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25 Hydro-Electrochemical Integration of the Higher Plant 377<br />

Energy Charge [ NADPH 2/NADP ]<br />

7<br />

6<br />

5<br />

4<br />

3<br />

2<br />

1<br />

0<br />

0.65<br />

0.56<br />

0<br />

4.5 days 24 h various hours (0 - 72)<br />

32.5°C / 10°C<br />

20°C 20°C<br />

12 24 36 48<br />

Darkness [h]<br />

[NADPH 2 / NADP]<br />

[ATP] + 0.5 [ADP]<br />

[ATP] + [ADP] + [AMP]<br />

60 72<br />

Fig.25.4. Circadian rhythm of NADPH2-to-NADP ratio and of “energy charge” of C. rubrum<br />

seedlings. The seedlings were germinated in constant light and alternating conditions of<br />

temperature for 4.5 days, thereafter transferred to constant 20 ◦ C for 24 h followed by a dark<br />

period of varied duration (3-h increments) (Wagner et al. 1975)<br />

1983, 1998). In C. rubrum there are semicircadian oscillations with specific<br />

phasing in enzyme activities involved in glycolysis, photosynthesis and<br />

respiration resulting in a circadian rhythm of adenylate energy charge and<br />

redox state (Table 25.2, Fig. 25.4).<br />

There is an inverse phase relationship between glyceraldehyde-3-phosphate<br />

dehydrogenase (GPD) linked to photosynthesis (NADP-GPD) and the<br />

enzyme linked to glycolysis (NAD-GDP) (Fig. 25.5). Adenylate kinase, a key<br />

enzyme in energy dissipation displays circadian/semicircadian oscillations<br />

in activity. The enzyme exists as isoenzymes in chloroplasts, mitochondria,<br />

thecytoplasmandthenucleus.Theenzymeactivitiesaremodulatedby<br />

light (phytochrome, cryptochrome), glucose feeding and photoperiod and<br />

thermoperiod (Figs. 25.6–25.8).<br />

The rhythm in enzyme activities may be amplitude-modulated by phytochrome.<br />

The phytochrome action is gated by the phasing of the endogenous<br />

rhythm reflecting a sequence of photophile and skotophile phases as<br />

conceived by Bünning (1977) for the functioning of the physiological clock<br />

(Figs. 25.9 and 25.10).

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