Unit 5

Energy transformation
5.1 Respiration
Minimum Competency
By the end of this section you should be able to:

 Describe the structure of ATP and its role in cellular metabolism.

 Explain how ATP is adapted to its role as an energy transfer molecule within a
cell.
 Describe how ATP is produced in a cell.
 Locate where the different processes of cellular respiration occur in the cell.
 Explain the role of electron donors and acceptors.
 Describe in detail each stage of aerobic respiration. Draw and label the structure
of a mitochondrion.
 Explain the processes of alcoholic fermentation and lactate production.
 Appreciate the importance of lactate production during running and other sports.

 Continued….
What is the ATP molecule like?
The full name for ATP is Adenosine Tri-Phosphate and it is clear that
ATP contains Adenosine (nucleic acid) and it is apparent that all
nucleic acids are composed of nucleotides, which in tern contains:
• a nitrogenous base (this one contains adenine)
• a pentose sugar
• a phosphate group
 That is why sometimes ATP is described as a phosphorylated
nucleotide. i.e. ATP is essentially the adenine nucleotide
with two extra phosphate groups added on – making
three in all.
 Continued….

Adding the extra phosphates requires energy, particularly when the

third phosphate is added. As a result, energy is stored in the ATP
molecule and when the bonds that hold this third phosphate in place
are broken, the energy is released again. When the third phosphate is
removed from ATP, we are still left with a phosphorylated nucleotide,
but this one only has two nucleotides. It is Adenosine Di-Phosphate
– or ADP.
 Continued….
How is ATP adapted to its role as an energy
transfer molecule in cells?
First, we must explain what we mean by an energy transfer
molecule. Sunlight energy cannot be used directly by plants (and
certainly not by other organisms) to ‘drive’ the synthesis of
proteins– or any other molecules. The same applies to the energy
held in a glucose molecule. These two energy sources must be
used to produce ATP, which is used to transfer the energy to the
relevant cellular process.
 Therefore, ATP is our energy currency and that is why all
living cells respire all the time to produce the ATP they
need. There are no exceptions.
 Continued….

ATP is adapted to this role because it:

releases energy in relatively small amounts that are
closely matched to the amounts of energy required in
many biological processes occurring inside cells.
releases energy in a single-step hydrolysis reaction,
so the energy can be released quickly.
is able to move around the cell easily, but cannot
escape from the cell
 Continued….

The following processes are examples of

processes that require energy from ATP:
the synthesis of macromolecules – such as proteins
active transport across a plasma membrane.
muscle contraction
conduction of nerve impulses
the initial reactions of respiration (the later reactions
release energy from glucose to form more ATP)
 Continued…..
How is ATP produced in a cell?
Almost all the ATP produced in cells is formed in the same
way.
It obviously involves ADP and Pi joining to form ATP and
this requires an input of energy. What we need to look
at is just how it is made to happen.
The formation of ATP involves an enzyme called ATP
synthase.

The ATP synthase in this diagram is in one of the

membranes of a mitochondrion, but it could be in a
 Continued….
o To understand how it works, you should think of it as a
kind of molecular ‘water wheel’. When the rotor is
made to spin by hydro- gen ions passing through it, the
energy of the spinning is used to activate sites in the
catalytic knob that convert ADP and Pi to ATP.
o In both photosynthesis and aerobic respiration, many
of the reactions generate the hydrogen ions that will
pass through the ATP synthase to produce ATP.
NOTE: About the ADP/ATP inter-conversion
More than one eminent biologist has said ‘the whole
biological world turns on the coupling and
uncoupling of the third phosphate of ATP’. This is
because this process is virtually the only way in which
energy can be harnessed and then released to drive
metabolic processes in cells.
 Continued….

How is ATP produced in respiration?

