A Level Biology Unit 7 page 1

Heckmondwike Grammar School Biology Department

Edexcel A-Level Biology B

Contents
Respiration ............................................................................... p4
Glycolysis and the Krebs Cycle .................................... p6
The electron transport chain ........................................ p8
Anaerobic respiration...................................................... p10
Photosynthesis......................................................................... p12
The light-dependent reactions ...................................... p14
The light-dependent reactions ...................................... p16
Factors affecting the rate of photosynthesis .............. p17

These notes may be used freely by biology students and teachers.

I would be interested to hear of any comments and corrections.
Neil C Millar ([email protected]) June 2016

Unit 1 Biochemistry Unit 2 Cells

Y12 Unit 3 Reproduction Unit 4 Transport
Unit 5 Biodiversity
Unit 6 Ecology
Y13 Unit 7 Metabolism Unit 8 Microbes
Unit 9 Control Systems Unit 10 Genetics

HGS Biology A-level notes NCM/5/15

A Level Biology Unit 7 page 2

Biology Unit 7 Metabolism

Specification
Aerobic respiration from one molecule of hexose sugar in aerobic
Cellular respiration yields ATP, which is used as a conditions compared with anaerobic conditions.
source of energy for metabolic reactions; and the How lactate as a by-product of anaerobic
process also generates heat. The different stages respiration affects mammalian muscle
in aerobic respiration, including: glycolysis in the contraction. How anaerobic respiration in plants
cytoplasm; link reaction; Krebs cycle; oxidative results in ethanol formation.
phosphorylation in mitochondria.
Glycolysis. The conversion of Photosynthesis
monosaccharides to pyruvate during glycolysis The structure of chloroplasts, including:
in the cytoplasm, including: the envelope, stroma, grana and lamellar structure.
phosphorylation of hexose molecules by ATP; Photosynthetic pigments. What is meant by
breakdown to glycerate 3-phosphate (GP); absorption and action spectra. Why many plants
production of reduced coenzyme (NADH) and have a variety of different photosynthetic
ATP (details of intermediate compounds and pigments.
other reactions are not required). Light-dependent stage. The role of the
Link reaction and Krebs cycle. The link thylakoid membranes in the light-dependent
reaction and Krebs cycle take place in the stage of photosynthesis. The processes of
mitochondrial matrix. During the complete cyclic and non-cyclic photophosphorylation
oxidation of pyruvate the events of the link result in the production of reduced NADP,
reaction and the Krebs cycle result in the ATP and oxygen.
removal of carbon atoms to produce: carbon Light-independent stage. The role of the
dioxide; reduced coenzyme (NADH); ATP stroma in the light-independent stage of
(detailed ledge of the intermediate compounds photosynthesis. How carbon dioxide is fixed
in the Krebs; cycle is not required). by combination with 5C ribulose bisphosphate
Oxidative phosphorylation. The electron (RuBP) to form glycerate 3-phosphate (GP)
transport chain takes place in the inner using the enzyme ribulose bisphosphate
mitochondrial membrane. The role of the carboxylase (RUBISCO). How reduced NADP
electron transport chain in generating ATP and ATP from the light-dependent stage are
(oxidative phosphorylation). The role of used to synthesise glyceraldehyde phosphate
oxygen as a terminal electron acceptor forming (GALP) from GP. How ribulose bisphosphate
water. How ATP is synthesised by is regenerated in the Calvin cycle (details of
chemiosmosis. The importance of intermediate compounds are not required).
mitochondrial membranes in this process. How GALP is used as a raw material in the
production of monosaccharides, amino acids
Anaerobic respiration and other molecules.
Anaerobic respiration is the partial breakdown of The factors that limit photosynthesis including
hexoses to produce a limited yield of ATP in the carbon dioxide, light intensity and temperature.
absence of oxygen. The difference in ATP yields

HGS Biology A-level notes NCM/5/15

A Level Biology Unit 7 page 3

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A Level Biology Unit 7 page 4

Respiration
All living cells require energy, and this energy is provided by the oxidation of glucose respiration.
exothermic
glucose + oxygen
carbon dioxide + water

The oxidation of glucose is strongly exothermic, but in respiration the energy is released not as heat, but in
the form of chemical energy in a compound called ATP (adenosine triphosphate). ATP is built up from ADP
3-
and phosphate ( PO4 , abbreviated to Pi). So all respiration really does is convert chemical energy stored in
glucose into chemical energy stored in ATP.

