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The light-independent

Carbon fixation in C4 and CAM plants

This worksheet requires you to study and compare the Calvin cycle with alternative ways of
fixing carbon dioxide. You will need to discuss the advantages of each new pathway and
understand the conditions that make them beneficial. You will need a good knowledge of the
Calvin cycle to complete this worksheet.

The problem of photorespiration

During the light-independent reaction of photosynthesis, carbon dioxide is accepted by
another molecule. In most plants this is ribulose bisphosphate (RuBP). The reaction forms
glycerate 3-phosphate which is then reduced by the NADP which is produced in the lightdependent reaction. Plants that use this pathway during the light-independent reaction of
photosynthesis are called C3 plants. The problem with this is that RuBP is not particularly
efficient, as it also has an affinity for oxygen.
The ability of RuBP to accept both carbon dioxide and oxygen means that, when CO 2 levels
in the leaf drop, RuBP combines with oxygen in a process called photorespiration. The
products of this reaction are CO2 and water, but no ATP. This makes photosynthesis much
less efficient, as intermediates required by the Calvin cycle are used up.

C4 and CAM pathways as solutions

C4 plants use an alternative molecule to combine with the carbon dioxide,
phosphoenolpyruvate (PEP), which has a higher affinity with it. This allows the plants to
store carbon dioxide even when concentrations of the gas are low within the leaf due to
stomatal closure. It can later be released within the leaf at the site of the light-independent
reaction, ensuring a high concentration where it is needed.
Within the mesophyll cells, PEP combines with carbon dioxide to form oxaloacetate. This is
converted to malate, a four-carbon acid-based molecule, which is moved to the cells
surrounding the xylem and phloem cells, called bundle sheath cells. The carbon dioxide is
then liberated before being fixed again in the normal pathway. Although this method allows
plants to store carbon dioxide so that they can use it later, more energy is required because
the gas is fixed twice.
C4 plants are adapted to conditions of extremely high light intensities, as carbon dioxide does
not become a limiting factor even when the amount available is very low. They also have a
higher rate of photosynthesis in warmer temperatures than C 3 plants. These very efficient
plants grow faster than C3 plants and have a higher yield.
An alternative to the C4 pathway is the crassulacean acid metabolism (CAM). This pathway is
found in many xerophytic plants (plants adapted to live in dry conditions). The CAM
pathway also uses PEP as the molecule which combines with carbon dioxide, but it is only
used at night. It occurs within the bundle sheath cells, which are found next to the xylem and
phloem cells. The Calvin cycle occurs as usual during the day using carbon dioxide liberated
from the malate formed during the night.
CAM plants are highly adapted to growing in conditions of drought. They are able to
conserve much more water than C4 or C3 plants as their stomata mainly open during the
cooler conditions of the night.

AQA Biology A2 Stretch and challenge Nelson Thornes Ltd 2009

The light-independent

1 Outline the events that take place in a C3 plant during the fixation of carbon dioxide
(the Calvin cycle).
2 Although C4 plants are more efficient at high light intensities and high temperatures, low light
intensities and low temperatures favour C 3 plants. Suggest why C4 plants are more efficient at
higher light intensities and higher temperatures.
3 The chain of reactions initiated when RuBP accepts oxygen is called photorespiration. Suggest why.
4 The molecule that combines with carbon dioxide in C4 plants is PEP but in C3 plants the molecule is
RuBP. PEP has a much higher affinity for carbon dioxide than RuBP and will not accept oxygen.
Explain how this helps C4 plants.
5 Gas exchange in CAM plants takes place mainly at night as this is when the CAM pathway fixes
carbon dioxide. Explain how this helps these plants avoid excessive water loss.
6 The main difference between C4 and CAM pathways is the way by which carbon dioxide supply to
the Calvin cycle is maintained. Explain how the carbon dioxide supply is maintained and what the
benefit to the plant is in each case.
7 In what ways does the C4 pathway differ from the CAM pathway?
8 Genetic modification of plants may make it possible to transfer the genes for the C 4 or CAM
pathways to C3 plants. As a result the genetically modified plants could be more efficient and better
adapted to the conditions in many developing countries. What would a scientist consider before
deciding to genetically modify a C3 plant with genes from one of the C4 or CAM pathways?

AQA Biology A2 Stretch and challenge Nelson Thornes Ltd 2009