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Photosynthesis proceeds in two parts: the light reactions and the dark reactions. The light reactions, discussed in Chapter 19, transform light energy into ATP and biosynthetic reducing power, NADPH. The dark reactions use the ATP and NADPH produced by the light reactions to reduce carbon atoms from their fully oxidized state as carbon dioxide to a more reduced state as a hexose. Carbon dioxide is thereby trapped in a form that is useful for many processes and most especially as a fuel. Together, the light reactions and dark reactions of photosynthesis cooperate to transform light energy into carbon fuel. The dark reactions are called either the Calvin–Benson cycle, after Melvin Calvin and Andrew Benson, the biochemists who elucidated the pathway, or simply the Calvin cycle. The components of the Calvin cycle are called the dark reactions because, in contrast with the light reactions, these reactions do not directly depend on the presence of light.

The second half of this chapter examines a pathway common to all organisms, known variously as the pentose phosphate pathway, the hexose monophosphate pathway, the phosphogluconate pathway, or the pentose shunt. The pathway provides a means by which glucose can be oxidized to generate NADPH, the currency of readily available reducing power in cells. The phosphoryl group on the 2′-hydroxyl group of one of the ribose units of NADPH distinguishes NADPH from NADH. There is a fundamental distinction between NADPH and NADH in biochemistry: NADH is oxidized by the respiratory chain to generate ATP, whereas NADPH serves as a reductant in bio-synthetic processes. The pentose phosphate pathway can also be used for the catabolism of pentose sugars from the diet, the synthesis of pentose sugars for nucleotide biosynthesis, and the catabolism and synthesis of less common four- and seven-carbon sugars. The pentose phosphate pathway and the Calvin cycle have in common several enzymes and intermediates that attest to an evolutionary kinship. Like glycolysis and gluconeogenesis, these pathways are mirror images of each other: the Calvin cycle uses NADPH to reduce carbon dioxide to generate hexoses, whereas the pentose phosphate pathway oxidizes glucose to carbon dioxide to generate NADPH.

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