longer wavelengths have less
energy than do shorter ones.

Figure 4.5 Wavelength and other suspects of the wave nature of light.

Figure 4.6 The electromagnetic spectrum.

The order of colors is determined by the

wavelength of light.
Visible light is one small part of the
electromagnetic spectrum.
The longer the wavelength of visible light, the
more red the color.
Likewise the shorter wavelengths are towards
the violet side of the spectrum.
Wavelengths longer than red are referred to
as infrared, while those shorter than violet are

Figure 4.7 Absorption and transmission of different

wavelengths of light by a hypothetical pigment.

Absorption Spectrum

Obtained by using a spectrophotometer.

When white light is dispersed using a prism to
produce a visible light spectrum of seven
colours ranging from violet to red.
The different wavelength of colour light are
illuminated to a test tube containing
chlorophyll extract one at a time.
The galvanometer shows the transmittance of
the different wavelengths of color light which
measures the extent in which a specific
wavelength of colour light is absorbed.

Action Spectrum

The action spectrum is a graph that shows the

effectiveness of different wavelength of light
involved in photosynthesis.
Is obtained by using the pure pigment or mixture
mixed with an electron acceptor.
Then it is exposed to different wavelength of
The time taken for a standard concentrations of
pigment and indicator solution to turn colourless
is taken as the rate of photosynthesis.
The amount of oxygen given out can also used
to determine the rate of photosynthesis.

Figure 4.10 Absorption spectrum of several plant pigments (left) and action
spectrum of elodea (right), a common aquarium plant used in
lab experiments about photosynthesis.

Stages of Photosynthesis

Photosynthesis is a two stage process.

The first process is the Light Dependent
Process (Light Reaction), requires the direct
energy of light to make energy carrier
molecules that are used in the second
The Light Independent Process (or Dark
Reaction) occurs when the products of the
Light Reaction are used to form C-C covalent
bonds of carbohydrates.

The Dark Reactions can usually occur in the

dark, if the energy carriers from the light
process are present.
The Light Reactions occur in the grana and
the Dark Reactions take place in the stroma
of the chloroplasts.

Figure 4.12

Figure 4.12 Overview of the two steps in the photosynthesis process


Light Reactions

In the Light Dependent Processes (Light

Reactions) light strikes chlorophyll a in such a
way as to excite electrons to a higher energy
In a series of reactions the energy is
converted (along an electron transport
process) into ATP and NADPH.
Water is split in the process, releasing
oxygen as a by-product of the reaction.

The ATP and NADPH are used to make C-C

bonds in the Light Independent Process
(Dark Reactions).
In the Light Independent Process, carbon
dioxide from the atmosphere (or water for
aquatic/marine organisms) is captured and
modified by the addition of Hydrogen to form
carbohydrates (general formula of
carbohydrates is [CH2O]n).
The incorporation of carbon dioxide into
organic compounds is known as carbon

The energy for this comes from the first

phase of the photosynthetic process.
Living systems cannot directly utilize light
energy, but can, through a complicated series
of reactions, convert it into C-C bond energy
that can be released by glycolysis and other
metabolic processes.
Photosystem are arrangements of chlorophyll
and other pigments packed into thylakoids.
Many Prokaryotes have only one
photosystem, Photosystem II (so numbered
because, while it was most likely the first to
evolve, it was the second one discovered).

Eukaryotes have Photosystem II plus

Photosystem I.
Photosystem I uses chlorophyll a, in the form
referred to as P700.
Photosystem II uses a form of chlorophyll b
known as P680.
Both "active" forms of chlorophyll a function
in photosynthesis due to their association
with proteins in the thylakoid membrane.

Figure 4.13

Figure 4.13 Action of a photosystem


Photophosphorylation is the process of

converting energy from a light-excited
electron into the phosphate bond of an ADP
This occurs when the electrons from water
are excited by the light in the presence of
The energy transfer is similar to the
chemiosmotic electron transport occurring in
the mitochondria.

Light energy causes the removal of an

electron from a molecule of P680 that is part
of Photosystem II.
The P680 requires an electron, which is
taken from a water molecule, breaking the
water into H+ ions and O-2 ions.
These O-2 ions combine to form the diatomic
O2 that is released.