There are two main pathways by which respiration can produce
ATP:
the aerobic pathway (aerobic respiration) – this requires
the presence of oxygen, and
the anaerobic pathway (anaerobic respiration and
fermentation)– this can take place in the absence of oxygen.
 Continued….
How is ATP produced in aerobic respiration?
A small amount of ATP is produced in a way that does not involve the ATP
synthase molecule; this method is called substrate level
phosphorylation. In this process, another molecule such as phosphoenol
pyruvate (the substrate) is able to transfer a phosphate group directly to
ADP.
NOTE:
 substrate level phosphorylation is when another molecule (substrate) is
able to transfer a phosphate group directly to ADP.
 The process is still catalyzed by an enzyme, it is just not ATP synthase .
 Continued….
o As already mentioned, this process only produces a
relatively small amount of the ATP produced in aerobic
respiration – in fact it produces about 10% of the total
ATP produced in aerobic respiration.
o As about 90% of the ATP produced in aerobic
respiration is produced by ATP synthase, many of the
reactions of this process are geared to producing the
hydrogen ions that will spin the rotor of the ATP
synthase molecule.
o Many different organic molecules can be respired –
they are called respiratory substrates. However,
glucose is the most commonly respired substrate and
so we will begin by looking at how this molecule is
respired.
 Continued

How are hydrogen ions transferred from glucose

to ATP synthase?
Two molecules are important in this transfer process:
Nicotinamide Adenine Dinucleotide (NAD).
Flavine Adenine Dinucleotide (FAD).
o Both are coenzymes and are capable of accepting
hydrogen ions. When this happens, we say that the
molecules have been reduced. We write the
reduced forms of the molecules as NADH and FADH or
NAD(reduced) and FAD(reduced).
o These molecules can release their hydrogen ions and
become oxidized again. The hydrogen ions can then
be used to turn the rotor of ATP synthase.
 Continued….

What are the stages of aerobic respiration of

glucose?
There are four stages in the aerobic respiration of glucose. These are:
glycolysis
the link reaction
Krebs cycle
electron transport and chemiosmosis

 Glycolysis
Glycolysis, takes place in the cytoplasm. It does not take place inside
the mitochondria because:
• the glucose molecule cannot diffuse through the mitochondrial
membranes (it is a medium-sized molecule and is not lipid soluble), and
• there are no carrier proteins to transport the glucose molecule across
the membranes.
Glycolysis (literally ‘glucose splitting’) results in glucose being
converted into a smaller molecule containing only three carbon atoms –
pyruvate. Pyruvate can enter the mitochondria and so all the
other stages take place inside the mitochondrion.
 Continued….

In the link reaction, pyruvate is then converted into a

two-carbon compound that enters into a cycle of
reactions – the Krebs cycle (named after Sir Hans
Krebs who discovered the reactions involved). Both
these stages take place in the fluid matrix of a
mitochondrion
 Continued….

In all three stages (glycolysis, the link reaction and

Krebs cycle), hydrogen atoms are transferred to NAD
to produce reduced NAD (NADH).
The Krebs cycle also produces reduced FAD (FADH).
These molecules later release their hydrogen atoms
as protons (hydrogen ions) and electrons in the final
stage of aerobic respiration. The electrons pass along
a series of molecules called an electron transport
chain.
NOTE: Electron transport chain a series of
molecules along which electrons travel.
The protons are used in the chemiosmotic synthesis of
ATP as they spin the rotor of the ATP synthase enzyme
located in the inner membrane of the mitochondrion
(Cristae). Eventually, the protons (hydrogen ions)
and electrons will combine with oxygen to form
 Continued…..
 Without the oxygen, this cannot happen as there is
nothing at the end of the electron transport chain to
accept the electrons. The electron transport chain
grinds to a halt and so does the production of ATP by
ATP synthase. Because it is oxygen-dependent, this
method of ATP production is called oxidative
phosphorylation.
 The link reaction, Krebs cycle and the reactions of the
electron transport chain all depend on the presence of
oxygen. None of these occurs in anaerobic respiration.
Glycolysis can take place in the absence of oxygen
and is the only energy-releasing process in anaerobic
respiration.
 Continued…..
What happens in glycolysis?
The reactions of glycolysis take place in the cytoplasm.
The following reactions take place in glycolysis:
o two molecules of ATP are used to ‘phosphorylate’
each molecule of glucose. This makes the glucose
more reactive.
o in the phosphorylation process, it is converted to
another six- carbon sugar (fructose 1,6-bisphosphate).
o the fructose 1,6-bisphosphate is split into two
molecules of the three-carbon sugar glyceraldehyde-3
phosphate (GP).
o each molecule of GP is then converted into pyruvate,
with the production of two molecules of ATP (by
substrate level phosphorylation) and one molecule of
reduced NAD.
Continued….
 Continued….