ATP is a nucleotide (one of the four found in DNA), but it also has this other function as an energy storage
molecule. So ATP is actually a bigger molecule than glucose, but it is very soluble and the energy it contains
can be released very quickly and easily. As we'll see later, over 30 molecules of ATP can be made from each
glucose molecule in respiration, so ATP stores a much smaller amount of energy than glucose. This is a
good thing, as these small packets of easily-released energy are more useful to cells and can be used to do
simple common jobs, as the next paragraph shows. An analogy would be that small change (ATP) is often
more useful than large bank notes (glucose).

What is the energy in ATP used for?

The processes in a cell that require energy can be put into three groups:
Muscle contraction and other forms of movement, such as cilia, flagella, cytoplasmic streaming, etc.
Each step of the muscle crossbridge cycle costs one ATP molecule.
Active transport. Each shape change in an active transport protein pump costs one ATP molecule.
Biosynthesis building up large molecules from smaller ones, e.g. protein synthesis, DNA replication,
starch synthesis, etc. Each monomer added to a growing polymer chain costs one ATP molecule.
Since these processes use ATP, they all involve ATPase enzymes. ATPases catalyse the hydrolysis of ATP to
ADP + Pi, and do work with the energy released.

All the thousands of chemical reactions taking place in a cell are referred to as Metabolism. To make the
reactions easier to understand, biochemists arrange them into metabolic pathways. The intermediates in
these metabolic pathways are called metabolites.
A Level Biology Unit 7 page 5

Mitochondria
Much of respiration takes place in the mitochondria. Mitochondria
have a double membrane: the outer membrane contains many
protein channels called porins, which let almost any small molecule
through; while the inner membrane is more normal and is
selectively permeable to solutes. The inner membrane is highly
folded into projections called cristae, giving a larger surface area.
The electron microscope reveals blobs on the inner membrane,
called stalked particles. These blobs have now been identified as
enzyme complexes that synthesise ATP, and are more correctly
called ATP synthase enzymes (more later). The space inside the inner membrane is called the matrix, and is
where the Krebs cycle takes place. The matrix also contains DNA, tRNA and ribosomes, and some genes
are replicated and expressed here.

Details of Respiration
The equation for cellular respiration is usually simplified to:
glucose + oxygen carbon dioxide + water (+ energy)
But in fact this is a summary of a complex metabolic pathway, comprising at least 30 separate steps. To
understand respiration in detail we can first break it up into 3 stages:

Before we look at these stages in detail, there are a few points to note from this summary.
The different stages of respiration take place in different parts of the cell. This compartmentation allows
the cell to keep the various metabolites separate, and to control the stages more easily.
The energy released by respiration is in the form of ATP.
Since this summarises so many separate steps (often involving H+ and OH- ions from the solvent water),
it is meaningless to try to balance the summary equation.
The release of carbon dioxide takes place before oxygen is involved. It is therefore not true to say that
respiration turns oxygen into carbon dioxide; it is more correct to say that respiration turns glucose
into carbon dioxide, and oxygen into water.
Stage 1 (glycolysis) is anaerobic respiration, while stages 2 and 3 are the aerobic stages.
A Level Biology Unit 7 page 6

Glycolysis and the Krebs Cycle

A Level Biology Unit 7 page 7

1. Glucose and other hexose sugars enter cells from the tissue fluid by facilitated diffusion using a specific
hexose carrier. This carrier can be controlled (gated) by hormones such as insulin, so that uptake of
sugars can be regulated.