4e + 4H+ +O2

The electron is "boosted" to a higher energy

state and attached to a primary electron
acceptor, which begins a series of redox
reactions, passing the electron through a
series of electron carriers, eventually
attaching it to a molecule in Photosystem I.
Light acts on a molecule of P700 in
Photosystem I, causing an electron to be
"boosted" to a still higher potential.

The electron is attached to a different primary

electron acceptor (that is a different molecule
from the one associated with Photosystem II).
The electron is passed again through a
series of redox reactions, eventually being
attached to NADP+ and H+ to form NADPH,
an energy carrier needed in the Light
Independent Reaction.
The electron from Photosystem II replaces
the excited electron in the P700 molecule.
There is thus a continuous flow of electrons
from water to NADPH.

This energy is used in Carbon Fixation.

Cyclic Electron Flow occurs in some
eukaryotes and primitive photosynthetic
No NADPH is produced, only ATP.
This occurs when cells may require additional
ATP, or when there is no NADP+ to reduce to
In Photosystem II, the pumping to H ions into
the thylakoid and the conversion of ADP + P
into ATP is driven by electron gradients
established in the thylakoid membrane.
Figure 4.14

Figure 4.14 Noncyclic photophosphorylation (top) and cyclic

photophosphorylation (bottom). These processes are better
known as the light reactions.


Photophosporylation in the chloroplast.

The thylakoid space contains high proton (H+)
concentration as a result of proton pump and
water is also split to form it.
When the proton passes down the concentration
gradient to the stroma through a channel protein
of enzyme ATP synthase, ATP is formed from
ADP and phosphate.
Further increase in proton concentration
gradient is from proton pump and is used in
formation of NADPH outside the thylakoid

After the light dependent stage, ATP and

NADPH are formed together with oxygen that
is released from leaves.
The ATP and NADPH2 will be used for the
light independent stage.

Dark Reaction

Carbon-Fixing Reactions are also known as

the Dark Reactions (or Light Independent
Carbon dioxide enters single-celled and
aquatic autotrophy through no specialized
structures, diffusing into the cells.
Land plants must guard against drying out
(desiccation) and so have evolved
specialized structures known as stomata to
allow gas to enter and leave the leaf.

The Calvin Cycle occurs in the stroma of

Carbon dioxide is captured by the chemical
ribulose biphosphate (RuBP).
RuBP is a 5-C chemical.
RuBP accept CO2 form a temporary sixcarbon molecules, which is unstable and
immediately breaks down to form two
phosphoglycerate (PGA) [3C chemical].

Figure 4.16 The first steps in the Calvin cycle


The first stable product of the Calvin Cycle is

3-phosphoglycerate (PGA), a 3-C chemical.
The energy from ATP and NADPH energy
carriers generated by the photosystems is
used to attach phosphates to (phosphorylate)
the PGA.

Eventually there are 12 molecules of

glyceraldehyde-3-phosphate (also known as
phosphoglyceraldehyde or PGAL, a 3-C), two
of which are removed from the cycle to make
a glucose.
The remaining PGAL molecules are
converted by ATP energy to reform 6 RuBP
molecules, and thus start the cycle again.

Metabolism of Glycerate Phosphate

( GP / PGA ) and Glyceraldehyde
Phosphate ( TP / PGAL )
Synthesis of carbohydrates
Glyceraldehyde-3-phosphate molecules are
converted to form monosaccharides eg.
Glucose and fructose combine to form
sucrose, transported in phloem sieve tubes.
The glucose molecules can be polymerized
into starch for storage or cellulose,
component of plant cell walls.

Synthesis of lipids

3-phosphoglycerate enters glycolysis

pathway and is converted to pyruvate.
Pyruvate is converted to 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
3-phosphoglycerate can also be converted to
Lipids such as triglycerides are esters of fatty
acids and glycerol. They are important
components of cell membranes.

Synthesis of proteins

3-phosphoglycerate is converted into acetyl

coenzyme A and enters into the Krebs cycle.
Some of its intermediates can produce
different amino acids by transamination
Amino acids are then polymerized into
proteins which are required for growth and
development , synthesis of enzymes and
structural components of the cell.

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.


Fixation of CO2 replaced by fixation of O2

When the [ ] of oxygen is high, oxygen will
act as a competitive inhibitor to compete
with CO2 for the active site of the enzyme
Photorespiration occurs when stomata
CO2 cant diffuse into the leaf & O2 is released
from FTS.

