Calvin and his co-workers then worked out the whole pathway and showed that the pathway operated in a cyclic manner; the RuBP was regenerated. Let us now see how the Calvin pathway operates and where the sugar is synthesised. Let us at the outset understand very clearly that the Calvin pathway occurs in all photosynthetic plants; it does not matter whether they have C3 or C4 (or any other) pathways (Figure 13.8).
For ease of understanding, the Calvin cycle can be described under three stages: carboxylation, reduction and regeneration.
1. Carboxylation – Carboxylation is the fixation of CO2 into a stable organic intermediate. Carboxylation is the most crucial step of the Calvin cycle where CO2 is utilised for the carboxylation of RuBP. This reaction is catalysed by the enzyme RuBP carboxylase which results in the formation of two molecules of 3-PGA. Since this enzyme also has an oxygenation activity it would be more correct to call it RuBP carboxylase-oxygenase or RuBisCO.
Figure 13.8 The Calvin cycle proceeds in three stages : (1) carboxylation, during which CO2 combines with ribulose-1,5-bisphosphate; (2) reduction, during which carbohydrate is formed at the expense of the photochemically made ATP and NADPH; and (3) regeneration during which the CO2 acceptor ribulose-1,5-bisphosphate is formed again so that the cycle continues
2. Reduction – These are a series of reactions that lead to the formation of glucose. The steps involve utilisation of 2 molecules of ATP for phosphorylation and two of NADPH for reduction per CO2 molecule fixed. The fixation of six molecules of CO2 and 6 turns of the cycle are required for the formation of one molecule of glucose from the pathway.
3. Regeneration – Regeneration of the CO2 acceptor molecule RuBP is crucial if the cycle is to continue uninterrupted. The regeneration steps require one ATP for phosphorylation to form RuBP.
Hence for every CO2 molecule entering the Calvin cycle, 3 molecules of ATP and 2 of NADPH are required. It is probably to meet this difference in number of ATP and NADPH used in the dark reaction that the cyclic phosphorylation takes place.
To make one molecule of glucose 6 turns of the cycle are required. Work out how many ATP and NADPH molecules will be required to make one molecule of glucose through the Calvin pathway.
It might help you to understand all of this if we look at what goes in and what comes out of the Calvin cycle.