In plants, carbon dioxide (
) enters the chloroplast through
the stomata and diffuses into the stroma of the chloroplast—the site of the Calvin cycle reactions where sugar is synthesized. The reactions are named after the scientist who discovered them,
and reference the fact that the reactions function as a cycle. Others call it the Calvin-Benson cycle to include the name of another scientist involved in its discovery (Figure 5.14).
The Calvin cycle reactions (Figure 5.14 ) can be organized into three
basic stages: fixation, reduction, and regeneration. In the stroma, in addition to 
, two other
chemicals are present to initiate the Calvin cycle: an enzyme abbreviated RuBisCO, and the molecule ribulose bisphosphate (RuBP). RuBP has five atoms of carbon and a phosphate group on each
end.
RuBisCO catalyzes a reaction between 
and RuBP, which forms a six-carbon compound that is immediately converted
into two three-carbon compounds. This process is called carbon fixation, because 
is “fixed” from its inorganic form into organic molecules.
ATP and NADPH use their stored energy to convert the three-carbon compound, 3-PGA, into another three-carbon compound called G3P. This type of reaction is called a reduction reaction, because
it involves the gain of electrons. A reduction is the gain of an electron by an atom or molecule. The molecules of ADP and 
, resulting from the reduction reaction, return to the light-dependent reactions to be re-energized.
One of the G3P molecules leaves the Calvin cycle to contribute to the formation of the carbohydrate molecule, which is commonly glucose (
). Because the carbohydrate molecule has six carbon atoms, it takes six turns of the
Calvin cycle to make one carbohydrate molecule (one for each carbon dioxide molecule fixed). The remaining G3P molecules regenerate RuBP, which enables the system to prepare for the
carbon-fixation step. ATP is also used in the regeneration of RuBP.
In summary, it takes six turns of the Calvin cycle to fix six carbon atoms from 
. These six
turns require energy input from 12 ATP molecules and 12 NADPH molecules in the reduction step and 6 ATP molecules in the regeneration step.
The following is a link (http://openstaxcollege.org/l/calvin_cycle2) to an animation of the Calvin cycle. Click Stage 1, Stage 2, and then Stage 3 to see G3P and ATP regenerate to form RuBP.
Evolution In Action
Photosynthesis
The shared evolutionary history of all photosynthetic organisms is conspicuous, as the basic process has changed little over eras of time. Even between the giant tropical leaves in the rainforest and tiny cyanobacteria, the process and components of photosynthesis that use water as an electron donor remain largely the same. Photosystems function to absorb light and use electron transport chains to convert energy. The Calvin cycle reactions assemble carbohydrate molecules with this energy.
However, as with all biochemical pathways, a variety of conditions leads to varied adaptations that affect the basic pattern. Photosynthesis in dry-climate plants (Figure 5.16) has evolved with adaptations that conserve water. In the harsh dry
heat, every drop of water and precious energy must be used to survive. Two adaptations have evolved in such plants. In one form, a more efficient use of 
allows plants to photosynthesize even when 
is in short supply, as when the stomata are closed on hot days. The other
adaptation performs preliminary reactions of the Calvin cycle at night, because opening the stomata at this time conserves water due to cooler temperatures. In addition, this adaptation has
allowed plants to carry out low levels of photosynthesis without opening stomata at all, an extreme mechanism to face extremely dry periods.
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