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

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178 6 Diffusion <strong>of</strong> Non-Electrolytes<br />

MX membranes this asymmetry factor was only 10, because desorption from the<br />

outer surface was higher. This is still a high asymmetry, since with a homogeneous<br />

membrane the factor should have been unity.<br />

This asymmetrical desorption pattern indicates that CM <strong>and</strong> MX membranes are<br />

composed <strong>of</strong> at least two compartments. The bulk <strong>of</strong> the 2,4-D was contained in<br />

the inner volume element, <strong>and</strong> it was desorbed through the inner surface. We call<br />

this inner domain <strong>of</strong> the cuticle the sorption compartment (soco), <strong>and</strong> morphologically<br />

it is identical with the cuticular layer(s) seen in TEM (Sect. 1.4). The outer<br />

domain across which only very small amounts <strong>of</strong> 2,4-D were desorbed is the cuticle<br />

proper, <strong>and</strong> we refer to it as limiting skin or limiting layer (Fig. 6.16). Volumes <strong>and</strong><br />

thicknesses <strong>of</strong> these two layers cannot be deduced from desorption plots. As in 4 h<br />

only 5% 2,4-D diffused across the outer surface <strong>of</strong> the CM, diffusion coefficients<br />

in the limiting skin must have been very low <strong>and</strong> much lower than in the sorption<br />

compartment. The data do not reveal if it is a wax layer on top <strong>of</strong> the cuticle or if the<br />

barrier consists <strong>of</strong> waxes embedded in the outer fraction <strong>of</strong> the MX. A combination<br />

<strong>of</strong> both is possible as well. In any event waxes are involved, since extracting them<br />

reduced asymmetry greatly (Fig. 6.15).<br />

When Mt/M0 (desorption through the inner surface) was plotted against the<br />

square root <strong>of</strong> time (Fig. 6.15b), plots were not linear up to Mt/M0 equal to 0.5, as<br />

would be expected with homogeneous membranes (Fig. 2.10b), but diffusion coefficients<br />

can still be estimated from the initial slope using (2.35). Since both CM <strong>and</strong><br />

MX are heterogeneous, these D-values are some kind <strong>of</strong> average characterising diffusion<br />

<strong>of</strong> 2,4-D in the sorption compartment (cuticular layer). For these calculations,<br />

Fig. 6.16 Schematic drawing <strong>of</strong> a cross section <strong>of</strong> a cuticle showing the thin limiting skin <strong>and</strong> the<br />

thick sorption compartment (not to scale). Modified from Bauer <strong>and</strong> Schönherr (1992)

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