2. The first step in glycolysis is the phosphorylation of the hexose to form hexose phosphate (usually
glucose phosphate), using phosphate from ATP. There are two reasons for this step. Firstly, it keeps
glucose in the cell by effectively removing pure glucose, so glucose will always diffuse down its
concentration gradient from the tissue fluid into the cell (glucose phosphate no longer fits the
membrane carrier). Secondly, it activates glucose for biosynthesis reactions: glucose phosphate is the
starting material for the synthesis of pentose sugars (and therefore nucleotides and DNA), glycogen and
starch.

3. Glucose is phosphorylated again (using another ATP) and split into two molecules of glyceraldehyde
phosphate (GALP), which is a triose (3 carbon) sugar. From now on everything happens twice per
original glucose molecule.

4. GALP is oxidised to form glycerate-3-phosphate (GP), a 3-carbon acid. In this step some energy is
released to form ATP, and a hydrogen atom is also released (which is why the reaction is an oxidation).
This hydrogen atom is very important as it stores energy, which is later used by the electron transport
chain to make more ATP. The hydrogen atom is taken up and carried to the electron transport chain by
the coenzyme NAD, which becomes reduced in the process.

(oxidised form of coenzyme ) NAD + H NADH ( reduced form of coenzyme)

5. GP is converted in a series of steps to form the 3-carbon compound pyruvate. Another ATP is made
during this process. Pyruvate marks the end of glycolysis, the first stage of respiration. Pyruvate can also
be turned back into glucose by reversing glycolysis, and this is called gluconeogenesis.

6. In the absence of oxygen pyruvate is converted into lactate or ethanol in anaerobic respiration (p 10).

7. In the presence of oxygen pyruvate enters the mitochondrial matrix to proceed with aerobic
respiration. Once pyruvate has entered the inside of the mitochondria (the matrix), it is converted to a
compound called acetyl coA. Since this step links glycolysis and the Krebs Cycle, it is referred to as the
link reaction. In this reaction pyruvate loses a CO2 and a hydrogen to form a 2-carbon acetyl compound,
which is temporarily attached to another coenzyme called coenzyme A (or just coA), so the product is
called acetyl coA. The CO2 diffuses through the mitochondrial and cell membranes by lipid diffusion, out
into the tissue fluid and into the blood, where it is carried to the lungs for removal. The hydrogen is
taken up by NAD again.

8. The acetyl CoA then enters the Krebs Cycle, named after Sir Hans Krebs, who discovered it in the
1940s at Sheffield University. It is one of several cyclic metabolic pathways, and is also known as the
citric acid cycle or the tricarboxylic acid cycle. The 2-carbon acetyl is transferred from acetyl coA to the
4-carbon oxaloacetate to form the 6-carbon citrate. Citrate is then gradually broken down in several
steps to re-form oxaloacetate, producing carbon dioxide and hydrogen in the process. Some ATP is also
made directly in the Krebs cycle. As before, the CO2 diffuses out the cell and the hydrogen is taken up
by NAD, or by an alternative hydrogen carrier called FAD. These hydrogen atoms are carried to the
inner mitochondrial membrane for the final part of respiration.
A Level Biology Unit 7 page 8

The Electron Transport Chain

The electron transport chain (or respiratory chain) is an unusual metabolic pathway in that it takes place
within the inner mitochondrial membrane, using integral membrane proteins. There are five of these
proteins, called I, II, III, IV and ATP synthase. They each contain up to 40 individual polypeptide chains,
which perform many different functions including enzymes and trans-membrane pumps.

In the electron transport chain the hydrogen atoms from NADH gradually release all their energy to form
ATP, and are finally combined with oxygen to form water.