What are the end products of glycolysis?

 There is a net gain of two ATP molecules per molecule of
glucose (two molecules are used initially and then four
are produced).
 Two molecules of reduced NAD are also produced (per
molecule of glucose).
 Two molecules of pyruvate are also produced. The
molecules of pyruvate pass into the mitochondria
through carrier molecules in the mitochondrial
membrane.
 Continued….
 The link reaction
o Location: It take place in the fluid matrix of
the mitochondrion.
o In the link reaction, a molecule of pyruvate reacts
with a molecule of coenzyme A (CoA) to form a
molecule of acetyl coenzyme A (acetyl CoA). In the
reaction:
 hydrogen is lost and reduced NAD is formed; removing
hydrogen from a molecule is dehydrogenation.
 a carbon atom is lost to form carbon dioxide; removing
carbon from a molecule is decarboxylation.
 Continued….

The acetyl coenzyme A then reacts with a C4 molecule

(a molecule containing four carbon atoms) called
oxaloacetate. In the reaction, acetyl CoA breaks down
into:
 a two-carbon ‘acetyl’ group, which reacts with the C4
compound oxaloacetate to form a C6 compound, and
 the original coenzyme A molecule, which is reused in
further reactions with other molecules of pyruvate.
 Products of Link reaction
o Acetyl * 2
o NADH* 2
o CO2 * 2
 Continued….
What happens in the Krebs cycle?
 the two-carbon group from acetyl coenzyme A reacts
with the four-carbon compound oxaloacetate to form a
six-carbon compound called citrate.
 citrate then loses a carbon atom (is decarboxylated)
to form a five-carbon compound and CO2 is produced.
 the five-carbon compound is then further
decarboxylated to form a four-carbon compound and
CO2 is again produced; a molecule of ATP is also
produced by substrate level phosphorylation.
 the four-carbon compound undergoes several
molecular transformations to regenerate the original
four-carbon compound (oxaloacetate) and the cycle is
complete and can begin again with oxaloacetate
reacting with another molecule of acetyl CoA.
Continued…..
 Continued….

End products of the Krebs cycle are:

 2CO2 * 2= 4
 3NADH * 2 = 6
 1FADH * 2= 2
 1 ATP * 2 = 2
 Continued….
 What happens in the electron transport chain and
chemiosmosis?
The electron transport chain and chemiosmosis together
make up the process of oxidative phosphorylation.
Whereas the reactions of the link reaction and Krebs
cycle take place in the fluid matrix of the mitochondrion,
the reactions of the electron transport chain and
chemiosmosis take place on the inner mitochondrial
membrane (Cristae).

.
 Continued…..

On the cristae, the following events take place:

the hydrogen atoms carried by reduced NAD and
reduced FAD are released and split into protons
(hydrogen ions) and electrons.
the electrons pass along a series of electron carriers
that form the transport chain; they lose energy as
they pass from one carrier to the next.
three of the electron carriers are proton pumps that
move protons from the matrix of the mitochondrion to
the inter-membrane space.
as the electrons are transferred through these three
proton pumps, the energy they lose powers the
pumps which move the protons into the inter-
membrane space.
electrons from reduced NAD make this happen at all
 Continued….

