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18 Oscillations in Plants 271<br />

prehensively reviewed by Lüttge and Hütt 2004). Termed as “deterministic<br />

chaos”, such behaviour is routine in many physical and hydrodynamics<br />

systems. It still remains unclear, however, to what extent biological systems<br />

functionally exploit this behaviour (Lüttge and Hütt 2004).<br />

Experimental evidence on chaos in plants is only gradually accumulating.<br />

The period doubling of leaf electric and temperature oscillations in<br />

response to rhythmical light (Shabala et al. 1986, 1997b), aperiodic leaflet<br />

movement reminiscent of homoclinic chaos (Chen et al. 1995) and chaotic<br />

behaviour in CO2 exchange (Lüttge and Beck 1992) were reported. Chaotic<br />

oscillations in the plant cell expansion rate were predicted by Kellershohn et<br />

al. (1996), and bifurcational regimes in stomatal oscillations were modelled<br />

by Rand et al. (1981).<br />

Such a dearth of experimental evidence of deterministic chaos in plants<br />

is more than surprising. Biological systems may gain benefits from exhibiting<br />

chaotic dynamics as it may contribute to the generation of diversity<br />

and hence adaptability (Lüttge and Hütt 2004). Advantages of chaotic dynamics<br />

were recently discussed (Lloyd 1997) and include greater flexibility<br />

in response towards external influences, phenotypic diversity, larger functional<br />

independence from external entrainment and higher dissipation of<br />

disturbance. Overall, exploitation of these benefits ensures the evolutionary<br />

survival of chaotic dynamics; thus, it can make “chaotic” behaviour in<br />

plants physiologically important.<br />

18.3.4<br />

Resonant Regimes<br />

The Dictionary of Physics defines resonance as a “condition in which a vibrating<br />

system responds with maximum amplitude to an alternating driving<br />

force”. Resonance phenomena are widely used in various areas of modern<br />

life (engineering, medicine, etc.). Keeping in mind the wide range of<br />

oscillatory activity in plants, we have to answer two questions: (1) are resonant<br />

responses possible in plants? and (2) will such behaviour be beneficial<br />

to plants?<br />

The answer to both questions is “yes”. There is no shortage of theoretical<br />

models predicting resonant plant responses to periodical disturbance.<br />

Examples include photosynthetic responses (Kocks and Ross 1995), membrane<br />

transport activity (Markevich and Sel’kov 1986; Tsong 1990) and<br />

energy transduction in glycolysis (Termonia and Ross 1982). These model<br />

predictions are further confirmed by experimental studies showing the<br />

resonant type of responses in stomatal conductance, water uptake and leaf<br />

surface electric potential to rhythmical light (Shabala 1989, 1997; Cardon<br />

et al. 1994) or root medium environment (Shabala et al. 1991). Preliminary

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