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18 Oscillations in Plants 267<br />

background with a period ranging from a few seconds to 10 min (Pearcy<br />

1990). On a larger scale, light fluctuations caused by cloud movements<br />

influence all plants growing in a particular area (Cardon et al. 1994), resulting<br />

in light fluctuations in the minute range of periods. It is not surprising,<br />

therefore, that leaf photosynthetic machinery is adjusted to such a “fluctuating”<br />

regime.<br />

Oscillations in photosynthesis are well-reported phenomena (Kocks and<br />

Ross 1995; Lüttge and Hütt 2004) and have been found at various levels of<br />

organisation. Rhythmical changes in chlorophyll a fluorescence, phosphorylation,<br />

oxygen evolution and CO2 assimilation are all examples of such<br />

oscillatory behaviour. The period range for such oscillations is usually<br />

around several minutes (Siebke et al. 1992). In most cases, such oscillations<br />

are strongly damped and, being induced by sudden changes in one<br />

of the environmental variables (light, CO2, etc.), disappear after several cycles<br />

(Siebke et al. 1992; Kocks and Ross 1995). However, non-damped (for<br />

several hours) oscillations have also been reported (Siebke and Weis 1995).<br />

18.2.2.8<br />

Osmotic Adjustment<br />

As discussed before, both leaf and axial organ movements are mediated<br />

by turgor and volume changes in the epidermal (in the case of nutations)<br />

or pulvini (in the case of leaf movement) cells. In addition, oscillatory<br />

ion transport mechanisms were shown to operate in planktonic diatoms<br />

for adjustment of buoyancy by appropriate uptake and release of ions<br />

(Gradmann and Boyd 1995). In higher plants, rapid (1−2 min period) cycles<br />

of K + uptake and release in osmotically stressed leaf mesophyll cells were<br />

reported (Shabala et al. 2000). All these facts point out the possibility of the<br />

“fine-tuning” mechanisms of osmotic adjustment being realised through<br />

oscillatory ion uptake across the plasma membrane.<br />

18.2.2.9<br />

Ultradian Rhythms in Time-Keeping<br />

Molecular and genetics aspects of circadian rhythms have been the subject<br />

of recent reviews (Webb 2003). Transcriptional/translational genetics<br />

models are favoured (Dunlap 1998). Several clock genes have been identified<br />

(Webb 2003). However, it appears that circadian systems will almost<br />

certainly be made up of more than one interconnected feedback loop, and<br />

there are many discomforting facts for the current genetics model. Dunlap<br />

(1998) himself called it as a “pleasing caricature of reality”.<br />

One of the striking features of oscillations in plants is the coexistence<br />

of ultradian and circadian modulations of the same physiological process.<br />

Can ultradian rhythms be a part of the circadian clock mechanism?

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