1 NADH molecules bind to protein I and release their hydrogen atoms as protons (H+) and electrons (e-).
FADH binds to complex II rather than complex I to release its hydrogen. The NAD and FAD molecules
then return to the Krebs Cycle to collect more hydrogen, so these coenzymes are constantly shuttling
between their oxidised and reduced forms:

2. The electrons are passed along the chain of proteins. The energy of the electrons is used to pump
protons across the inner mitochondrial membrane by active transport through the proteins, forming a
proton gradient across the membrane.
3. Finally, the electrons are combined with protons and molecular oxygen (O2) to form water, the final
end-product of respiration. The oxygen diffused in from the tissue fluid, crossing the cell and
mitochondrial membranes by lipid diffusion. Oxygen is only involved at the very last stage of respiration
as the final electron acceptor, but without it the whole electron transport chain stops.
4. The energy of the electrons is now stored in the form of the proton gradient across the inner
mitochondrial membrane. Its a bit like using energy to pump water uphill into a high reservoir, where it
is stored as potential energy. And just as the potential energy in the water can be used to generate
A Level Biology Unit 7 page 9

electricity in a hydroelectric power station, so the potential energy in the proton gradient can be used
to generate ATP in the ATP synthase enzyme. The ATP synthase enzyme has a proton channel through
it, and as the protons fall down this channel their energy is used to make ATP, spinning the globular
head as they go. It takes four protons to synthesise one ATP molecule.

This synthesis of ATP is called oxidative phosphorylation because it uses oxygen to phosphorylate ADP.

Chemiosmosis
The method of storing energy by creating a proton gradient across a membrane is called chemiosmosis, and
was discovered by Peter Mitchell in the 1960s, for which work he got a Nobel prize in 1978. Some poisons
act by making proton channels in mitochondrial membranes, so giving an alternative route for protons and
stopping the synthesis of ATP. This also happens naturally in the brown fat tissue of new-born babies and
hibernating mammals: respiration takes place, but no ATP is made, with the energy being turned into heat
instead.

Summary of Oxidative Phosphorylation

1. NADH releases its H and is oxidised to NAD, which returns to the Krebs cycle
(NADH NAD + H+ + e-).
2. The electron is passed along the chain of proteins in the inner mitochondrial membrane, releasing its
energy as it goes.
3. Oxygen combines with hydrogen to form water (O2 + H+ + e- H2O).
4. The energy of the electron is used to make ATP in the ATP synthase enzyme (ADP + Pi ATP).
A Level Biology Unit 7 page 10

Anaerobic Respiration
If there is no oxygen (anaerobic conditions) then the final reaction to make water cannot take place, no
electrons can leave the electron transport chain, and so NADH cannot unload any hydrogens to the
electron transport chain. This means that there is no NAD in the cell; its all in the form of NADH.
Without NAD as a coenzyme, some of the enzymes of the Krebs cycle and glycolysis cannot work, so the
whole of respiration stops.

Some cells can get round this problem using anaerobic respiration. This adds an extra step to the end of
glycolysis that regenerates NAD, so allowing glycolysis to continue and some ATP to be made. Anaerobic
respiration only makes 2 ATPs per glucose, but its better than nothing. There are two different kinds of
anaerobic respiration:

Lactic Acid Anaerobic Respiration Ethanolic Anaerobic Respiration

In animals and bacteria the extra step converts In plants and fungi the extra steps converts pyruvate
pyruvate to lactate (or lactic acid). This is a to ethanol. This is also a reduction, so NADH is
reduction, so NADH is used and NAD is used and NAD is regenerated, to be used in
regenerated, to be used in glycolysis. The reaction is glycolysis. Ethanol is a two-carbon compound and
reversible, so the energy remaining in the lactate carbon dioxide is also formed. This means the
molecule can be retrieved when oxygen becomes reaction is irreversible, so the energy in the ethanol
available and the lactate is oxidised via the rest of cannot be retrieved by the cells.
aerobic respiration.
Ethanolic anaerobic respiration is also known as
The bacteria used to make yogurt use this reaction, fermentation, and we make use of fermentation in
as do muscle cells and red blood cells in humans. yeast to make ethanol in beer and wine.
Lactic acid is a fairly strong acid so it lowers the pH
in muscles cells, which slows enzymes and
contributes to muscle fatigue and cramp. So
anaerobic respiration in muscle cells cannot be
continued for very long.
A Level Biology Unit 7 page 11

How Much ATP is made in Respiration?