2 phosphoglycolate (2C)
GP (3C) + C


2 Phosphoglucolate are converted to 1

molecule of 3-phosphoglycerate requiring
O2 is used and the fourth organic carbon is
converted into CO2

There is no net production of ATP, NADPH

and other energy rich metabolites.
Prevention of photorespiration is to increase
the [ ] of CO2
i) C4 plants, the Hatch-slack pathway
ii) CAM plants close the stomata during the
day & fixation of CO2 occurs at night.

Figure 4.18 Photorespiration


C4 Leaf Compared with C3 leaf

1. Example of leaf C3 plant is Helianthus (sunflower).

Figure 4.19 Leaf anatomy of a C3 (top) and C4 (bottom) plant.

2. Kranz anatomy is exhibited in C4 plants.

Kranz (=crown) anatomy is the internal
structure of Grammineae leaves, which have
a ring of bundle sheath cells surrounding the
vascular tissue. Leaves with Kranz anatomy
exhibit dimorphic chloroplasts, i.e. two
different types each in mesophyll cells and
bundle sheath cells.

3. Differences between the chloroplasts found in that of the

mesophyll and that of bundle sheath are summarised in
the table below.
Table 4.3

Chloroplasts of

Chloroplasts of bundle

i) Granal activities

They are higher

They have none or little


ii) Photosystem II

It is active, produces
plenty of ATP, NADPH2
and O2

It is little , produces little

NADPH2 and O2

iii) RuBP carboxylase

It has almost none

It is of high

iv) CO2 fixation

It is nil

It is active

v) Starch

There is little starch

There is abundant of

C4 Plant

C-4 Pathway / Hatch-slack


Some plants have developed a preliminary

step to the Calvin Cycle (which is also
referred to as a C-3 pathway), this preamble
step is known as C-4.
While most C-fixation begins with RuBP, C-4
begins with a new molecule,
phosphoenolpyruvate (PEP), a 3-C chemical
that is converted into oxaloacetic acid (OAA,
a 4-C chemical) when carbon dioxide is
combined with PEP.

The OAA is converted to Malic Acid and then

transported from the mesophyll cell into the
bundle-sheath cell, where OAA is broken
down into PEP plus carbon dioxide.
The carbon dioxide then enters the Calvin
Cycle, with PEP returning to the mesophyll
The resulting sugars are now adjacent to the
leaf veins and can readily be transported
throughout the plant.

Figure 4.17 C-4 photosynthesis involves the separation of carbon

fixation and carbohydrate synthesis in space and time.

Crassulacean acid metabolism ( CAM ) plants

CAM plants only contain mesophyll cells and
no Krantz anatomy.
Photosynthetic pathway is the same as C4
plants but fixation of carbon dioxide takes
place at night when stomata are open.
Phosphoenolpyruvate combines with carbon
dioxide to reduce oxaloacetate. Oxaloacetate
is reduced to form malate. Malate is stored in
the cell vacuole at night to prevent pH
changes in the cytoplasm.
During day time, malate is oxidized producing
pyruvate and carbon dioxide.

Concentration of carbon dioxide increases in

the mesophyll cells and photorespiration is
Carbon dioxide is used in Calvin cycle
producing organic molecules.
Stomata of CAM plants are closed during
daytime. They are open at night when lower
temperatures and higher humidities reduce
the rate of transpiration.

The pathway was first discovered in a plant

belonging to the family Crassulaceae and is
therefore referred to as crassulacean acid
metabolism ( CAM ).
The same pathway has been found in some
other plants belonging to different families
also living in hot, arid conditions. Eg of CAM
plants are succulent plants such as the cacti
and pineapples.

