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bundles (Table 20.1). Since the depolarization of a SWP lingers longer than
that of an AP, SWPs may be more effective than APs in triggering the
opening of excitable, i.e., voltage-gated Ca 2+ and anion channels needed
for AP induction and propagation. SWPs in Vicia , cucumber and sunflower
plants are frequently accompanied by action spikes (Roblin 1985; Roblin
and Bonnemain 1985; Stahlberg and Cosgrove 1994, 1997a; Stankovic and
Davies 1998; Stankovic et al. 1997). Conversely, the fact that no depolariza-
tion has ever been reported to cause a propagating SWP suggests that APs
are unable to trigger SWPs.
20.4
The Propagation of SWPs
According to previously introduced considerations (Figs. 20.3 and 20.4),
SWPs only undergo an apparent propagation with an appearance and range
that is determined by the gradual decline of the inducing pressure signal.
Unlike flames and pressure chambers, wounding by puncturing, surface
cuts and organ excision generates a hydraulic signal that is based on exist-
ing pressure gradients in the plants (Stahlberg and Cosgrove 1997c). This
provides useful information of how, e.g., species, age, light and other envi-
ronmental conditions affect the ability of plants to generate SWPs as well
as their size and range (Alarcon and Malone 1994; Stahlberg and Cosgrove
1994, 1995; Stahlberg et al. 2005). An example is given in Fig. 20.6.
Figure 20.6, example B shows the rapid ascent of a SWP from a punc-
ture wound in the hypocotyl to the upper hypocotyl, epicotyl and leaf in
an intact, illuminated sunflower plant. When the nonsubmersed part of
a hypocotyl was punctured, the limited water supply near the punctured
vascular bundle supported only a SWP with a short range limited to the
hypocotyl (Fig. 20.6, example A). When punctured under water, the gener-
ated SWP had an increased, systemic range. The increased range is a clear
indication of the hydraulic nature of SWP propagation. Using a stimulus
that is close to the small scale of insect-inflicted wounds, we present here
a short SWP that could be more representative than those published before
as a consequence of larger injuries. It propagates with the same rate as those
induced by root excision and shows the same range extension to wounding
under water (Stahlberg et al. 2005b).
Although propagation rates of SWPs and APs overlap (Roblin 1985; Rob-
lin and Bonnemain 1985; Stankovic et al. 1997), SWPs have four distinctive
features that clearly separate their movement from that of APs. First, SWPs
propagate with a measurable decrement (loss of amplitude) along the plant
surface. Excision-induced SWPs in sunflower stems have a decrement of
2.5% cm −1 (Stahlberg et al. 2005b). Second, SWPs propagate from the stem
 
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