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Yoshida - 1981 - Fundamentals of Rice Crop Science

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196 FUNDAMENTALS OF RICE CROP SCIENCE<br />

This reaction does not require light and can proceed in darkness.<br />

Carbon dioxide is converted into carbohydrates during photosynthesis by two<br />

biochemical processes: C-3 and C-4 pathways (Hatch and Slack 1970, Bjorkman<br />

and Berry 1973). In the C-3 pathway, which is <strong>of</strong>ten called the Calvin Cycle,<br />

carbon dioxide is first incorporated into 3-phosphoglycerate (PGA):<br />

Note that the carbon atom in the CO 2 molecule (marked with an asterisk) is<br />

incorporated into the carboxyl group <strong>of</strong> one PGA molecule. In subsequent reactions,<br />

some PGA is converted into carbohydrates and some is used to regenerate<br />

RuDP molecules to serve as CO 2 acceptors. PGA is a compound with three carbon<br />

atoms per molecule; hence, the pathway is named the three-carbon photosynthetic<br />

pathway (C-3 pathway).<br />

In the C-4 pathway, CO 2 is first incorporated into compounds with four carbon<br />

atoms:<br />

+ P (5.5)<br />

CO 2 fixation and the formation <strong>of</strong> C-4 acids, malate and aspartate, occur in the<br />

mesophyll cells. The C-4 acids, transported to inner bundle sheath cells, are<br />

converted into CO 2 and pyruvate (C-3 compound):<br />

(5.6)<br />

The released CO 2 is fixed again in the Calvin cycle to produce PGA and carbohydrate.<br />

The pyruvate returns to the mesophyll cells, where it acquires a phosphate

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