The molecules that act as electron carriers in the

electron transport chain are:
reduced NAD dehydrogenase (also a proton pump)
ubiquinone (also a proton pump), and
a number of carriers called cytochromes (these are
proteins that contain iron); two of them form a
complex that acts as the third proton pump.
Continued…..
 Continued….
o At the end of the electron transport chain, the electrons combine
with protons and with oxygen to form molecules of water. Because
of this, oxygen is known as the terminal electron acceptor.

o Whereas reduced NAD is dehydrogenated by the NAD

dehydrogenase complex, reduced FAD is dehydrogenated by
ubiquinone. So electrons from reduced FAD only operate two
of the three proton pumps.

o Because of the action of the proton pumps, protons accumulate in

the inter-membrane space creating a higher concentration there
than in the matrix (on the other side of the membrane). This proton
gradient results in protons diffusing through the ATP synthase
molecule (down the concentration gradient) making the synthase
rotor ‘spin’ and produce ATP from ADP and Pi. The diffusion of
hydrogen ions through the ATP synthase is chemiosmosis.
o The oxidation of one molecule of reduced NAD results
in six protons passing through ATP synthase and so
leads to the synthesis of three molecules of ATP.

o The oxidation of one molecule of reduced FAD results

in four protons passing through ATP synthase and so
leads to the synthesis of just two molecules of ATP.

o By adding up the number of molecules of ATP

produced, the model of aerobic respiration we have
discussed predicts that there will be a net yield of 38
molecules of ATP per molecule of glucose.
o
 Continued…
Even though most biology text books states that 38ATP
molecules can be made per oxidation of glucose (2 from
glycolysis, 2 from krebs cycle and about 34 from electron
transport system), however, this maximum yield is never
quite reached because:
 loss of protons due to leaky membranes.
 the cost of moving pyruvate and ADP in to the
mitochondria fluid matrix.
Therefore, the actual estimate ranges from 28 to 30 ATP,
per glucose.
 Continued…
The following Figure summarizes the
production of ATP in aerobic
respiration.
 Continued…
 Respirometers
Respirometers come in several different forms, but
they all work on the principle that oxygen is used in
aerobic respiration and carbon dioxide is produced.
The overall summary equation for the aerobic
respiration of glucose is:
C6H12O6 + 6O2 ➞ 6CO2 + 6H2O + 36ATP
 Continued…
What happens in the anaerobic pathway?
o If there is no oxygen present, the final reaction of oxidative
phosphorylation cannot take place. As a result, the electron
transport chain comes to a halt. No protons are
pumped and the action of ATP synthase also stops.
o If the electron transport chain does not function, NAD is not
regenerated from reduced NAD and FAD is not regenerated
from reduced FAD. Very quickly, the Krebs cycle and the
link reaction come to a halt as both NAD and FAD are
required in their oxidized forms for the Krebs cycle to function.
NAD is also required in the link reaction and so this comes to a
halt also.
o However, glycolysis can continue even though it also requires
NAD. This is because the reduced NAD formed during
glycolysis can be regenerated under anaerobic
conditions by converting the pyruvate into another
product in a reduction reaction. Reduced NAD supplies
the hydrogen for this reduction and becomes oxidized itself. It
is therefore regenerated and can be used again in glycolysis.
Continued…
 Continued….
 Different organisms produce different fermentation
end products. Animal cells produce lactate (lactic
acid) when they ferment glucose. Yeast cells produce
ethanol (ethyl alcohol). But both only produce two
molecules of ATP per molecule of glucose.
 Continued….
Lactate formation during exercise
o During exercise, the energy demand of muscle cells increases
greatly. More glucose is respired to meet the demand. However,
sometimes, aerobic respiration is insufficient to meet this energy
demand.
o Fermentation of glucose supplies the extra energy. But it also
forms lactate and as this accumulates, it leads to muscle
fatigue. Also, fermentation only yields 2 molecules of ATP per
molecule of glucose whereas aerobic respiration yields 38.
o However, fermentation is a much faster process and can
produce a lot of ATP quickly, over a short period of time.
The ATP used in sprints and short-distance runs is nearly
all generated anaerobically.
 Continued…..
o But, due to muscle fatigue, this cannot be sustained.
Longer races must be run slower to allow aerobic
respiration to produce the ATP at its slower rate.
o Lactate, once formed, can be used to regenerate
glucose or be metabolized as an energy source by the
liver.
Note: Lactate is transported back to the liver
where it is converted in to pyruvate by the cori
cycle using the enzyme lactate dehydrogenase. Then
pyruvate is used to generate glucose.
Continued…..
 Continued….
5.2 How do plants harness light energy in Photosynthesis
By the end of this section you should be able to:
Draw, label and describe a chloroplast.
Locate where light-dependent and -independent processes occur in the
chloroplast.
Name the products of the light-dependent and -independent processes.
Explain how the structure of a photosystem is related to its function.
Explain what is meant by a photosynthetic unit.
Describe how glucose is synthesized in the light- independent reactions
of photosynthesis.
Describe the factors that affect the rate of photosynthesis and explain
why they affect the rate.
Separate photosynthetic pigments by paper chromatography.
Explain photorespiration and how it is related to higher temperatures.
Distinguish between C3 and C4 plants and give at least three examples
of each.
Appreciate the importance of C4 plants in Ethiopia.
Describe the CAM photosynthetic pathway and explain why this brings
added benefits to plants living in desert conditions.
 Continued….
Photosynthesis
o In photosynthesis, light energy is used in a series of
reactions that lead to the synthesis of a range of organic
molecules. The energy that entered the system as light
is now held in the organic molecules produced. It is now
chemical energy. When energy is changed from one
form to another, we say it has been transduced.
o Light energy is absorbed by special photosensitive
pigments such as chlorophyll in the chloroplasts.
o The light-dependent reactions take place in the
membranes of the thylakoids in the chloroplasts. The
liquid stroma is the site of the light-independent
reactions, in which carbohydrates are synthesized.
Chemical reactions like these take place most effectively
in solution, rather than if some were fixed in
membranes.
 Continued…

