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

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4.6 <strong>Water</strong> <strong>Permeability</strong> <strong>of</strong> Isolated Astomatous Cuticular Membranes 111<br />

could be responsible for the difference. Unfortunately, in the two sets <strong>of</strong> experiments<br />

different species were used. Nevertheless, it is interesting that these two independent<br />

approaches yield estimates for Dw <strong>of</strong> the same order <strong>of</strong> magnitude. It is an<br />

advantage <strong>of</strong> these approaches that solubility <strong>of</strong> water in cuticular wax (wax/water<br />

partition coefficients) are not needed for estimating Dw. Unfortunately, assumptions<br />

concerning the thickness <strong>of</strong> the limiting skin must be made, <strong>and</strong> these assumptions<br />

enter as the square.<br />

4.6.4 <strong>Water</strong> <strong>Permeability</strong> <strong>of</strong> Paraffin Waxes<br />

Model III A postulates that wax in <strong>and</strong> on top <strong>of</strong> the CM completely determine water<br />

permeability, while aqueous pores are not involved. This model is difficult to test,<br />

because there are no data on water permeability <strong>of</strong> reconstituted waxes. The evidence<br />

presented is indirect, <strong>and</strong> relies on co-penetration in CM <strong>of</strong> lipophilic solutes<br />

<strong>and</strong> water (Figs. 4.16 <strong>and</strong> 4.17). The observation that cuticular transpiration is correlated<br />

with diffusion <strong>of</strong> stearic acid in cuticular wax (Fig. 4.15) agrees with model<br />

III A, <strong>and</strong> the correlation between water <strong>and</strong> benzoic acid permeances is also consistent<br />

with model III A, even though (4.17) indicates that, in CM <strong>of</strong> species with<br />

relatively high Pw, water transport in aqueous pores may have been involved. On<br />

the other h<strong>and</strong>, data <strong>of</strong> Table 4.8 are consistent with model IIIC, which assumes<br />

that a significant amount <strong>of</strong> water crosses the CM via aqueous pores, which can be<br />

blocked by silver chloride precipitates. There is no reason why CM <strong>of</strong> all species<br />

investigated so far should meet model III A, B or C criteria. The question arises<br />

which thickness <strong>of</strong> a wax barrier could account for observed water permeances.<br />

Since we have no data for reconstituted wax, we shall take a look at data obtained<br />

with paraffin waxes.<br />

4.6.4.1 <strong>Water</strong> Permeance <strong>of</strong> Polyethylene <strong>and</strong> Paraffin Wax<br />

In Table 4.7, data on wax coverage for CM <strong>of</strong> several species are given. They<br />

range from 11.8µgcm −2 (Citrus aurantium) to 1,317µgcm −2 (Malus). For Allium<br />

(Table 4.6), only 1µgcm −2 has been determined using gas chromatography. Multiplying<br />

the wax amounts by specific weight (0.9gcm −3 ), we obtain wax layers<br />

having a thickness from 11 nm (Allium), 131 nm (Citrus) <strong>and</strong> 14.6µm (Malus). For<br />

most species in Table 4.7, wax amounts varied from 30 to 100µgcm −2 , resulting in<br />

wax layers <strong>of</strong> 333 nm to 1.1µm. Can we estimate Pwv for such wax layers?<br />

Waxes are partially crystalline, hydrophobic <strong>and</strong> they consist mainly <strong>of</strong> CH2<br />

<strong>and</strong> CH3 groups. This is also true for polyethylene, which is partially crystalline<br />

but exclusively composed <strong>of</strong> CH2 <strong>and</strong> CH3 groups. Pwv <strong>of</strong> polyethylene is 3.3 ×<br />

10 −11 m 2 s −1 (Table 4.1). A membrane <strong>of</strong> a comparable physico-chemical structure<br />

is parafilm (polyethylene covered with paraffin) <strong>and</strong> water permeability Pwv <strong>of</strong><br />

127µm-thick parafilm is 3.33 × 10 −7 ms −1 (Santrucek et al. 2004), which results in

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