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

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96 4 <strong>Water</strong> <strong>Permeability</strong><br />

only permeances have been measured, but not diffusion <strong>and</strong> partition coefficients in<br />

the transport-limiting barrier.<br />

<strong>Water</strong> permeability <strong>of</strong> CM has been studied using different approaches. With the<br />

cup method (Schönherr <strong>and</strong> Lendzian 1981) the inner surface <strong>of</strong> the CM is in contact<br />

with water, <strong>and</strong> the outer surface is exposed to dry air <strong>of</strong> nearly 0% humidity<br />

(Sect. 9.7). The amperometric method (Becker et al. 1986) used nitrogen gas saturated<br />

with water vapour as donor at the morphological inner surface <strong>of</strong> the CM, <strong>and</strong><br />

dry nitrogen served as receiver facing the morphological outer surface <strong>of</strong> the CM.<br />

Permeances obtained with these two methods are minimum permeances, because<br />

the outer surface <strong>of</strong> the CM is not or only weakly hydrated. Other experiments<br />

had been conducted with partial vapour pressures between 0.02 <strong>and</strong> 1.0 or with<br />

liquid water in contact with the outer surface <strong>of</strong> the CM while the inner surface<br />

was wet by water (Niederl et al. 1998; Schönherr 1976b; Schönherr <strong>and</strong> Schmidt<br />

1979; Schreiber 2001). These experiments show if permeance increases with partial<br />

vapour pressure. With 100% humidity or liquid water on the receiver side, maximum<br />

permeances are measured.<br />

4.6.2.1 Chemical Composition <strong>of</strong> Wax <strong>and</strong> Its Relationship<br />

to <strong>Water</strong> <strong>Permeability</strong><br />

Since neither thickness <strong>of</strong> the CM nor amounts <strong>of</strong> wax provide a satisfactory<br />

explanation for the large variability in water permeability <strong>of</strong> CM from different<br />

plant species, some workers in the field have postulated that permeability might be<br />

related to wax composition. Qualitative <strong>and</strong> quantitative wax composition can vary<br />

considerably with leaf development <strong>and</strong> with growing conditions. Wax amounts<br />

<strong>and</strong> composition can be measured with high accuracy using modern capillary gas<br />

chromatography/mass spectrometry. Each plant species has its own specific wax<br />

composition, with homologues varying in chain lengths distribution <strong>and</strong> substance<br />

class composition. This might account for differences in cuticular transpiration<br />

observed with different species. By analysing wax composition <strong>and</strong> measuring water<br />

permeability in parallel, this hypothesis can be tested. Suitable data sets are available<br />

for astomatous isolated CM from Citrus aurantium (Geyer <strong>and</strong> Schönherr 1990;<br />

Riederer <strong>and</strong> Schneider 1990) <strong>and</strong> Hedera helix (Hauke <strong>and</strong> Schreiber 1998).<br />

Wax composition <strong>of</strong> Citrus aurantium leaves varied with leaf age <strong>and</strong> depended<br />

on growing conditions (Riederer <strong>and</strong> Schneider 1990). Increased day temperatures<br />

(15–35 ◦ C) resulted in smaller wax amounts (alkanes, alcohols, acids <strong>and</strong> esters)<br />

by factors <strong>of</strong> 2–3, whereas increased night temperatures had the opposite effect.<br />

Humidity (50% or 90%) had no effect on wax composition. <strong>Water</strong> permeability<br />

determined using the cup method (Sect. 9.7) was measured at 25 ◦ C with subsets <strong>of</strong><br />

CM from identical isolations. Even though wax coverage <strong>and</strong> composition depended<br />

on growing conditions, water permeability <strong>of</strong> CM was not affected. <strong>Permeability</strong><br />

<strong>of</strong> all sets <strong>of</strong> CM decreased during storage <strong>of</strong> CM at 8 ◦ C. During about 50 days<br />

<strong>of</strong> storage, permeance decreased to about 40% <strong>of</strong> the initial Pw. This reduction in<br />

permeance was attributed to a change in wax structure during storage. Possibly the

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