Mismatch between cuticle deposition and area expansion in fruit skins allows potentially catastrophic buildup of elastic strain.

MAIN CONCLUSION: The continuous deposition of cutin and wax during leaf and fruit growth is crucial to alleviate elastic strain of the cuticle, minimize the risk of failure and maintain its barrier functions. The cuticular membrane (CM) is a lipoidal biopolymer that covers primary surfaces of terrestrial plants. CMs have barrier functions in water and solute transfer and pathogen invasion. These require intact CMs throughout growth. This is a challenge particularly for fruit, because they increase in area from initiation through to maturity. Our paper investigates the effects of cutin and wax deposition on strain buildup in the CM. We use developing fruits and leaves of apple (Malus × domestica) and sweet cherry (Prunus avium) as models. The hypothesis was that the continuous deposition of the CM prevents the buildup of excessive elastic strain in fruit and leaves. Strains were quantified from decreases in surface area of CMs after isolation from epidermal discs, after wax extraction and from increases in surface area during development. Cuticle mass per unit area increased throughout development in apple fruit, and leaves of apple and sweet cherry. In sweet cherry fruit, however, CM mass increased only initially, but thereafter decreased as the surface expanded rapidly. The release of strain on CM isolation was low in apple fruit and leaves and sweet cherry leaves, but high in sweet cherry fruit. Conversely, strains fixed by the deposition of wax and cutin were high in apple fruit and leaves and sweet cherry leaves, but low in sweet cherry fruit. Our results indicate that in expanding organs, deposition of cutin and wax in the CM allows conversion of elastic to plastic strain. Hence, any lack of such deposition allows buildup of high, potentially catastrophic, elastic strain.

Evidence for a radial strain gradient in apple fruit cuticles.

MAIN CONCLUSION: The morphological outer side of the apple fruit cuticle is markedly more strained than the inner side. This strain is released upon wax extraction. This paper investigates the effect of ablating outer and inner surfaces of isolated cuticular membranes (CM) of mature apple (Malus × domestica) fruit using cold atmospheric pressure plasma (CAPP) on the release of strain after extraction of waxes. Strain release was quantified as the decrease in area of CM discs following CAPP treatment and subsequent solvent extraction of wax. Increasing duration of CAPP treatment proportionally decreased CM mass per unit area. There was no difference in mass loss rate between CAPP treatments of outer or inner surfaces. Also, there was no difference in surface area of CMs before and after CAPP treatment. However, upon subsequent wax extraction, surface area of CMs decreased indicating the release of strain. Increasing the duration of CAPP treatment resulted in increasing strain release up to 47.7 ± 8.0 % at 20 min when CAPP was applied to the inner surface. In contrast, strain release was independent of CAPP duration averaging about 12.1 ± 0.6 % when applied to the outer surface of the CM. Our results provide evidence for a marked gradient of strain between the outer side (strained) and the inner side of the CM (not strained) of mature apple fruit.

Intracuticular wax fixes and restricts strain in leaf and fruit cuticles.

This paper investigates the effects of cuticular wax on the release of strain and on the tensile properties of enzymatically isolated cuticular membranes (CMs) taken from leaves of agave (Agave americana), bush lily (Clivia miniata), holly (Ilex aquifolium), and ivy (Hedera helix) and from fruit of apple (Malus × domestica), pear (Pyrus communis), and tomato (Lycopersicon esculentum). Biaxial strain release was quantified as the decrease in CM disc area following wax extraction. Stiffness, maximum strain and maximum force were determined in uniaxial tensile tests using strips of CM and dewaxed CMs (DCMs). Biaxial strain release, stiffness, and maximum strain, but not maximum force, were linearly related to the amount of wax extracted. Apple CM has the most wax and here the effect of wax extraction was substantially accounted for by the embedded cuticular wax. Heating apple CM to 80°C melted some wax constituents and produced an effect similar to, but smaller than, that resulting from wax extraction. Our results indicate that wax 'fixes' strain, effectively converting reversible elastic into irreversible plastic strain. A consequence of 'fixation' is increased cuticular stiffness.