Difference between Hatch-Slack pathway

( C4 Cycle ) and CAM



1) Family

It occurs in Grammineae

It occurs in Crussulaceae

2) Example

An example is maize

Cacti and pineapple

3) Cells involved

Two types i.e. mesophyll and

bundle sheath cells are involved

One type i.e. mesophyll

cells are involved only

4) Chloroplast involved

There are two types involved

One type is involved

5) Separation of

It is a spatial separation, by

It is a temporal separation,
by time

6) Regeneration of PEP

It does not involve starch

It involves starch

The capture of carbon dioxide by PEP is

mediated by the enzyme PEP carboxylase,
which has a stronger affinity for carbon
dioxide than does RuBP carboxylase When
carbon dioxide levels decline below the
threshold for RuBP carboxylase, RuBP is
catalyzed with oxygen instead of carbon

The product of that reaction forms glycolic

acid, a chemical that can be broken down by
photorespiration, producing neither NADPH
nor ATP, in effect dismantling the Calvin
C-4 plants, which often grow close together,
have had to adjust to decreased levels of
carbon dioxide by artificially raising the
carbon dioxide concentration in certain cells
to prevent photorespiration.

C-4 plants evolved in the tropics and are

adapted to higher temperatures than are the
C-3 plants found at higher latitudes.
Common C-4 plants include crabgrass, corn,
and sugar cane.
Note that OAA and Malic Acid also have
functions in other processes, thus the
chemicals would have been present in all
plants, leading scientists to hypothesize that
C-4 mechanisms evolved several times
independently in response to a similar
environmental condition, a type of evolution
known as convergent evolution.

4. The physiological differences between C3 and C4 plants

Table 4.4

In C3 leaf

In C4 leaf

a) CO2 fixation

It has one type, C3 cycle

It has two types, C4 and C3


b) First product of photosynthesis

It is PGA

It is oxaloacetate

c) O2 inhibition

It has a high O2 inhibition

It has low O2 inhibition

d) Photorespiration

It has a higher rate of


It has low rate of


e) Maximum rate of

This can be achieved with

lower light intensity

A higher maximum is obtained

with higher light intensity

f) Effect of temperature
raised from 25C to 35C

There is no change at higher


There is a 50% higher rate at

higher temperature

g) Compensation point

It requires 40-60ppm of CO2 to

achieve this

It just requires a few ppm to

achieve this

h) water loss

A lot of water is lost during


Little water is lost during


i) Efficiency

It is less efficient

It is more efficient

Limiting Factors on the Rate of

Wavelenght of light (light quality) can influence
the rate of photosynthesis.

Visible light of different wavelengths or

different colours affect the rate of
photosynthesis differently.

Red light of around 450 nm and blue light of

680 nm are most effective for
photosynthesis whereas green light of 540
nm is least effective.

This is because red and blue lights are

absorbed by the chlorophyll whereas green
light is not absorbed at all.
The effects of different wavelengths of light
on photosynthesis can be seen from the
action spectrum below.
The action spectrum is corresponding to the
absorption spectrum as shown.
This shows that light absorbed by
chlorophyll is used for photosynthesis.

Absorption spectrum
Rate of


% of light
absorbed by





Wavelength / nm

Rate of Photosynthesis

Light intensity or quality of light affects the rate

of photosynthesis.

The effect of light intensity on the rate of

photosynthesis is as shown.

Light Intensity / lux

When light intensity is low, the rate of

photosynthesis is proportional to light
intensity. This is because the light supplies
energy for photosynthesis.
At a higher light intensity depending of on
the type of plants, the rate of photosynthesis
becomes maximum due to atmospheric
carbon dioxide is always a limiting factor at
0.04 %.
If the carbon dioxide concentration is
increased to 0.4%, the rate of
photosynthesis will be increased a few
times as shown.

Rate of Photosynthesis

0.4%CO2CH 25oC

0.04%CO2CH 25oC

Light intensity / lux

If the light intensity is further increased, the

rate of photosynthesis will fall rapidly
because the temperature may be too high
that denatures enzymes or the chlorophyll is
destroyed by photo-oxidation.

Rate of photosynthesis

Temperature is an important factor in


The effect of different temperatures on the rate of

photosynthesis is as shown.







At 0C, photosynthesis is very low because

there is very little kinetic energy for any
enzymic reaction to take place.
At optimum temperature, which may be
between 25C to 40C, the rate of
photosynthesis is maximum. This is
because such temperature provides the
most suitable amount of kinetic energy for

Between 0C to optimum temperature, the

temperature quotient ( Q10 ) is equal to 2.
This is same as to say for every 10C
increased in temperature, there is a
corresponding double increase in the rate of
When the temperature is increased further,
there is a rapid decrease in the rate of
photosynthesis. This is because enzymes are
denatured by high temperature.

