C3 Photosynthesis

Plants which use only the Calvin cycle for fixing the carbon dioxide from the air are known
as C3 plants. In the first step of the cycle CO2 reacts with RuBP to produce two 3-carbon
molecules of 3-phosphoglyceric acid (3-PGA). This is the origin of the designation C3 or C3
in the literature for the cycle and for the plants that use this cycle.

Index

Photosynthesis
Concepts

Reference
Moore, et al.
Ch 7

The entire process, from light energy capture to sugar production occurs within the
chloroplast. The light energy is captured by the non-cyclic electron transport process which
uses the thylakoid membranes for the required electron transport.

About 85% of plant species are C3 plants. They include the cereal grains: wheat, rice, barley,
oats. Peanuts, cotton, sugar beets, tobacco, spinach, soybeans, and most trees are C3 plants.
Most lawn grasses such as rye and fescue are C3 plants.

C3 plants have the disadvantage that in hot dry conditions their photosynthetic efficiency
suffers because of a process called photorespiration. When the CO2 concentration in the
chloroplasts drops below about 50 ppm, the catalyst rubisco that helps to fix carbon begins to
fix oxygen instead. This is highly wasteful of the energy that has been collected from the
light, and causes the rubisco to operate at perhaps a quarter of its maximal rate.

The problem of photorespiration is overcome in C4 plants by a two-stage strategy that keeps

CO2 high and oxygen low in the chloroplast where the Calvin cycle operates. The class of
plants called C3-C4 intermediates and the CAM plants also have better strategies than C3
plants for the avoidance of photorespiration.

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C4 Photosynthesis
Sugarcane is a
champion at
photosynthesis
under the right
conditions and
is a prime
example of a
C4 plant, one
which uses C4
photosynthesis.
Sugarcane has
been recorded
at 7%
photosynthetic
efficiency.

C4 plants almost never saturate with light and under hot, dry conditions much outperform C3
plants. They use a two-stage process were CO2 is fixed in thin-walled mesophyll cells to form a
4-carbon intermediate, typically malate (malic acid). The reaction involves phosphoenol
pyruvate (PEP) which fixes CO2 in a reaction catalyzed by PEP-carboxylate. It forms
oxaloacetic acid (OAA) which is quickly converted to malic acid. The 4-carbon acid is actively
pumped across the cell membrane into a thick-walled bundle sheath cell where it is split to CO2
and a 3-carbon compound.

This CO2 then enters the Calvin cycle in a chloroplast of the bundle sheath cell and produces
G3P and subsequently sucrose, starch and other carbohydrates that enter the cells energy
transport system.

Index

Photosynthesis
Concepts

Reference
Moore, et al.
Ch 7

The advantage that comes from this two-stage process is that the active pumping of carbon into
the bundle sheath cell and the blocking of oxygen produce an environment with 10-120x as
much CO2 available to the Calvin cycle and the rubisco tends to be optimally utilized. The high
CO2 concentration and the absence of oxygen implies that the system never experiences the
detractive effects of photorespiration.

The drawback to C4 photosynthesis is the extra energy in the form of ATP that is used to pump
the 4-carbon acids to the bundle sheath cell and the pumping of the 3-carbon compound back to
the mesophyll cell for conversion to PEP. This loss to the system is why C3 plants will
outperform C4 plants if there is a lot of water and sun. The C4 plants make some of that energy
back in the fact that the rubisco is optimally used and the plant has to spend less energy
synthesizing rubisco.

Moore, et al. say that only about 0.4% of the 260,000 known species of plants are C4 plants.
But that small percentage includes the important food crops corn, sorghum, sugarcane and
millet. Also inluded are crabgrass and bermuda. Many tropical grasses and sedges are C4
plants.

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C3-C4 Intermediate Photosynthesis

Moore, et al. point to Flaveria (Asteraceae), Panicum (Poaceae) and Alternanthera
(Amarantheceae) as genera that contain species that are intermediates between C3 and
C4 photosynthesis. These plants have intermediate leaf anatomies that contain bundle
sheath cells that are less distinct and developed than the C4 plants.

