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Literature review<br />

BLANK and YOUNG (1998) reported that smoke increases the permeability to<br />

solutes <strong>of</strong> a subdermal seed membrane for some species <strong>of</strong> California chaparral.<br />

They also stated that these specific mechanisms <strong>of</strong> fire cue stimulation may be<br />

species dependent. This response involves triggering via elevated nutrient content or<br />

via the presence <strong>of</strong> stimulating gases in the smoke or triggering chemicals that<br />

permeate the embryo and induce enzymatic changes that trigger germination. In an<br />

experiment done by BROWN and VAN STADEN (1997) the dormancy <strong>of</strong> celery<br />

seeds was broken by a combination <strong>of</strong> plant-derived smoke, benzyladenine and<br />

gibberellins in the dark at temperatures between 18 and 26°C. From these results it<br />

could be argued that smoke extracts act in a similar way to cytokinins in the celery<br />

seed as it enhances gibberellin activity.<br />

In a study conducted at the ultrastructure level by EGERTON-WARBURTON (1998),<br />

the causal factor(s) associated with seed dormancy and the stimulation <strong>of</strong><br />

germination were investigated for Emmenanthe penduliflora (Hydrophyllaceae)<br />

seeds. It was found that a short exposure to smoke resulted in two major<br />

morphological changes. These changes are closely associated with the stimulation<br />

and acceleration <strong>of</strong> germination. The first major and most visible smoke induced<br />

morphological change observed was an intense chemical scarification <strong>of</strong> the external<br />

cuticle (EGERTON-WARBURTON, 1998). This has a direct and destabilising effect<br />

on the external cuticle and is manifested as the formation <strong>of</strong> oil-like spheres or<br />

micelles. These micelles increase the surface area <strong>of</strong> the seed for the exchange <strong>of</strong><br />

water and solutes, as well as altering the hydrophobicity <strong>of</strong> the seed surface<br />

(EGERTON-WARBURTON, 1998).<br />

The second, more important smoke induced morphological change occurred at the<br />

internal cuticle. Here the exposure to smoke stimulus caused a significant increase in<br />

the number and diameter <strong>of</strong> permeate channels in the cuticle <strong>of</strong> Emmenanthe<br />

penduliflora seeds (EGERTON-WARBURTON, 1998). The creation <strong>of</strong> such channels<br />

within the cuticle increases the permeability <strong>of</strong> this layer. It was shown through the<br />

use <strong>of</strong> the fluorescent apoplastic tracer dye, lucifer yellow (carbohydrazide), that<br />

51

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