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26 Signals and Signalling Pathways in Plant Wound Responses 397<br />

spondingly were not active importers via the phloem. These small wounds<br />

were not accompanied by electrical activity (Table 26.1). Steam girdling<br />

the petiole of leaf 1 prevented the systemic induction of PI activity by<br />

a small mechanical wound to that leaf (Table 26.1). These results are consistent<br />

with the report (Nelson et al. 1983) from Ryan’s group that systemic<br />

signalling was prevented by prior hot air (80 ◦ C) treatment of the petiole<br />

of the leaf that was to be wounded; the wounding protocol used by that<br />

group is, in our view, similar to wounding method 3, i.e. a small mechanical<br />

wound. We conclude that for small mechanical wounds the systemic<br />

signal is a chemical elicitor that is exported from the wounded leaf in the<br />

phloem.<br />

Electrical events are associated only with severe wounds. Malone (1996)<br />

has provided evidence, on several grounds, that such electrical events are<br />

likely to be associated with hydraulic dispersal. On this basis, and from<br />

our evidence presented earlier for the similarities of the patterns of hydraulic<br />

dispersal (Fig. 26.1) and electrical activity, we conclude that the<br />

electrical events could be responses to chemicals transported in the xylem<br />

by hydraulic dispersal from the wound site, rather than action potentials<br />

propagated from that site. This is contrary to the conclusion in an earlier<br />

paper reporting work from our laboratory in which we used only severe<br />

wounds (Wildon et al. 1992); see also Davies (2004).<br />

The conclusions here raise interesting questions about the results reported<br />

in our previous paper (Rhodes et al. 1996), which was the most<br />

detailed study to date of electrical events at the cell level in the wounded<br />

plant. In that work we used intracellular recording from all the cell types in<br />

the petiole of unwounded leaf 1 to further characterise electrical events following<br />

a severe wound (heat, wound-type 5) to cotyledon 1. The recording<br />

setup is shown in Fig. 26.2.<br />

Fig.26.2. Schematic diagram of the arrangement of electrodes on the petiole of leaf 1 of<br />

a tomato seedling, used to record wound-induced electrical events. Each plant was wounded,<br />

using heat produced by passing an electric current for 30 s through a small wire element<br />

held over, but not touching, cotyledon 1 (Reproduced from Rhodes et al. 1996, with the<br />

permission of Springer-Verlag)

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