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

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108 4 <strong>Water</strong> <strong>Permeability</strong><br />

Table 4.9 <strong>Water</strong> permeance (Pwv) <strong>and</strong> diffusion coefficients (Dw) <strong>of</strong> cuticular membranes at 25 ◦ C<br />

Species te (s) ℓ(µm) Pwv(ms −1 ) Dw(m 2 s −1 ) Cw(gkg −1 ) Cw(gkg −1 ) a<br />

Schefflera L. 930 2.96 8.19 × 10 −7 1.57 × 10 −15 33 –<br />

Clivia L. 770 6.50 1.14 × 10 −6 9.14 × 10 −15 17 –<br />

Hedera L. 933 4.33 2.66 × 10 −6 3.35 × 10 −15 73 –<br />

Nerium L. 960 12.81 3.25 × 10 −6 2.84 × 10 −14 34 24<br />

Ficus L. 512 5.68 4.25 × 10 −6 1.05 × 10 −14 48 30<br />

Citrus L. 264 2.87 1.20 × 10 −5 5.20 × 10 −15 139 63<br />

Pyrus L. 550 3.12 1.22 × 10 −5 2.95 × 10 −15 270 43<br />

Solanum F. 640 6.47 2.23 × 10 −5 1.08 × 10 −14 244 –<br />

Capsicum F. 361 8.03 9.28 × 10 −5 2.98 × 10 −14 523 38<br />

Lycopersicon F. 215 8.00 1.42 × 10 −4 4.96 × 10 −14 477 43<br />

D was calculated from the hold-up time <strong>and</strong> the total thickness (ℓ) <strong>of</strong> CM; Cw is sorption <strong>of</strong> water<br />

at 25 ◦ C <strong>and</strong> 100% humidity calculated from P/D [(3.17) <strong>and</strong> (3.20)] using the data by Becker<br />

et al. (1986). L is leaf CM <strong>and</strong> F is fruit CM. (Data from Becker et al. 1986)<br />

a Data taken from Chamel et al. (1991) derived from water vapour sorption<br />

Permeance varied by a factor <strong>of</strong> 173, <strong>and</strong> ranged from 8.19 × 10 −7 (Schefflera<br />

leaf) to 1.42 × 10 −4 ms −1 (tomato fruit). Diffusion coefficients varied by a factor <strong>of</strong><br />

only 32, <strong>and</strong> ranged from 1.57 × 10 −15 (Schefflera) to 4.96 × 10 −14 m 2 s −1 (tomato<br />

fruit). According to theory [Chap. 2; (2.18)] one might conclude that variations in<br />

membrane thickness <strong>and</strong> partition coefficient caused these differences. The Dw values<br />

are much lower than those for synthetic polymers, PVA being the only exception<br />

(Table 4.1). With some CM (Nerium <strong>and</strong> Ficus), water sorption (Cw) derived from<br />

the Pwv/Dw ratio is low <strong>and</strong> similar to sorption in CM when determined gravimetrically<br />

(Table 4.9, last column). With the other CM, water concentration calculated<br />

from Pwv/Dw is unreasonably high, <strong>and</strong> much larger than that determined by a<br />

sorption experiment. This indicates structural heterogeneity <strong>of</strong> these membranes.<br />

When calculating Dw from hold-up time, the thickness <strong>of</strong> the CM enters into consideration.<br />

Becker et al. (1986) used the weight average thickness, while the waxy<br />

limiting skin or the cuticle proper is the limiting barrier both in water <strong>and</strong> solute<br />

diffusion (Sects. 4.6 <strong>and</strong> 6.5). The thickness <strong>of</strong> the limiting skin in CM <strong>of</strong> various<br />

plant species is not known, but plausible estimates are 100–500 nm (Sect. 1.4). In<br />

calculating Dw from (2.5), thickness <strong>of</strong> the limiting barrier enters as ℓ 2 , <strong>and</strong> using a<br />

thickness less than total thickness <strong>of</strong> the CM would result in lower diffusion coefficients<br />

than those in Table 4.9. In Sect. 6.5.2 we estimate the diffusion coefficient<br />

<strong>of</strong> water in cuticular wax as 1.2 × 10 −16 m 2 s −1 . This is about a factor <strong>of</strong> 10 lower<br />

than Dw values in leaf CM estimated from the hold-up time (Table 4.9). With Citrus<br />

CM, a Dw <strong>of</strong> 1.2 × 10 −16 m 2 s −1 would have been obtained with ℓ equal to 0.44µm.<br />

This amounts to 15% <strong>of</strong> the total thickness <strong>of</strong> the CM <strong>and</strong> appears plausible. This<br />

neglects the fact that the waxy diffusion path <strong>of</strong> water is tortuous, but in calculating<br />

Dw for synthetic polymers membrane thickness (ℓ) is used (Table 4.1) <strong>and</strong> tortuosity<br />

is also disregarded.<br />

We can further test these cuticles for homogeneity by plotting Pwv vs 1/ℓ <strong>and</strong> Dw<br />

vs 1/6× hold-up time. In homogenous membranes, these plots should be linear. The

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