Carbon dioxide fixing, rubisco enzyme

Fig. 6.3. Carbon dioxide-fixing pathway in the C4-plants (prepared mainly on the basis of Hatch et al., 1967). Circled numbers 1, phosphoenolpyruvate carboxylase 2, malate dehydrogenase (NADP ) 3, malate dehydrogenase (NADP ) (OAA decarboxylating) (= malic enzyme) 4, Rubisco 5, pyruvate orthophosphate dikinase. Pi, phosphate PPi, diphosphate...

The dark reactions (carbon-fixation reactions) use the ATP and NADPH produced by the light reactions to fix carbon dioxide as carbohydrate sucrose and starch. The reactions form a cycle (the Calvin cycle) in which the enzyme ribulose bisphosphate carboxylase (rubisco), located in the stroma, condenses a C02 molecule with ribulose 1,5-bisphosphate to produce two molecules of 3-phosphoglycerate. Other reactions then regenerate the ribulose... 

Among the proteins found in the chloroplast stroma are the enzymes of the Calvin cycle, which functions in fixing carbon dioxide into carbohydrates during photosynthesis (Chapter 8). The large (L) subunit of ribulose 1,5-bisphosphate carboxylase (rubisco) is encoded by chloroplast DNA and synthesized on chloroplast ribosomes in the stromal space. The small (S) subunit of rubisco and all the other Calvin cycle enzymes are encoded by nuclear genes and transported to chloroplasts after their synthesis in the cytosol. The precursor forms of these stromal proteins contain an N-terminal stromal-import sequence (see Table 16-1). 

The fundamental basis of photosynthetic carbon metabolism is the incorporation of carbon dioxide by ribulose-bisphosphate carboxylase (rubisco). This leads to the synthesis of three-carbon sugars which are either exported from the chloroplast or metabolized to regenerate the acceptor ribulose bisphosphate. Rubisco is a bifunctional enzyme in that, in parallel to carboxylation, it catalyzes an oxygenation reaction that leads to phospho-glycolate. This is the starting point for photorespiratory metabolism, which will be discussed below (Section 1.6.2). In C4 plants, the conventional C3 pattern of the photosynthetic carbon reduction Calvin cycle is confined to the bundle sheath cells. The surrounding mesophyll cells act as an ancillary carbon dioxide pump, fixing carbon dioxide via phosphoenolpyruvate carboxylase into C4 acids. These are transported to the bundle sheath for decarboxylation.In this way, photorespiration is limited because of the elevated carbon dioxide levels. 

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