We can now summarise respiration and see how much ATP is made from each glucose molecule. ATP is
made in two different ways:
Some ATP molecules are made directly by the enzymes in glycolysis or the Krebs cycle. This is called
substrate level phosphorylation (since ADP is being phosphorylated to form ATP).
Most of the ATP molecules are made by the ATP synthase enzyme in the electron transport chain. Since
this requires oxygen it is called oxidative phosphorylation. Scientists dont yet know exactly how many
protons are pumped in the electron transport chain, but the current estimates are: 10 protons pumped
by NADH; 6 by FADH; and 4 protons are needed by ATP synthase to make one ATP molecule. This
means that each NADH can make 2.5 ATPs (10/4) and each FADH can make 1.5 ATPs (6/4). Previous
estimates were 3 ATPs for NADH and 2 ATPs for FADH, and these numbers still appear in most
textbooks, although they are now probably wrong.
Two ATP molecules are used at the start of glycolysis to phosphorylate the glucose, and these must be
subtracted from the total.

The table below is an ATP account for aerobic respiration, and shows that 32 molecules of ATP are
made for each molecule of glucose used in aerobic respiration. This is the maximum possible yield; often
less ATP is made, depending on the circumstances. Anaerobic respiration only produces the 2 molecules of
ATP from the first two rows. You dont need to learn this table, but you should understand it.
Final ATP yield Final ATP yield
Stage molecules produced per glucose
(old method) (new method)
Glycolysis 2 ATP used -2 -2
4 ATP produced (2 per triose phosphate) 4 4
2 NADH produced (1 per triose phosphate) 6 5
Link Reaction 2 NADH produced (1 per pyruvate) 6 5
Krebs Cycle 2 ATP produced (1 per acetyl coA) 2 2
6 NADH produced (3 per acetyl coA) 18 15
2 FADH produced (1 per acetyl coA) 4 3
Total 38 32

Other substances can also be used to make ATP. Glycogen of course is the main source of glucose in
humans. Triglycerides are broken down to fatty acids and glycerol, both of which enter the Krebs Cycle. A
typical triglyceride molecule might make 50 acetyl CoA molecules, yielding 500 ATP molecules. Fats are
thus a very good energy store, yielding 2.5 times as much ATP per g dry mass as carbohydrates. Proteins
are not normally used to make ATP, but in starvation they can be broken down and used in respiration.
They are first broken down to amino acids, which are converted into pyruvate and Krebs Cycle
metabolites and then used to make ATP.
A Level Biology Unit 7 page 12

Photosynthesis
Photosynthesis is essentially the reverse of respiration. It is usually simplified to:
carbon dioxide + water (+ light energy) glucose + oxygen
But again this simplification hides numerous separate steps. To understand photosynthesis in detail we can
break it up into 2 stages:

The light-dependent reactions use light energy to split water and make ATP, oxygen and energetic
hydrogen atoms. This stage takes place within the thylakoid membranes of chloroplasts, and is very
much like the electron transport chain, only in reverse.
The light-independent reactions dont need light, but do need the products of the light-dependent stage
(ATP and H), so they stop in the absence of light. This stage takes place in the stroma of the
chloroplasts and involves the fixation of carbon dioxide and the synthesis of glucose.
Like respiration, oxygen and carbon dioxide are quite separate. Plants do not turn carbon dioxide into
oxygen; they turn carbon dioxide into glucose, and water into oxygen.
We shall see that there are many similarities between photosynthesis and respiration, and even the same
enzymes are used in some steps.

