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24 Electrophysiology and Phototropism 353<br />

The huge amount of experimental material testifies that the main laws<br />

of excitability such as the inducement of nonexcitability after excitation<br />

and the summation of subthreshold irritations were developed in the vegetative<br />

and animal kingdoms in protoplasmatic structures earlier than the<br />

morphological differentiation of nervous tissues (Volkov 2000). These protoplasmatic<br />

excitable structures consolidated into the organs of a nervous<br />

system and adjusted the interaction of the organism with the environment.<br />

Volkov and Haack (1995) studied the role of electrical signals induced by<br />

insects in long-distance communication in plants and confirmed the mechanism<br />

by which electrical signals can directly influence both biophysical<br />

and biochemical processes in remote tissues.<br />

Bose (1925) has established the availability of a reflex arch in plants<br />

such as Mimosa pudica. When plants are excited sensory cells generate<br />

impulses which terminate at motor cells. The character of their distribution<br />

depends upon the physiological condition of plants. The signal<br />

from a beam of the sun is transmitted to the tissues of a stem at an<br />

extremely high speed and consequently the stem will curve toward the<br />

source of light. After excitation, the illuminated top of a stem causes an<br />

impulse to be distributed among the tissues (Volkov et al. 2005). When<br />

the impulse reaches motor cells, the stem bends. Thus, after electrochemical<br />

signals have reached the cell, deep cytophysiological reactions occur.<br />

24.2<br />

Phototropism and Photosensors<br />

Lightisanessentialsourceofenergyonwhichmanyofthebiologicalfunctions<br />

of plants depend. The sun’s radiant energy optimizes germination,<br />

photosynthesis, flowering, and other processes needed to maintain homeostasis.<br />

The first experiment on phototropism is probably lost in antiquity.<br />

Most likely someone noticed an indoor potted plant bending toward a window<br />

and rotated the pot 180 ◦ and then noticed later that the plant again<br />

bentbacktowardthewindow.Thefirstscientiststoreallymakeprogress<br />

in explaining phototropism were Charles and Francis Darwin (1888). They<br />

grew canary grass seedling and placed little caps of tinfoil or this glass<br />

painted black on the tips of the coleoptiles and determined what portion<br />

of the coleoptile had to receive light in order for phototropism to occur.<br />

Darwin (1888) found the tip was the light-sensitive part, but the bending<br />

also occurred well below the tip. Darwin (1888) hypothesized that an<br />

“influence” was translocated from the illuminated tip to the part where<br />

bending occurred.

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