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18 Oscillations in Plants 269<br />

The foundations of the systems theory suggestthatiftherelationship<br />

between several parameters within the system is described by linear equations,<br />

such a system should eventually reach its stable state (Stucki and<br />

Somogyi 1994). Once disturbed, such a system will eventually return to<br />

a new steady state through a series of damped oscillations. This is often observed<br />

for plant photosynthetic responses (Kocks and Ross 1995), changes<br />

in stomatal aperture (Barrs 1971) or cell electrophysiological characteristics<br />

(Tyerman et al. 2001).<br />

The story is quite different if the system is governed by non-linear mechanisms.<br />

In that case, a limited cycle (a two-dimensional attractor), rather<br />

than a singular point, will be a stable condition (Stucki and Somogyi 1994).<br />

Thus,self-sustainedoscillationsareexpectedtobefoundinsuchnon-linear<br />

systems.<br />

Thereisnodoubtthatmostphysiologicalprocessesinplantsaregoverned<br />

by non-linear mechanisms. Physiologically it means that within some<br />

narrow range of parameters (light intensity, ambient temperature, water<br />

and nutrient availability, etc.), plant responses might be linear. In this<br />

case, a sudden perturbation within this range will cause only a brief series<br />

of damped oscillations in plant physiological responses. As soon as<br />

the disturbance is beyond the range of the linear response, non-damping<br />

self-sustained oscillations are expected.<br />

18.3.2<br />

Advantages of Oscillatory Strategy<br />

Every plant physiologist will probably agree that circadian rhythmicity<br />

in plants is a result of evolutionary adaptation of energy metabolism to<br />

optimise energy conservation with respect to daily environmental cycles of<br />

energy supply (Wagner et al. 1975). Researchers are less in agreement when<br />

discussing the functional role of ultradian oscillations in plants. Although<br />

there is no lack of theoretical investigations (see later) showing advantages<br />

of oscillatory strategy, direct evidence for plants is still rather rudimentary.<br />

Earlier Rapp (1987) described at least five positive functional advantages<br />

which periodic behaviour confers to living organisms. These are revisited<br />

next.<br />

18.3.2.1<br />

Temporal Organisation<br />

Separate temporal compartments may be necessary where mutually incompatible<br />

biochemical reactions occur in an identical spatial (subcellular)<br />

compartment. Examples may include protein synthesis and degradation.

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