Chloroplasts
Photosynthesis takes place entirely within chloroplasts. Like
mitochondria and nuclei, chloroplasts are surrounded by a
double membrane called an envelope, but in addition
chloroplasts have a third membrane called the thylakoid
membrane. This membrane is folded into disk-shaped vesicles
called thylakoids, enclosing small spaces called the thylakoid
lumen. The thylakoid vesicles are usually layered in stacks
called grana and the grana are linked by numerous thylakoid
lamellae. The thylakoid membrane contains the same ATP
synthase particles found in mitochondria. Chloroplasts also
contain DNA, tRNA and ribososomes, and they often store the
products of photosynthesis as starch grains and lipid droplets.
A Level Biology Unit 7 page 13

Photosynthetic Pigments
Chloroplasts contain a number of different photosynthetic pigments that absorb the light energy used in
photosynthesis. There are two general types of pigment:
The chlorophylls, including chlorophyll a (blue-green) and chlorophyll b (olive green).
The carotenoids, including -carotene (red-orange) and xanthophyll (orange-red).
Chlorophyll a is the most important pigment in photosynthesis, while the others act as accessory pigments,
absorbing different wavelengths of light so that more of the visible spectrum is used.

The absorption spectra of pure samples of some of these

pigments are shown in the graph on the left. A low absorption
means that those wavelengths are not absorbed and so cannot
be used, but instead are reflected or transmitted. Different
species of plant have different combinations of photosynthetic
pigments, giving rise to different coloured leaves.

By measuring the rate of photosynthesis using different

wavelengths of light, an action spectrum is obtained. The
action spectrum can be well explained by the absorption
spectra above, showing that these pigments are responsible
for photosynthesis.

Photosystems
Chlorophyll is a fairly small molecule (not a protein) with a structure similar to haem, but with a magnesium
atom instead of iron. Chlorophyll and the other pigments are arranged in complexes with proteins, called
photosystems. Each photosystem contains some 200 chlorophyll molecules and 50 molecules of accessory
pigments, together with several protein molecules (including enzymes) and lipids. These photosystems are
located in the thylakoid membranes and they hold the light-absorbing pigments in the best position to
maximise the absorbance of photons of light. The chloroplasts of green plants have two kinds of
photosystem called photosystem I (PSI) and photosystem II (PSII). These absorb light at different
wavelengths and have slightly different jobs in the light dependent reactions of photosynthesis, as we shall
see.
A Level Biology Unit 7 page 14

The Light-Dependent Reactions

The light-dependent reactions (or photophosphorylation) take place on the thylakoid membranes using
four membrane-bound proteins: photosystem I (PSI), photosystem II (PSII), cytochrome (C) and ferredoxin
NADP reductase (FNR). In these reactions light energy is used to split water, oxygen is given off, hydrogen
is produced and ADP is phosphorylated to make ATP.

1. Chlorophyll molecules in PSII absorb photons of light, exciting chlorophyll electrons to a higher energy
level and causing a charge separation within PSII. This charge separation drives the splitting (or
photolysis) of water molecules to make oxygen (O2), protons (H+) and electrons (e-):
photolysis
2H2O O2 + 4H+ + 4e-
Water is a very stable molecule and it requires the energy from 4 photons of light to split one water
molecule. The oxygen produced diffuses out of the chloroplast and eventually into the air; the protons
build up in the thylakoid lumen causing a proton gradient; and the electrons from water replace the
excited electrons that have been ejected from chlorophyll.

2. The excited, high-energy electrons are passed along the chain of proteins in the thylakoid membrane, in
a similar way to the electron transport chain. In PSI more light energy is absorbed and passed to the
electron, which gains energy as it goes. The energy of the electrons is used to pump protons from
stroma to lumen, creating a proton gradient across the thylakoid membrane.

3. At the final complex, the electron is recombined with a proton to form a hydrogen atom, which is taken
up by the coenzyme NADP, reducing it to NADPH.
NADP + H+ + e- NADPH
Note that while respiration uses the coenzyme NAD to carry hydrogen, photosynthesis always uses its
close relative, NADP.

