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Water and Solute Permeability of Plant Cuticles: Measurement and ...

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24 1 Chemistry <strong>and</strong> Structure <strong>of</strong> <strong>Cuticles</strong> as Related to <strong>Water</strong> <strong>and</strong> <strong>Solute</strong> <strong>Permeability</strong><br />

The chemical nature <strong>of</strong> the CP is a matter <strong>of</strong> debate (Jeffree 2006). Some workers<br />

believe that electron-lucent lamellae are waxes, while the more dense lamellae are<br />

made <strong>of</strong> cutin. This is pure speculation, because it has not been established that<br />

typical ester cutin is present in the CP at all. The mass <strong>of</strong> the CP is very small relative<br />

to the total mass <strong>of</strong> the CM, <strong>and</strong> constituents that occur only in traces might be lost<br />

during processing. After transesterification <strong>of</strong> cutin from positions 2–3 cm where<br />

the CP contributes most <strong>of</strong> the mass <strong>of</strong> the CM (Fig. 1.8), only nine n-fatty acid<br />

(C12, C14, C16 <strong>and</strong> C18) homologues have been identified which amounted to 52%<br />

<strong>of</strong> the total mass <strong>of</strong> cutin. The most frequent cutin acid (11%) was 9,10-epoxy-18hydroxyoctadecanoic<br />

acid (Riederer <strong>and</strong> Schönherr 1988). It is difficult to envision<br />

a polymer composed <strong>of</strong> 50% <strong>of</strong> simple fatty acids.<br />

The CP <strong>of</strong> Clivia cuticle survived extraction <strong>of</strong> CM with chlor<strong>of</strong>orm (Fig. 1.9)<br />

<strong>and</strong> exhibited heavy contrast. Electron-lucent lamellae were preserved, <strong>and</strong> this is<br />

unlikely to happen if they were made <strong>of</strong> waxes. The CL was differentiated into an<br />

external (ECL) <strong>and</strong> internal (ICL) layer, with large differences in contrast. Since<br />

the specimen was extracted, penetration by KMnO4 was not hindered by waxes,<br />

<strong>and</strong> failure <strong>of</strong> the ECL to develop contrast indicates that reactive groups had been<br />

eliminated during cutan formation.<br />

Fig. 1.9 Transverse section <strong>of</strong> a polymer matrix membrane obtained from the adaxial surface <strong>of</strong><br />

a young Clivia miniata leaf. The MX was stained with KMnO4 prior to embedding, <strong>and</strong> sections<br />

were stained with uranyl acetate <strong>and</strong> lead citrate. (Taken from Schmidt <strong>and</strong> Schönherr 1982)

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