How is the structure of a chloroplast suited

to its function?
o The chlorophyll and other photosensitive pigment
molecules are arranged in special photosystems that
are linked to electron transport chains (ETCs).
o The molecules of the photosystems and the electron
transport chains are fixed in the membranes of the
thylakoids. This makes the process much more efficient
than if they were just floating around in a solution.
o There are two different photosystems, each sensitive to
light of a different wavelength and linked to a
different electron transport chain. These are called
photosystem I and photosystem II.
Continued….
 What is the structure of a photosystem?
o A photosystem consists of a number of pigment molecules all
clustered around one particular chlorophyll molecule called
the reaction center molecule. This cluster of pigment
molecules is called an antenna complex.

NOTE:
 Photosystem is biochemical mechanism by which
chlorophyll absorbs light energy.
 Reaction center molecule is where light-dependent
reactions begin.
 Antenna complex is an array of protein and
chlorophyll light- harvesting molecules embedded in
the thylakoid membrane.
o Only the reaction center molecule is positioned next
to the electron transport chain. Energy absorbed by
other molecules in the photosystem is transferred to the
reaction center molecule, where the light-dependent
 Continued…
o Different pigment molecules in the antenna complex
can absorb different wavelengths of light, making the
whole system more efficient.
o The pigments in the antenna complex include
chlorophyll a, chlorophyll b and carotenoids. The
reaction center molecule is always chlorophyll a.
o The range of wavelengths each molecule absorbs is its
absorption spectrum.
 Continued….
What happens in the light-dependent
reactions?
The light-dependent reactions use light energy to ‘drive’
the synthesis of two molecules that will, in turn, drive the
light- independent reactions. These two molecules are:
 ATP – this provides the energy for the reactions,
and
 reduced NADP – this provides the hydrogen ions
for a key reduction reaction.
Note: NADP(Nicotinamide Adenine Dinucleotide
Phospahate) is very similar to NAD that is used in
respiration and it has the same function – transporting
hydrogen ions.
 Continued….
The main events in the light-dependent
reactions are Summarized in the following
figure primary
acceptor
primary
acceptor Fd 6.
2e
–

1. Pq NADP
2e– 3.
NADP +
reductase 2 H+
2. splitting of water cytochrome
releases oxygen complex NADP
2e– +
H 2O 5. H+
2 H+ Pc
+ light
of electrons

1/ O 2e– 4.
2 2
P700
energy

ATP photosystem I
photosystem II
 Continued….
Photosystem I and photosystem II
1. Electrons (e–) in chlorophyll molecules in photosystem
II are excited by the energy in photons of light – they
become more energetic. Because of the extra energy,
they escape from the chlorophyll and pass to an electron
acceptor (the primary electron acceptor).
2. The conditions created in the chloroplast cause the
following reaction to occur:
2H2 O → O2 + 4H+ + 4e–

