Strawberry fruit skins are far more permeable to osmotic water uptake than to transpirational water loss.

Water movements through the fruit skin play critical roles in many disorders of strawberry (Fragaria × ananassa Duch.) such as water soaking, cracking and shriveling. The objective was to identify the mechanisms of fruit water loss (dry skin, transpiration) and water uptake (wet skin, osmosis). Fruits were held above dried silica gel or incubated in deionized water. Water movements were quantified gravimetrically. Transpiration and osmotic uptake increased linearly with time. Abrading the thin cuticle (0.62 g m-2) increased rates of transpiration 2.6-fold, the rates of osmotic uptake 7.9-fold. The osmotic potential of the expressed juice was nearly the same for green and for white fruit but decreased in red fruit stages. Fruit turgor was low throughout development, except for green fruit. There was no relationship between the rates of water movement and fruit osmotic potential. The skin permeance for transpiration and for osmotic uptake were both high (relative to other fruit species) but were two orders of magnitude greater for osmotic uptake than for transpiration. Incubating fruit in isotonic solutions of osmolytes of different sizes resulted in increases in fruit mass that depended on the osmolyte. The rate of osmotic uptake decreased asymptotically as molecular size of the osmolyte increased. When transpiration and osmotic uptake experiments were conducted sequentially on the same fruit, the rates of transpiration were higher for fruit previously incubated in water. Fluorescence microscopy revealed considerable microcracking in a fruit previously incubated in water. Our findings indicate that the high permeance for osmotic uptake is accounted for by an extremely thin cuticle and by viscous water flow through microcracks and along polar pathways.

Correction: Crack initiation and propagation in sweet cherry skin: A simple chain reaction causes the crack to 'run'.

[This corrects the article DOI: 10.1371/journal.pone.0219794.].

Crack initiation and propagation in sweet cherry skin: A simple chain reaction causes the crack to 'run'.

Rain cracking severely affects the commercial production of many fleshy-fruit species, including of sweet cherries. The objectives were to investigate how the gaping macroscopic cracks (macrocracks) of a rain-cracked fruit can develop from microscopic cracks in the cuticle (microcracks). Incubating fruit in deionized water is well known to cause significant macrocracking. We found that after a lag phase of 2 h, the numbers and lengths of macrocracks increased. Macrocrack number approached an asymptote at 12 h, whereas macrocrack length continued to increase. The rate of macrocrack propagation (extension at the crack tip) was initially 10.8 mm h-1 but then decreased to a near-constant 0.5 mm h-1. Light microscopy revealed three characteristic zones along a developing macrocrack. In zone I (ahead of the crack), the cuticle was intact, the epidermal cells were unbroken and their cell walls were thin. In zone II, the cuticle was fractured, the first epidermal cells died and their cell walls began to thicken (swell). In zone III, most epidermal cells had died, their cell walls were swollen and cell:cell separation began along the middle lamellae. The thickness of the anticlinal epidermal cell walls and the percentage of intact living cells along a crack were closely and negatively related. Cracks were stained by calcofluor white, but there was no binding of monoclonal antibodies (mAbs) specific for hemicelluloses (LM11, LM21, LM25). Strong binding was obtained with the anti-homogalacturonan mAb (LM19), indicating the presence of unesterified homogalacturonans on the crack surface. We conclude that macrocrack propagation is related to cell death and to cell wall swelling. Cell wall swelling weakens the cell:cell adhesion between neighbouring epidermal cells, which separate along their middle lamellae. The skin macrocrack propagates like a 'run' in a fine, knitted fabric.

Patterns of microcracking in apple fruit skin reflect those of the cuticular ridges and of the epidermal cell walls.