Rate of Photosynthesis

Carbon dioxide concentration is always a

limiting factor

The effect of carbon dioxide concentration on the

rate of photosynthesis is shown .





CO2 concentration / %

When the carbon dioxide concentration is

increased from 0 to 0.1%, the rate of
photosynthesis increases proportionally.
This is because carbon dioxide is a reactant
for the process.
0.1% concentration of carbon dioxide
concentration is the optimum concentration.
If the concentration is increased from 0.1 to
1%, the rate of photosynthesis remains
more or less the same as other factor may
be limiting.

Further increase in concentration of carbon

dioxide will result in a rapid decrease in the
rate of photosynthesis. This is because
when too much carbon dioxide dissolved in
water, it produces too low a pH that would
denature enzymes.

Water Availability

Water is required in the light-dependent

reactions of photosynthesis.

However it is also required in many other

plant physiological processes. It is difficult to
measure its direct effect on photosynthesis.

The closure of stomatal pores of wilted

leaves may decrease the rate of
photosynthesis by preventing the diffusion
of carbon dioxide to the mesophyll tissue.

Chlorophyll concentration

under normal conditions, chlorophyll

concentration does not limit photosynthesis.

In the absence of ions like magnesium and

nitrogen which are essential in chlorophyll
synthesis, the chlorophyll concentration is

Chlorophyll pigments may also be

destroyed by very high light intensity
resulting in reduced rate of photosynthesis.

Oxygen concentration

oxygen acts as a competitive inhibitor and

competes with carbon dioxide for the active
site of the enzyme ribulose bisphosphate

High oxygen concentration and low carbon

dioxide concentration will lower the rate of
photosynthesis in C3 plants

Compensation point
Compensation point is a state in plants when
the rate of photosynthesis is same as the rate
of respiration.
This happens under one or all the following
This is usually occurs in the morning or evening or
whenever when the light intensity is low.
It can be artificially induced when the carbon
dioxide concentration is reduced at certain light
This occurs when temperature is artificially
reduced at certain light intensity.

It has the following characteristics:

at compensation point, the carbon dioxide

produced by respiration is used for
the oxygen produced in photosynthesis is
used for respiration
there is no net production of sugar as
what is produced by photosynthesis is
used up by respiration.
There is no net gaseous exchange.
Different species of plants have different
compensation points partly due to different
chlorophyll respiration.

Plants exhibiting compensation point a

lower light intensity shows more
efficiency in photosynthesis than plants
that need higher light intensity.
Similarly, plants are more efficient in
photosynthesis if they need low carbon
dioxide concentration and temperature
to achieve compensation point.

Sun-loving plants

Adapted for growth in areas that receives

strong sunlight and need more energy to
form and maintain thick leaves.
Have a higher rate of respiration and reach
the compensation point at high light
Have more palisade cells in the leaves,
which allow more light to be absorbed so that
more carbohydrate can be produced.

Shaded plants

Adapted for growth in shady areas with

fewer cells and less chlorophyll.
Less energy is needed for leaves formation
and maintenance.
Have a lower rate of respiration and reach
the compensation point faster at low light
Less carbohydrate can be produced.

Roles of
C3, C4 and CAM plants
on the increasing carbon
dioxide emission and
global warming

C3 plants

Most plants in the world are C3 plants with

RuBP carboxylase in the stroma of
chloroplasts to fix carbon dioxide.
C3 plants play a very important role to reduce
increasing carbon dioxide emission and global
They increase in the rate of carbon fixation
when carbon dioxide [ ] increase.
Shows higher photosynthetic rate as [ ] of CO2

C4 plants

C4 plants are grass family with PEP

carboxylase to fix carbon dioxide in the
cytoplasm of mesophyll cells.
The role of C4 plants is not important as C3
PEP carboxylase work at a high rate when the
[ ] of CO2 is low.
C4 plants helps to increase food production at
higher warm temperature.

CAM plants

CAM plants open their stomata at night to fix

carbon dioxide.
Close their stomata to reduce transpiration
during day time.
Prevent photorespiration.

Global warming

is linked to an increase in the CO2 [ ].

Result in rising sea levels, increase in the
melting of ice, changes in vegetation and
unusual weather patterns.
As the sea level increase,
a) C3 plants help to reduce CO2 [ ]
b) C4 plants adapted in increasing salt [ ] &
important to produce more food.