This light-dependent splitting of water is called

photolysis. The electrons replace those lost from
the chlorophyll molecule.
 Continued….
3. The primary electron acceptor passes the electrons to
the next molecule in an electron transport chain
(Plastoquinone or ‘Pq’). The electrons then pass along
a series of cytochromes (similar to those in the
mitochondrial electron transport chain) and finally to
Plastocyanin (Pc) – the last carrier in the chain. The
electrons lose energy as they are passed from one
carrier to the next.
4. One of the molecules in the cytochromes complex is a
proton (hydrogen ion) pump.
As electrons are transferred to and then transferred from
this molecule, the energy they lose powers the pump
which moves protons from the stroma of the chloroplast
to the space inside the thylakoid. This leads to an
accumulation of protons inside the thylakoid, which
drives the chemiosmotic synthesis of ATP.
 Continued….
5. Electrons in chlorophyll molecules in photosystem I
are excited (as this photosystem absorbs photons of
light) and escape from the molecule. They are replaced
by the electrons that have passed down the electron
transport chain from photosystem II.
6. The electrons then pass along a second electron
transport chain involving ferredoxin (Fd) and NADP
reductase.
At the end of this electron transport chain, they(the
electrons) can react with protons (hydrogen ions) and
NADP in the stroma of the chloroplast to form reduced
NADP.
 Continued….
The following diagram shows how the molecules
are arranged in a photosynthetic unit
 Continued….
Remember
 A photosynthetic unit is a unit of pigments,
electron carriers and ATP synthase that is
capable of carrying out all the reactions in the
light-dependent stage of photosynthesis. The
formation of ATP in the way described above is
called non-cyclic photophosphorylation. This is
because:
 the phosphorylation (formation of ATP) is light-
dependent.
 the electrons lost from the chlorophyll are not
recycled in any way.
 Continued….
 Plants sometimes generate ATP by cyclic
photophosphorylation. In cyclic
photophosphorylation, only photosystem I is
used. No oxygen and no reduced NADP are
formed. Figure shows this system.
photosystem I redox chain
e–  Here, you can see that
ADP+P
Chl Fd
+ i
electrons lost from the
chlorophyll molecule are
of molecules

Pq

e– Cyt AT P returned to it. Hence the

name ‘cyclic’.
Pc
 This process usually only
energy

Chl
e–
happens when sugars cannot
be synthesized for some
reason – such as lack of
photon
carbon dioxide.
 Continued….
A summary of the light-dependent
reactions
Light energy is used to excite electrons which
then:
 cause the transfer of protons to the inside of the
thylakoid membrane as they pass along the first
electron transport chain; this eventually leads to
the formation of ATP, and
 react with hydrogen ions and NADP at the end of
the second electron transport chain to form
reduced NADP; this reaction could only happen
because of the extra energy possessed by the
electrons.
 The ATP and reduced NADP are used to drive the
synthesis of carbohydrates in the light-
 Continued….

How is carbohydrate synthesized in the

light-independent reactions
The light-independent reactions of photosynthesis
occur in the stroma of the chloroplasts.
They comprise a complex cycle of reactions that
involves the addition of carbon dioxide to a pre-
existing five-carbon molecule (RuBP or Ribulos
Bisphosohate) within the chloroplast.
The resulting molecules are modified to regenerate the
original molecule whilst, at the same time,
synthesizing glucose.
The sequence of reactions was discovered by Melvin
Calvin, an American biologist. Because of his work,
the light-independent reactions are also referred to as
the Calvin cycle
 Continued….
The main stages of the light-independent reactions are:
 carbon dioxide reacts with ribulose bisphosphate
(RuBP) – a five-carbon compound in the stroma; the
reaction is catalyzed by the enzyme Rubisco.
 two molecules of the three-carbon compound GP or
GPA (Phosphoglycerate) are formed from this
reaction.
 the cycle runs 3 times and produce 6PGA(GPA)
 each molecule of GP is converted to TP (triose
phosphate – another three-carbon compound); this is
a reduction reaction using hydrogen ions from
reduced NADP and energy from ATP.
 some of the TP formed is used to regenerate the RuBP
or Ribulose bisphosphate a 5C molecule (ATP is
again required) whilst some is used to form glucose
and other useful organic compounds.
 Continued….

