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

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Problems 27<br />

10. One <strong>of</strong> them is eliminated during hydroxylation in synthesis <strong>of</strong> cutin acids <strong>of</strong><br />

the C16-family. Thus, the specific radioactivity <strong>of</strong> the cutin acids was only half as<br />

high as that <strong>of</strong> 3 H-hexadecanoic acid, <strong>and</strong> the maximum rate <strong>of</strong> 3 H-cutin synthesis<br />

should be twice as high, that is 0.6µgcm −2 h −1 . At this rate, 14.4µgcm −2 <strong>of</strong><br />

3 H-cutin could be synthesised in 24 h. Between positions 3 <strong>and</strong> 5, the ester cutin<br />

fraction increased by about 50µg (Fig. 1.2). We do not know the growth rate <strong>of</strong><br />

leaves <strong>and</strong> epidermis cells. If they need a day for 1 cm, about one third <strong>of</strong> the cutin<br />

synthesised would belong to the C16-family.<br />

About 2% <strong>of</strong> the weight <strong>of</strong> the CM were C16-cutin acids at position 2–3 cm<br />

(Riederer <strong>and</strong> Schönherr 1988). At 5–6 cm, C16-cutin acids amounted to about 25%.<br />

This shows that C16-cutin acids occur in Clivia CM, but it is also clear that ester<br />

cutin is made up primarily by C18-cutin acids. Some <strong>of</strong> the 3 H-hexadecanoic acid<br />

molecules may have been elongated <strong>and</strong> turned into C18-cutin acids. Unfortunately,<br />

the radioactive cutin acids were not identified chemically.<br />

These studies demonstrate that Clivia cuticles are very dynamic structures that<br />

greatly change during development. The CP appears first <strong>and</strong> is maintained, but<br />

thickness <strong>of</strong> the CL <strong>and</strong> its chemical composition undergo major changes. Cutin synthesis<br />

occurred even at the tip <strong>of</strong> the leaf. After cell expansion is complete, non-ester<br />

cutin occurs in large amounts because ester cutin is converted into non-ester cutin<br />

(Fig. 1.10). There are no comparable studies <strong>of</strong> epidermis <strong>and</strong> cuticle development<br />

in other plant species.<br />

1.4.4 Lateral Heterogeneity<br />

The cuticle over ordinary epidermal cells (pavement cells) covers the surfaces <strong>of</strong><br />

stems, leaves, flowers <strong>and</strong> fruits. Many mechanistic studies into permeability <strong>of</strong> cuticles<br />

were carried out using isolated cuticular membranes which lack trichomes <strong>and</strong><br />

stomata. Much less is known about the involvement <strong>of</strong> specialised epidermal cells in<br />

water <strong>and</strong> solute permeation. However, trichomes <strong>and</strong> stomata occur on most leaves<br />

<strong>and</strong> stems (Glover <strong>and</strong> Martin 2000: Bird <strong>and</strong> Gray 2003), <strong>and</strong> it is well-established<br />

that permeability <strong>of</strong> cuticles over these special structures differs from that over pavement<br />

cells (Strugger 1939; Bauer 1953; Franke 1960; 1967; Meier-Maercker 1979).<br />

The cuticle over guard cells <strong>and</strong> trichomes is <strong>of</strong>ten traversed by aqueous pores, <strong>and</strong><br />

they play a decisive role in foliar penetration <strong>of</strong> ionic solutes (Schönherr 2006). This<br />

aspect <strong>of</strong> foliar penetration is treated comprehensively in Chap. 5.<br />

Problems<br />

1. What is the average thickness <strong>of</strong> a cuticle, <strong>and</strong> what are the upper <strong>and</strong> lower<br />

values <strong>of</strong> very thick <strong>and</strong> very thin cuticles?

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