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The Calvin cycle (or Calvin-Benson cycle or carbon fixation) is a series of biochemical reactions that takes place in the stroma of chloroplasts in photosynthetic organisms. It was discovered by Melvin Calvin and Andrew Benson at the University of California, Berkeley with James Bassham also contributing. It is one of the light-independent reactions or dark reactions.
The enzymes in the Calvin cycle are functionally equivalent to many enzymes used in other metabolic pathways such as gluconeogenesis and the pentose phosphate pathway, but they are to be found in the chloroplast stroma instead of the cell cytoplasm, separating the reactions. They are activated in the light (which is why the name "dark reaction" is misleading), and also by products of the light-dependent reaction. These regulatory functions prevent the Calvin cycle from operating in reverse to respiration, which would create a continuous cycle of carbon dioxide being reduced to carbohydrates, and carbohydrates being respired to carbon dioxide. Energy (in the form of ATP) would be wasted in carrying out these reactions that have no net productivity.
The sum of reactions in the Calvin cycle is the following:
It should be noted that hexose (six carbon) sugars are not a product of the Calvin cycle. Although many texts list a product of photosynthesis as C6H12O6, this is mainly a convenience to counter the equation of respiration, where six-carbon sugars are oxidized in mitochondria. The carbohydrate products of the Calvin Cycle are three-carbon sugar phosphate molecules, or "triose phosphates," specifically, glyceraldehyde-3-phosphate.
Steps of the Calvin cycle
(Simplified versions of the Calvin cycle integrate the remaining steps, except for the last one, into one general step - the regeneration of RuBP - also, one G3P would exit here.)
Thus, of 6 G3P produced, three RuBP (5C) are made totalling 15 carbons, with only one available for subsequent conversion to hexose. This required 9 ATPs and 6 NADPH per 3 CO2.
RuBisCO also reacts competitively with O2 instead of CO2 in photorespiration. The rate of photorespiration is higher at high temperatures. "photorespiration" turns RuBP into 3PGA and 2-phosphoglycolate, a 2-carbon molecule which can be converted via glycolate and glyoxalate to glycine. Via the glycine cleavage system and tetrahydrofolate, two glycines are converted into serine +CO2. Serine can be converted back to 3-phosphoglycerate. Thus, only 3 of 4 carbons from two phosphoglycolates can be converted back to 3PGA. Obviously photorespiration has very negative consequences for the plant, because rather than fixing CO2), this process leads to loss of CO2). C4 carbon fixation evolved to circumvent photorespiration, but can only occur in certain plants living in very warm or tropical climates.
Products of the Calvin cycle
The immediate product of the Calvin cycle is glyceraldehyde-3-phosphate (G3P) and water. Two G3P molecules (or one F6P molecule) that have exited the cycle are used to make larger carbohydrates. In simplified versions of the Calvin cycle they may be converted to F6P or F5P after exit, but this conversion is also part of the cycle.
Hexose isomerase converts about half of the F6P molecules into glucose-6-phosphate. These are dephosphorylated and the glucose can be used to form starch, which is stored in, for example, potatoes, or cellulose used to build up cell walls. Glucose, with fructose, forms sucrose, a non-reducing sugar which is a stable storage sugar, unlike glucose.
|This article is licensed under the GNU Free Documentation License. It uses material from the Wikipedia article "Calvin_cycle". A list of authors is available in Wikipedia.|