1 TP

Glucose
 Continued….
Summary the light-independent reactions
of photosynthesis
three ‘turns of the cycle’ result in an output of one
molecule of TP. Six turns of the cycle would give an
output of two molecules of TP – enough to make one
molecule of glucose.
TP can also be converted to lipids, amino acids and
from these into nucleotides and all the other organic
molecules found in plants. TP is the basis for the
synthesis of all organic molecules.
 Continued….

DID yoU kNoW?

How the light-dependent and
light- independent reactions are
related light
During the light-independent reactions,
reduced NADP is reoxidised to NADP
light-dependent reactions light-independent reactions
and ATP is hydrolysed to ADP and
Pi. These are then reused in the light-
dependent reactions to regenerate ATP
and reduced NADP to be used again
in the light-independent reactions …
and so on. Figure 5.38 summarises
the relationship between the light-
dependent reactions and the light-
independent reactions. dependent and light-independent
reactions
 Continued….
What factors affect the rate of
photosynthesis?
Photosynthesis is dependent on a number of factors. The
main ones, and their effects, are shown in the table
Factor Effect on photosynthesis
below.
Light intensity Low light intensity can limit the light- dependent reactions by reducing the
number of electrons in chlorophyll molecules that are photo-excited.

Carbon dioxide concentration Can limit the light-independent reactions by influencing the rate of the initial
reaction with RuBP.

Temperature Can limit the rate of enzyme action, for example, ATP synthase (light-
dependent reactions) and Rubisco (light-independent reactions).
 Continued….
As well as the major factors discussed above, a
number of other factors influence the rate of
photosynthesis. These include:
 the wavelength of the light; photosynthesis takes
place faster in ‘red’ and ‘blue’ wavelengths than
in other wavelengths because these wavelengths
are absorbed more efficiently than others; leaves
are green because green wavelengths are
reflected.
 the amount of chlorophyll present.
 Continued….
Are there any other ways of
photosynthesizing?
 C3 photosynthesis and photorespiration
What we have just described is the method of
photosynthesis that takes place in plants living in
temperate environments, such as those found in
Europe.
It is called C3 photosynthesis – because the first
compound formed in the light-independent reactions
of the Calvin cycle is GP, which contains three carbon
atoms.
C3 plants have leaves that are adapted to this method
of photosynthesis. These leaves are generally
broad, to catch as much sunlight as possible.
 Continued….
o The cells that contain most chloroplasts (the palisade
cells) are nearest to the upper surface of the leaf (to
absorb as much light as possible). The stomata are
mainly on the lower surface, to minimize water loss.
o During the day, the stomata are open for most of the
time to allow the entry of carbon dioxide, but they can
be closed if the water loss is too great on a hot day.
The spongy mesophyll has air spaces that allow easy
diffusion of carbon dioxide and oxygen between the
palisade layer and the stomata. The following figure
shows the structure of the leaf of a C3 plant.
 Continued….
o However, plants in the tropics have a problem. Here,
it can be very hot and the leaves close their stomata
to minimise water loss.
o When C3 plants do this, the concentration of carbon
dioxide in the leaves falls and the enzyme Rubisco
starts to behave in an unusual way.
o In the low concentrations of carbon dioxide, Rubisco
binds with oxygen, not carbon dioxide. This means
that RuBP is oxidised to one molecule of GP (not
two) and a molecule of phosphoglycolate. In
addition, carbon dioxide is produced in the process.
o The process is called photorespiration because
it involves oxidation of carbon.
Ribulose bisphosphate + oxygen Rubisco GP
+phosphoglycolate + CO2
 Continued….
It is not necessary to try to remember all these reactions.
Instead, think of the two phases of photorespiration:
1. Rubisco catalyses a reaction between oxygen and
RuBP to form one molecule of GP (not two) and one
molecule of phosphoglycolate.
2. The phosphoglycolate is converted to GP in reactions
in the chloroplast, peroxisome and mitochondrion.
Photorespiration reduces the efficiency of photosynthesis
for several reasons, including:
the carbon is oxidised, which is the reverse of
photosynthesis – the reduction of carbon to
carbohydrate
the ribulose bisphosphate must be resynthesised
and the phosphoglycolate removed
ATP is used in the resynthesis of RuBP.
 Continued….
 C4 photosynthesis
To get round the problem of photorespiration reducing
the efficiency of photosynthesis, plants that grow in
tropical areas like Ethiopia (such as maize,
crabgrass, sorghum and sugar cane) have evolved
a different photosynthetic pathway called C4
photosynthesis.
As the name suggests, the first compound formed in
the light- independent reactions is a C4 compound
(contains four carbon atoms) not GP (a C3 compound).
There is a difference in how glucose is synthesized in
the light-independent reactions.
 Continued….
First, look at the structure of the leaf of a C4 plant
in figure

