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Literature review<br />

these channels permit the rapid exchange <strong>of</strong> water and solutes between the external<br />

environment and the seed.<br />

Even though these major morphological changes have a large influence on the<br />

function <strong>of</strong> the cell, they do not significantly alter the general shape or dimensions <strong>of</strong><br />

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

speculated that the mechanism by which such morphological changes benefit the<br />

plant may be a synergy between the observed increased cuticular permeability and a<br />

simultaneous leaching <strong>of</strong> endogenous inhibitors <strong>of</strong> germination during imbibition.<br />

They also further described the mechanism <strong>of</strong> formation <strong>of</strong> permeate channels.<br />

Because the permeate channels occurs in the cuticle, the formation <strong>of</strong> channels has<br />

to be the result <strong>of</strong> a preceding reaction between certain constituents <strong>of</strong> smoke and<br />

the semi-crystalline structure <strong>of</strong> waxes (EGERTON-WARBURTON, 1998). This is a<br />

rather promising argument as the pyrolysis <strong>of</strong> cellulose alone produces a collection <strong>of</strong><br />

compounds, such as aromatic hydrocarbons, ketones and a number <strong>of</strong> organic acids,<br />

which all have the potential to dissolve or modify waxes.<br />

Smoke also contains a number <strong>of</strong> compounds that may act as surfactants (e.g.<br />

alcohols) (EGERTON-WARBURTON, 1998). These surfactants modify cuticular<br />

layers by plasticizing the molecular structure <strong>of</strong> waxes (EGERTON-WARBURTON,<br />

1998). The sorption <strong>of</strong> such surfactants to the cuticle also creates hydrophilic<br />

channels. This increases the area <strong>of</strong> channels within the cuticle that leads to<br />

accelerated transcuticular penetration <strong>of</strong> solutes in several species.<br />

Using liquid chromatography, BLANK and YOUNG (1998) discerned over 30 organic<br />

anions in aqueous extracts <strong>of</strong> soil heated between 250 to 450°C. BLANK and<br />

YOUNG (1998) speculated that these unknown compounds or combinations <strong>of</strong><br />

compounds could be cueing agents.<br />

FLEMATTI et al. (2004) identified the UV absorbance maximum and molecular<br />

weight <strong>of</strong> a germination-enhancing compound in smoke. They also assigned a<br />

molecular formula <strong>of</strong> C8H6O3 to the compound. They also identified the compound as<br />

the butenolide, 3-methyl-2H-furo[2,3-c]pyran-2-one and were able to synthesize it.<br />

52

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