Cycles Calvin 248

The set of reactions that transforms 3-phosphoglycerate into hexose is named the Calvin-Benson cycle (often referred to simply as the Calvin cycle) for its discoverers. The reaction series is indeed cyclic because not only must carbohydrate appear as an end product, but the 5-carbon acceptor, RuBP, must be regenerated to provide for continual COg fixation. Balanced equations that schematically represent this situation are... 

Each number in parentheses represents the number of carbon atoms in a compound, and the number preceding the parentheses indicates the stoichiometry of the reaction. Thus, 6(1), or 6 COg, condense with 6(5) or 6 RuBP to give 12 3-phosphoglycerates. These 12(3)s are then rearranged in the Calvin cycle to form one hexose, 1 (6), and regenerate the six 5-carbon (RuBP) acceptors. 

The Calvin cycle enzymes serve three important ends ... 

Most of the enzymes mediating the reactions of the Calvin cycle also participate in either glycolysis (Chapter 19) or the pentose phosphate pathway (Chapter 23). The aim of the Calvin scheme is to account for hexose formation from 3-phosphoglycerate. In the course of this metabolic sequence, the NADPH and ATP produced in the light reactions are consumed, as indicated earlier in Equation (22.3). 

Balancing the Calvin Cycle Reactions To Account for Net Hexose Synthesis... 

FIGURE 22.27 Light-induced pH changes in chloroplast compartments. Illumination of chloroplasts leads to proton pumping and pH changes in the chloroplast, such that the pH within the thylakoid space falls and the pH of the stroma rises. These pH changes modulate the activity of key Calvin cycle enzymes. 

Compartmentation of these reactions to prevent photorespiration involves the interaction of two cell types, mescrphyll cells and bundle sheath cells. The meso-phyll cells take up COg at the leaf surface, where Og is abundant, and use it to carboxylate phosphoenolpyruvate to yield OAA in a reaction catalyzed by PEP carboxylase (Figure 22.30). This four-carbon dicarboxylic acid is then either reduced to malate by an NADPH-specific malate dehydrogenase or transaminated to give aspartate in the mesophyll cells. The 4-C COg carrier (malate or aspartate) then is transported to the bundle sheath cells, where it is decarboxylated to yield COg and a 3-C product. The COg is then fixed into organic carbon by the Calvin cycle localized within the bundle sheath cells, and the 3-C product is returned to the mesophyll cells, where it is reconverted to PEP in preparation to accept another COg (Figure 22.30). Plants that use the C-4 pathway are termed C4 plants, in contrast to those plants with the conventional pathway of COg uptake (C3 plants). 

Eormadon of PEP by pyruvate Pi dikinase reini-dates the cycle. The CO9 liberated in the bundle slieadi cell is used to syndiesize hexose by die convendonal rubisco-Calvin cycle series of reacdons. 

Write a balanced equation for the synthesis of a glucose molecule from ribulose-l,5-bisphosphate and COg that involves the first three reactions of the Calvin cycle and subsequent conversion of the two glyceraldehyde-3-P molecules into glucose. 

Photosynthesis in green plants occurs in two basic processes. In the dark (the Calvin cycle) carbon dioxide is reduced by a strong reducing agent, the reduced form of nicotinamide adeninedinucleotide phosphate, NADPH2, with the help of energy obtained from the conversion of ATP to ADP ... 

Fig. 6.3 The Calvin cycle or the dark reactions of photosynthesis see Cooper and also Stryer in Further Reading.

Much interest has recently been shown in artificial photosynthesis. Photosynthesis is a system for conversion or accumulation of energy. It is also interesting that some reactions occur simultaneously and continuously. Fujishima et al. [338] pointed out that a photocatalytic system resembles the process of photosynthesis in green plants. They described that there are three important parts of the overall process of photosynthesis (1) oxygen generation by the photolysis of water, (2) photophosphorylation, which accumulates energy, and (3) the Calvin cycle, which takes in and reduces carbon dioxide. The two reactions, reduction of C02 and generation of 02 from water, can occur simultaneously and continuously by a sonophotocatalytic reaction. 

Figure 38, Chapter 3. A bifurcation diagram for the model of the Calvin cycle with product and substrate saturation as global parameters. Left panel Upon variation of substrate and product saturation (as global parameter, set equalfor all irreversible reactions), the stable steady state is confined to a limited region in parameter space. All other parameters fixed to specific values (chosen randomly). Right panel Same as left panel, but with all other parameters sampled from their respective intervals. Shown is the percentage r of unstable models, with darker colors corresponding to a higher percentage of unstable models (see colorbar for numeric values).

Figure 39, Chapter 3. Bifurcation diagrams for the model of the Calvin cycle for selected parameters. All saturation parameters are fixed to specific values, and two parameters are varied. Shown is the number of real parts of eigenvalues larger than zero (color coded), with blank corresponding to the stable region. The stability of the steady state is either lost via a Hopf (HO), or via saddle node (SN) bifurcations, with either two or one eigenvalue crossing the imaginary axis, respectively. Intersections point to complex (quasiperiodic or chaotic) dynamics. See text for details.

Bistability and switching are crucial concepts of cellular regulation [80, 98] and can often be detected using the graphical methods outlined above. See also [98,287] for several illustrative examples. For a number of pathways, transitions between different states were observed, either in silico, in vivo, or both. Examples include the glycolytic pathway [273, 294], the Calvin cycle [113, 125], and models of the human erythrocytes [295, 296],... 

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