These intermediates are

characterized by their
resistance to
photorespiration so that they
can operate in higher
temperatures and dryer
environments than C3 Index
plants. At right, the ranges
of CO2 compensation points Photosynthesis
Concepts
for the three types of plants
are shown. These
Reference
compensation points are the
Moore, et al.
values at which the plants
Ch 7
cease to provide net
photosynthesis.

The connection to hot and dry conditions comes from the fact that all the plants will
close their stomata in hot and dry weather to conserve moisture, and the continuing
fixation of carbon from the air drops the CO2 dramatically from the atmospheric
concentration of nominally 380 ppm (2004 value). If the CO2 compensation point is
lower on the above scale, the plant can operate in hotter and dryer conditions. The
limits are placed by the fact that rubisco begins to fix oxygen rather than CO2, undoing
the work of photosynthesis. C4 plants shield their rubisco from the oxygen, so can
operate all the way down to essentially zero CO2 without the onset of photorespiration.

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Crassulacean Acid Metabolism (CAM)

The CAM plants represent a metabolic strategy adapted

to extremely hot and dry environments. They represent
about 10% of the plant species and include cacti,
orchids, maternity plant, wax plant, pineapple, Spanish
moss, and some ferns. The only agriculturally
significant CAM plants are the pineapple and an Agave
species used to make tequila and as a source of fiber.

The sketch below of the day-night cycle of the CAM

plants is patterned after Moore, et al. The name
Crassulacean Acid Metabolism came from the fact that
this strategy was discovered in a member of the
Crassulaceae which was observed to become very
acidic at night and progressively more basic during the
day.

Index

Photosynthesis
Concepts

Reference
Moore, et al.
Ch 7

The acidity was found to arise from the opening of their stomata at night to take in CO2
and fix it into malic acid for storage in the large vacuoles of their photosynthetic cells.
It could drop the pH to 4 with a malic acid concentration up to 0.3M . Then in the heat
of the day, the stomata close tightly to conserve water and the malic acid is
decarboxylated to release the CO2 for fixing by the Calvin cycle. PEP is used for the
initial short-term carbon fixation as in the C4 plants, but the entire chain of reactions
occurs in the same cell rather than handing off to a separate cell as with the C4 plants.
In the CAM strategy, the processes are separated temporally, the initial CO2 fixation at
night, and the malic acid to Calvin cycle part taking place during the day.

With stomata open only at night when the

temperature is lower and the relative
humidity higher, the CAM plants use
much less water than either C3 plants or
C4 plants. Some varieties convert to C3
plants at the end of the day when their
acid stores are depleted if they have
adequate water, and even at other times
when water is abundant.

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Photorespiration
Respiration refers to the metabolism of oxygen and the release of carbon dioxide. In cellular
respiration it is a positive term, a process vital to life. But photorespiration is an entirely
negative term because it represents a severe loss to the process of using light energy in
photosynthetic organisms to fix carbon for subsequent carbohydrate synthesis. By leading to
the loss of up to half of the carbon that has been fixed at the expense of light energy,
photorespiration undoes the work of photosynthesis.

Photorespiration happens in C3 plants when the CO2 concentration drops to about 50 ppm.
The key enzyme that accomplishes the fixing of carbon is rubisco, and at low concentrations
of CO2 it begins to fix oxygen instead.

Index

Photosynthesis
Concepts

Reference
Moore, et al.
Ch 7

Under moderate temperature conditions when C3 plants have sufficient water, the supply of
carbon dioxide is abundant and photorespiration is not a problem. The CO2 concentration of
the atmosphere as of 2004 was about 380 ppm and this CO2 freely diffuses through the
stomata of leaves and across the membranes of the chloroplasts while water diffuses out
through the stomata. But during hot and dry conditions, the stomata close to prevent excessive
water loss and the continuing fixation of carbon in the Calvin cycle dramatically reduces the
relative concentration of CO2. When it reaches a critical level of about 50 ppm the rubisco
stops fixing CO2 and begins to fix O2 instead. Even though the detoured process feeds some
PGA back into the cycle, the photorespiration process causes rubisco to operate at only about
25% of its optimal rate.

The C4 plants and CAM plants avoid photorespiration and therefore operate at much higher
efficiencies in hot and dry climates.

Photorespiration wiki
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