4. The combination of the water splitting and proton pumping causes a proton gradient across the
thylakoid membrane. This gradient is used to make ATP using the ATP synthase enzyme in exactly the
same way as respiration. This synthesis of ATP is called photophosphorylation because it uses light
energy to phosphorylate ADP (ADP + Pi ATP).
A Level Biology Unit 7 page 15

Cyclic and Non-Cyclic Photophosphorylation

There is an alternative light-dependent reaction that plants use sometimes, called cyclic
photophosphorylation, which uses photosystem I but not photosystem II. In this reaction, chlorophyll
molecules in PSI absorb light and the excited chlorophyll electrons are passed to the cytochrome complex,
where their energy is used to pump protons across the thylakoid membrane before being returned to
photosystem I. The proton gradient is used to make ATP using chemiosmosis and the ATP synthase
enzyme in exactly the same way as normal.

This process is called cyclic photophosphorylation because electrons just cycle between photosystem I and
cytochrome. By contrast the full light-dependent reaction is called non-cyclic photophosphorylation
because the electrons dont cycle, but are passed from water along the chain of proteins and finally to
NADPH.

What is the purpose of cyclic photophosphorylation? Non-cyclic photophosphorylation makes roughly

equal quantities of ATP and NADPH, but plant cells actually use more ATP than NADPH, so when ATP is
running out they can switch to cyclic photophosphorylation to just make ATP. In cyclic
photophosphorylation no NADPH or oxygen is made and no water is split.

Summary of Non-Cyclic Photophosphorylation

1. Light energy is absorbed by chlorophyll and used to photolyse water (H2O O2 + H+ + e-).
2. The high-energy electron is passed along the chain of proteins in the thylakoid membrane, gaining energy
from light as it goes.
3. The electron is taken up by NADP, which is reduced to NADPH (NADP + H+ + e- NADPH).
4. The energy from the light is used to make ATP in the ATP synthase enzyme (ADP + Pi ATP).
A Level Biology Unit 7 page 16

The Light-Independent Reactions

The light-independent, or carbon-fixing reactions, of photosynthesis take place in the stroma of the
chloroplasts and comprise another cyclic pathway, called the Calvin Cycle, after the American scientist who
discovered it.

1. Carbon dioxide binds to the 5-carbon sugar ribulose bisphosphate (RuBP) to form 2 molecules of the 3-
carbon acid glycerate phosphate (GP), the same compound as that found in glycolysis. This carbon-fixing
reaction is catalysed by the enzyme ribulose bisphosphate carboxylase, always known as rubisco. It is a
very slow and inefficient enzyme, so large amounts of it are needed (recall that increasing enzyme
concentration increases reaction rate), and it comprises about 50% of the mass of chloroplasts, making
it the most abundant protein in nature.

2. Glycerate phosphate (C3H4O4.PO3) is not a carbohydrate, so it is reduced by NADPH and activated by

ATP to form the sugar glyceraldehyde phosphate (GALP) (C3H6O3.PO3), the same triose sugar as that
found in glycolysis. Two ATP and two NADPH molecules from the light-dependent reactions provide
the energy for this step. The ADP and NADP return to the thylakoid membrane for recycling.

3. GALP is a branching point. Most of the GALP continues through a complex series of reactions to
regenerate the RuBP and complete the cycle. 5 GALP molecules (5 x 3C = 15 carbon atoms) combine
to form 3 RuBP molecules (3 x 5C = 15 carbon atoms).

4. Every 3 turns on average of the Calvin Cycle 3 CO2 molecules are fixed to make 1 new GALP molecule
(3CO2 + 6H C3H6O3). This GALP leaves the cycle, and two of these GALP molecules combine to
form one glucose molecule using the glycolysis enzymes in reverse.