The structure is essentially similar to that of a C3

plant, but there is one important difference. The cells
of the bundle sheath contain chloroplasts (but
chloroplast with no thylakoid), which they don’t
in C3 plants.
Having no thylakoids means that the light-dependent
reactions cannot occur here and so oxygen is not
 Continued….
The light-dependent reactions in the C4 pathway also
involve a set of reactions not found in C3 plants.
These reactions take place in the mesophyll
cells(Palisade mesophyll), which have
chloroplasts with thylakoids and so can carry out the
light-dependent reactions. However, they do not
have the enzymes to catalyze the reactions of the
Calvin cycle. Instead, the following reactions take
place:
1. Carbon dioxide reacts with a C3 compound called
PEP (Phosphoenolpyruvate) to form the C4
compound oxaloacetate. This is catalyzed by the
enzyme PEP carboxylase.
2. Oxaloacetate is converted into another C4
compound (malate), which then passes from the
 Continued….
3. In the bundle sheath cell, malate is converted to
pyruvate with the release of a molecule of carbon
dioxide, which starts the reactions of the Calvin cycle by
binding with RuBP.
4. The pyruvate is converted back to PEP; this reaction
requires ATP.
 Continued….
Overall, the C4 cycle uses two more molecules of
ATP to deliver a molecule of carbon dioxide to
Rubisco than does the C3 cycle.
During active photosynthesis in the tropics, this is not
a problem,as the high light intensity generates much
ATP from the light-dependent reactions.
C4 photosynthesis is most efficient in conditions of:
low carbon dioxide concentration
high light intensity
high temperature
 Continued….
A comparison of C3 and C4
photosynthesis

 The crop plants that are grown in Ethiopia (such as

sorghum and wheat) are all C4 plants and are,
therefore, well adapted to photosynthesize efficiently
in the hot, bright days found in this country.
 Crop plants that are grown in temperate areas (such
as peas and carrots) would not photosynthesize as
efficiently, b/c they are C3 plants. They would,
therefore, not produce high yields.
 Continued….
Cacti and CAM (Crassulacean Acid
Metabolism) photosynthesis
In the extreme heat of deserts, having stomata open
during the day is a sure path to desiccation and death
for the plants. But if they don’t open their stomata, how
will they get the carbon dioxide they need for
photosynthesis? The answer is obvious really
– open them at night when temperatures fall.
Cacti use what is essentially the same set of reactions as
C4 plants, but they separate the two stages not by
carrying them out in different cells, but by carrying
them out at different times. The CAM photosynthesis
cycle is as follows:
 Continued….
1. At night, the plants open their stomata to
allow in CO2, which then reacts with PEP in
mesophyll cells to form oxaloacetate, and then
malate just as in the C4 pathway.
2. The malate is then stored in the vacuoles of
these cells overnight.
3. During the day, the light-dependent reactions
generate ATP and reduced NADP so that the
Calvin cycle can continue.
4. Malate is released from the vacuoles and is
broken down to glycerate, releasing carbon
dioxide for the reactions of the Calvin cycle.
Continued….
Continued….
Continued….