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400 J.D.Rhodes,J.F.Thain,D.C.Wildon<br />

rations of the electrical events recorded by the surface and bath electrodes<br />

would normally lead to their classification as variation potentials with the<br />

implication that they reflect depolarisations in the underlying tissues due<br />

to chemicals that have travelled in the xylem from the wound site. However,<br />

given our current ignorance of detailed mechanisms at the cell level,<br />

they could reflect the summation of action potentials travelling at slightly<br />

different times in the different STE/CC complexes of the petiole.<br />

26.3<br />

Conclusions and Future Prospects<br />

For severe wounding, the results are consistent with the distribution of<br />

elicitors of PI synthesis and electrical activity by hydraulic dispersal in the<br />

xylem. We conclude that the electrical events are likely to be responses to<br />

chemicals transported in the xylem by hydraulic dispersal from the wound<br />

site, rather than action potentials propagated from the wound site.<br />

Following a small crushing wound, the pattern of systemic PI synthesis<br />

was consistent with the transport of a chemical elicitor in the phloem from<br />

the wound site.<br />

The lack of an electrical event following a small wound indicates that<br />

electrical signals are not an essential part of the systemic signalling system<br />

that induces PI synthesis.<br />

The fact that a severe wound leads to large rapid action-potential-like<br />

depolarisations in the cells of the STE/CC complexes in the petiole, but not<br />

in other cell types, is intriguing. Do these depolarisations have a function?<br />

In the absence of detailed studies at the cell level, the description of<br />

wound-induced electrical events recorded with surface-contact electrodes<br />

as either variation potentials or action potentials could carry misleading<br />

implications about their cellular mechanisms.<br />

A significant step forward in plant signalling would be the identification<br />

of the cellular mechanism of wound-induced electrical events; this could be<br />

achieved using a similar approach to that described here and in Rhodes et<br />

al. (1996). However, since the mapping of its genome, Arabidopsis may be<br />

the preferred model organism; Arabidopsis does show electrical events following<br />

severe wounds (our unpublished observations). The identification<br />

of the ion channels responsible for the electrical events could be studied by<br />

the use of a range of agonists and inhibitors which could be introduced via<br />

the xylem stream and their electrical consequences in the phloem could be<br />

monitored using microelectrodes.<br />

Acknowledgements. The work from our laboratory was funded by the<br />

Biotechnology and Biological Sciences Research Council, UK.

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