5. The light-independent reactions are now finished, and the glucose can now be transported out of the
chloroplast and used to make all the other organic compounds that the plant needs (cellulose, lipids,
proteins, nucleic acids, etc). Some of these need the addition of mineral elements like N, P or S. Plants
are very self-sufficient!
A Level Biology Unit 7 page 17

Factors affecting the rate of Photosynthesis

The rate of photosynthesis by a plant or alga can be measured by recording the amount of oxygen
produced, or carbon dioxide used, in a given period of time. But these measurements are also affected by
respiration, which plants do all the time, so the respiration rate must be measured separately. Weve
already seen that a plants growth (or productivity) depends on the difference between the rates of
photosynthesis and respiration. The conditions at which the rates of photosynthesis and respiration are
equal, so there is no net change in oxygen or carbon dioxide concentration, is called the compensation
point. Many of the environmental factors that affect photosynthesis also affect respiration.

Temperature. Temperature affects the rates of all enzyme

reactions, so the rates of photosynthesis and respiration are
both affected. Photosynthesis is more sensitive to
temperature with an optimum of about 30-35C, whereas
respiration often has an optimum nearer to 45C. So there is
a temperature compensation point around 40C (A), and
above this temperature plants lose mass as the rate of
respiration is greater than the rate of photosynthesis.

Carbon dioxide concentration. Carbon dioxide is the

substrate for the enzyme rubisco in the light-independent
stages of photosynthesis, so the higher the carbon dioxide
concentration the faster the rate of the Calvin cycle. The rate
of respiration is not affected by carbon dioxide
concentration, and the carbon dioxide compensation point is
usually very low, at about 50ppm (A). Normal carbon dioxide
concentration in the air is about 400ppm (B), whereas the optimum concentration for most plants is
nearer to 1000ppm, so carbon dioxide is often the limiting factor.

Light intensity. Light is the source of energy for the

production of ATP and NADPH in the light-dependent stages
of photosynthesis, so the higher the light intensity the faster
the rate of photosynthesis. The rate of respiration is not
affected by light intensity, and the light compensation point is
usually low. Shade plants are adapted to growing in low light
conditions (such as a forest floor), so have a very low light
compensation point (A) and a low optimum intensity. Shade plants make good house plants, since they
are adapted to the low light intensities indoors. Sun plants have a higher compensation point (B), and
have a very high optimum near the light intensity of a bright summers day (C).
A Level Biology Unit 7 page 18

Time of Day. Both photosynthesis and respiration are

affected by time of day: photosynthesis by changes in light and
respiration by changes in temperature. At night respiration
exceeds photosynthesis, while during the day photosynthesis
exceeds respiration, so there are two compensation points
each day (A and B). Over a 24-hour period the amount of
photosynthesis is greater than the amount of respiration, so
plants gain mass and have a net uptake of carbon dioxide.

Understanding how factors affect photosynthesis and respiration is very important for farmers and
commercial growers. For example in a closed greenhouse with lots of plants the carbon dioxide
concentration can fall very low, so it can be worth increasing the CO2 concentration in the greenhouse to
increase the rate of photosynthesis. This is most efficiently done by burning a fuel, since this releases CO2
and raises the temperature. Its hard to beat the intensity of daylight, but day length can be increased with
artificial lighting.

Limiting Factors
Although all these factors (and many others) could affect the rate of photosynthesis, at any given time there
can only be one factor that is actually controlling the rate the limiting factor. This is the factor that is in
shortest supply, or furthest from its optimum. Its a bit like a chain that is only as strong as its weakest link.
We can observe this in an experiment to investigate the rate of photosynthesis.

1. At low light intensities the rate of photosynthesis increases as the light intensity increases. This must
mean that light is the limiting factor, since the rate depends on it.
2. At higher light intensities the rate of photosynthesis stays the same even if the light intensity increases.
This means that light is not the limiting factor, since the rate doesnt depend on it. This isnt surprising
since there is now plenty of light. The rate of photosynthesis must be limited by some other factor.
3. If we repeat the experiment at a higher concentration of carbon dioxide we get a higher rate, showing
that a higher rate is possible, and that carbon dioxide concentration was rate limiting at point 2.