Crassulacean acid metabolism

• Crassulacean acid metabolism, also known as CAM photosynthesis,

is a carbon fixation pathway that evolved in some plants as an
adaptation to arid conditions.
• In a plant using full CAM, the stomata in the leaves remain shut
during the day to reduce evapotranspiration, but open at night to
collect carbon dioxide (CO2), and allow it to diffuse into the
mesophyll cells. The CO2 is stored as the four-carbon acid malate in
vacuoles at night, and then in the daytime, the malate is transported
to chloroplasts where it is converted back to CO 2, which is then used
during photosynthesis. The pre-collected CO2 is concentrated around
the enzyme RuBisCO, increasing photosynthetic efficiency. The
mechanism was first discovered in plants of the family Crassulaceae.
CAM is an adaptation for increased efficiency in the
use of water, and so is typically found in plants
growing in arid conditions
During the night
During the night, a plant employing CAM has its
stomata open, allowing CO2 to enter and be fixed as
organic acids by a PEP reaction similar to the C4
pathway. The resulting organic acids are stored in
vacuoles for later use, as the Calvin cycle cannot
operate without ATP and NADPH, products of light-
dependent reactions that do not take place at night
During the day
• During the day the stomata close to conserve water,
and the CO2-storing organic acids are released from
the vacuoles of the mesophyll cells. An enzyme in
the stroma of chloroplasts releases the CO2, which
enters into the Calvin cycle that photosynthesis may
take place
 Steps of CAM Photosynthesis

• CAM photosynthesis begins at night, when the

plant’s stomata open and CO2 gas is able to diffuse
into the cytoplasm of CAM mesophyll cells.
• In the cytoplasm of those cells, the CO2 molecules
encounter hydroxyl ions, OH−, which they combine
with to become HCO3 the enzyme
phosphoenolpyruvate carboxylase (PEP carboxylase).
• CO2 + OH− → HCO3
2. The PEP carboxylase enzyme catalyzes the following reaction to add
the CO2 to a molecule called phosphoenolpyruvate (PEP).
PEP + HCO3− → OXALOACETATE
3. Oxaloacetate then receives an electron from NADH and becomes a
molecule of malate. This reaction is catalyzed by the enzyme Malate
Dehydrogenase (MDH). That reaction looks like:
OXALOACETATE + NADPH + MDH → MALATE + NADP+
Interestingly, malate dehydrogenase catalyzes a reversible reaction,
meaning that it can either add electrons to oxaloacetate, or take electrons
away from molecules of malate.
4. Malate is now stored in vacuoles within the plant cells, until the sun
rises and photosynthesis begins. When that happens, malate enters the 
Calvin Cycle, just like 3-phosphoglycerate would in a plant using a 3-
carbon, or “C3” pathway for carbon fixation.
 Examples of CAM Plants

CAM metabolism is common in plants that live in hot, dry

environments where water is difficult to gain and conserve.
Examples include:
Cacti
• Typical cacti have a rounded shape, which minimizes the
surface area through which they can lose water during the
day. Many also have spines to stab any animals that might
want to eat them and consume their delicious water.
• It makes sense, then, that cacti would also make use of the
CAM cycle to prevent them from opening their stomata and
losing water during the day!
Agave
• Agave –also uses CAM to survive in desert
environments.
• It looks more like a leafy green plant than a
cactus, but like cacti, it has developed thick flesh
to reduce its surface area and conserve water,
and spines along the edges of its leaves to
discourage animals from eating them.
 Benefits

• The most important benefit of CAM to the plant is the

ability to leave most leaf stomata closed during the day.
Plants employing CAM are most common in arid
environments, where water comes at a premium. Being
able to keep stomata closed during the hottest and driest
part of the day reduces the loss of water through
evapotranspiration, allowing such plants to grow in
environments that would otherwise be far too dry. Plants
using only C3 carbon fixation, for example, lose 97% of
the water they uptake through the roots to transpiration - a
high cost avoided by plants able to employ CAM.