MAIN CONCLUSION: Microcracks in the cuticle of developing apples are aligned with ridges on the inner cuticle surface and are indicative of stress-strain concentrations above the anticlinal cell walls. Microcracks occur in cuticles of most fruits. Growth strains are considered causal. In apples (Malus × domestica), microcracks usually form a mesh pattern similar to that formed by cuticular ridges. Ridge patterns are similar to those of the epidermal cells' anticlinal walls. Our aim was to identify the mechanistic bases for these pattern similarities. By quantifying ridge depth, ridge width, and the areas enclosed by ridges, we reveal the presence of major and minor ridges. Major ridges enclose two-to-four epidermal cells, minor ridges only one cell. There are similar and overlying patterns of microcracking on the cuticle's outer surface and of ridges on its inner surface-microcracks generally follow the outlines of the major ridges. In biaxial tensile tests at 20 kPa, strains were low and microcracks shallow, but at > 40 kPa, strains were higher and microcracks deeper. Microcracks traversing the cuticle are usually aligned with the anticlinal walls of the underlying epidermal cells. In general, increased skin strain is associated with increased skin transpiration. Transpiration increases are reversible for low strains but irreversible for high strains. The alignment of cuticular microcracks with the major ridges, and these with the anticlinal cell walls, indicates associated stress/strain concentrations.

Cell wall swelling, fracture mode, and the mechanical properties of cherry fruit skins are closely related.

MAIN CONCLUSION: Cell wall swelling, fracture mode (along the middle lamellae vs. across cell walls), stiffness, and pressure at fracture of the sweet cherry fruit skin are closely related. Skin cracking is a common phenomenon in many crops bearing fleshy fruit. The objectives were to investigate relationships between the mode of fracture, the extent of cell wall swelling, and the mechanical properties of the fruit skin using sweet cherry (Prunus avium) as a model. Cracking was induced by incubating whole fruit in deionised water or by fracturing exocarp segments (ESs) in biaxial tensile tests. The fracture mode of epidermal cells was investigated by light microscopy. In biaxial tensile tests, the anticlinal cell walls of the ES fractured predominantly across the cell walls (rather than along) and showed no cell wall swelling. In contrast, fruit incubated in water fractured predominantly along the anticlinal epidermal cell walls and the cell walls were swollen. Swelling of cell walls also occurred when ESs were incubated in malic acid, in hypertonic solutions of sucrose, or in water. Compared to the untreated controls, these treatments resulted in more frequent fractures along the cell walls, lower pressures at fracture (p fracture), and lower moduli of elasticity (E, i.e., less stiff). Conversely, compared to the untreated controls, incubating the ES in CaCl2 and in high concentrations of ethanol resulted in thinner cell walls, in less frequent fractures along the cell walls, higher E and p fracture. Our study demonstrates that fracture mode, stiffness, and pressure at fracture are closely related to cell wall swelling. A number of other factors, including cultivar, ripening stage, turgor, CaCl2, and malic acid, exert their effects only indirectly, i.e., by affecting cell wall swelling.

Biaxial tensile tests identify epidermis and hypodermis as the main structural elements of sweet cherry skin.

The skin of developing soft and fleshy fruit is subjected to considerable growth stress, and failure of the skin is associated with impaired barrier properties in water transport and pathogen defence. The objectives were to establish a standardized, biaxial tensile test of the skin of soft and fleshy fruit and to use it to characterize and quantify mechanical properties of the sweet cherry (Prunus avium) fruit skin as a model. A segment of the exocarp (ES) comprising cuticle, epidermis, hypodermis and adhering flesh was mounted in the elastometer such that the in vivo strain was maintained. The ES was pressurized from the inner surface and the pressure and extent of associated bulging were recorded. Pressure : strain responses were almost linear up to the point of fracture, indicating that the modulus of elasticity was nearly constant. Abrading the cuticle decreased the fracture strain but had no effect on the fracture pressure. When pressure was held constant, bulging of the ES continued to increase. Strain relaxation upon releasing the pressure was complete and depended on time. Strains in longitudinal and latitudinal directions on the bulging ES did not differ significantly. Exocarp segments that released their in vivo strain before the test had higher fracture strains and lower moduli of elasticity. The results demonstrate that the cherry skin is isotropic in the tangential plane and exhibits elastic and viscoelastic behaviour. The epidermis and hypodermis, but not the cuticle, represent the structural 'backbone' in a cherry skin. This test is useful in quantifying the mechanical properties of soft and fleshy fruit of a range of species under standardized conditions.