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222 S.J. Roux, C. Song, C. Jeter<br />

7-pass TM heterotrimeric G-protein linked receptors. The binding of ATP<br />

or ADP (among other NTPs and NDPs) activates these receptors, initiating<br />

secondary messenger systems and downstream signaling cascades, thereby<br />

affecting changes in gene expression and culminating in the induction of<br />

cell-type specific responses.<br />

The first recognized physiological activity of these signaling agents was<br />

that of a co-neurotransmitter, and so this type of signaling was originally<br />

known as purinergic transmission (Burnstock 1972). Derivatives of ATP,<br />

including ADP and adenosine, were also shown to have biological effects.<br />

For example, adenosine functions as a negative regulator of neurotransmitter<br />

release at specialized P1 purinceptors, functioning together with ATP<br />

to modulate smooth muscle contraction; and ADP signals blood platelet<br />

aggregation during thrombosis (reviewed in Burnstock 1996; Ralevic and<br />

Burnstock 1998).<br />

Extracellular ATP (eATP) has been reported to have numerous effects<br />

on the physiology of plants, too, altering both developmental programs<br />

andresponsestoenvironmentalstimuli.Earlystudiesshowedthatexogenous<br />

application of ATP could induce the closure of the Venus’s-flytrap<br />

(Jaffe 1973), affect cytoplasmic streaming in Chara cells (Williamson 1975),<br />

modulate stomatal aperture in Commelina communis (Nejidat et al. 1983),<br />

and stimulate pollen tube generative nuclear divisions in Lilium lingiflorum<br />

(Kamizyo and Tanaka 1982). However, most of these reports assumed that<br />

the applied ATP was somehow, directly or indirectly, altering the energy<br />

charge of the cell, and thus was still playing its standard role of driving<br />

energy-dependent reactions.<br />

More recent results have revealed that the hydrolysis of eATP is not<br />

required to induce responses in plant cells (Demidchik et al. 2003; Jeter<br />

et al. 2004). This has led plant scientists to recognize the potential role<br />

of eATP as an agonist, exerting its effects through interaction with cell<br />

surface receptors, similar to what happens in animal cells. The purpose of<br />

this chapter is to review this more recent literature and discuss potential<br />

mechanisms by which extracellular nucleotides could alter plant growth<br />

and development as cell–cell signaling molecules.<br />

15.2<br />

Rapid Responses of Plants to Applied Nucleotides<br />

15.2.1<br />

Induced Changes in the Concentration of Cytoplasmic Calcium Ions<br />

In animal cells, activation of purinoceptors by extracellular nucleotides<br />

rapidly leads to changes in membrane potential and increases in the con-

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