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18 Oscillations in Plants 265<br />

Driessche 2000). Recent experiments in our laboratory have found a strong<br />

(R >0. 9) correlation between root circumnutation patterns and rhythmic<br />

changes in K + uptake in vertically grown maize roots (Shabala 2003), with<br />

periodical (t ∼ 50 min) K + flux changes from influx to efflux at two opposite<br />

sides in opposite phase.<br />

Plant circumnutations show strong dependence on temperature (Q 10 ∼ 2;<br />

Schuster and Engelmann 1997), light intensity (Anderson-Bernadas et al.<br />

1997) and quality (Schuster and Engelmann 1997), mechanical stimulation<br />

(Anderson-Bernadas et al. 1997) and chemical composition of the<br />

medium (Buer et al. 2000). All these observations point out the possibility<br />

of “encoding” environmental information by nutational patterns.<br />

The traditional view of the physiological role of plant nutations is that<br />

nutational growth of an organ maximises the penetration of the medium in<br />

which it grows (Barlow et al. 1994). Recently, Inoue et al. (1999) showed that<br />

in rice root circumnutation was important in creating a larger pushing force<br />

of the seminal root without causing floating of the seedling on flooded and<br />

very soft soil, thus leading to a higher seedling establishment percentage in<br />

the field. More direct evidence is needed to fully reveal the role of nutations<br />

in plant axial organs.<br />

18.2.2.4<br />

Root Nutrient Acquisition<br />

Despite the widely reported substantial diurnal changes in the rates of nutrient<br />

uptake by roots (reviewed by Shabala 2003), ultradian oscillations in<br />

root nutrient acquisitions remains underexplored. Nonetheless, such oscillations<br />

have been reported in several plant species (Knaritonashvili et al.<br />

1997; Macduff and Dhanoa 1996; Shabala et al. 1997; Shabala and Knowles<br />

2002; Shabala 2003), with periods ranging from 5 min to several hours.<br />

Importantly, such oscillations appear to be extremely sensitive to environmental<br />

conditions, with their periods showing a strong dependence on solution<br />

pH, temperature and osmolality (Shabala 2003). This may be strong<br />

evidence for a frequency-encoding mechanism operating in plant roots,<br />

similar to one reported for guard cells (McAinsh et al. 1995; Blatt 2000).<br />

18.2.2.5<br />

Growth<br />

There is increasing evidence that axial growth of many plant organs is also<br />

rhythmically modulated. Ultradian oscillations in stem growth with periods<br />

of several minutes were reported for stems, hypocotyls, roots and sporangiphores<br />

(see Shabala 2003 for references). Oscillations in membranetransport<br />

activity are likely to be the major driving force (Shabala et al.<br />

1997a; Tyerman et al. 2001). It has been suggested that periodical fluctua-

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