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376 E. Wagner et al.<br />

The circadian rhythmic cell (cyanobacterial and eukaryotic) is considered<br />

to be a hydro-electrochemical oscillator (Wagner et al. 1997) synchronised<br />

by the daily light–dark cycles, with temporal compartmentation of<br />

metabolism and a network of metabolic sequences to compensate for oxidative<br />

stress in adapting to the light environment, e.g. by separating nitrogen<br />

fixation from photosynthetic oxygen production (Sherameti et al. 2002).<br />

25.5<br />

Circadian Rhythmic Organisation of Energy<br />

Metabolism in C. rubrum and the Gating<br />

of Photoreceptor (Phytochrome) Action<br />

In C. rubrum a circadian rhythm in overall energy transduction has been<br />

observed. The rhythm results from an oscillatory network between glycolysis<br />

and oxidative phosphorylation, coupled to photophosphorylation. This<br />

network produces a circadian rhythm in adenylate energy charge and redox<br />

state (NADP/NADPH2). The nucleotide ratios themselves could act as<br />

rate effectors in compartmental feedback and thus fulfil the requirements<br />

of precise temperature-compensated time-keeping (Wagner et al. 1975,<br />

Table 25.2. Rhythmic organisation of metabolism in Chenopodium rubrum (the period<br />

lengths of the subpeaks are in parentheses) (Complemented after Bünning 1977)<br />

Phenomenon Period length (h)<br />

Photoperiodic light sensitivity 30<br />

Betacyanine accumulation 24–30 (15)<br />

Betacyanine turnover 24–30<br />

Adenylate kinase activity 30 (15)<br />

Energy charge (ATP + 1/2 ADP/ATP + ADP + AMP) 21–24 (11–13)<br />

NADPH2/NADP ratio 21–24<br />

Dark respiration 21–24<br />

Chlorophyll accumulation 15<br />

Net photosynthesis 15<br />

Triose phosphate dehydrogenase activity (NADH2;NADPH2) 15<br />

Malate dehydrogenase activity 12–15<br />

Glutamate dehydrogenase activity 12–15<br />

Glucose-6-phosphate dehydrogenase activity 12–15<br />

Gluconate-6-phosphate dehydrogenase activity 12–15<br />

Pyridine nucleotide, pool size [NAD(H2); NADP(H2)]<br />

Turgor-controlled growth phenomena<br />

12–15 (6)<br />

Stem extension rate 23–27<br />

Leaf movement 23–27

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