Microcracking of strawberry fruit cuticles: mechanism and factors.

Microscopic cracks in the cuticle (microcracks) are the first symptom of the strawberry fruit disorder 'water soaking' in which the fruit surface appears watery, translucent, and pale. Water soaking severely impacts fruit quality. The objective was to investigate the factors and mechanisms of cuticular microcracking in strawberry. Fluorescence microscopy revealed numerous microcracks in the achene depressions, on the rims between depressions and at the bases of trichomes. Microcracks in the achene depressions and on the rims were either parallel or transversely oriented relative to a radius drawn from the rim to the point of attachment of the achene. In the achene depression, the frequency of microcracks with parallel orientation decreased from the calyx end of the fruit, towards the fruit tip, while the frequency of those with transverse orientation remained constant. Most microcracks occurred above the periclinal cell walls of the epidermal cells. The long axes of the epidermal cells were primarily parallel-oriented. Microcracking increased during fruit development. Cuticle mass per fruit remained constant as fruit surface area increased but cuticle thickness decreased. When fruit developed under high relative humidity (RH) conditions, the cuticle had more microcracks than under low RH conditions. Exposing the fruit surface to increasing RHs, increased microcracking, especially above 75% RH. Liquid-phase water on the fruit surface was markedly more effective in inducing microcracking than high vapor-phase water (high RH). The results demonstrate that a combination of surface area growth strain and water exposure is causal in inducing microcracking of the strawberry cuticle.

Necked strawberries are especially susceptible to cracking.

Fruit cracking is a commercially important disorder that reduces both quantity and quality of strawberries (Fragaria × ananassa Duch.). The objective was to identify the physiological mechanism of cracking and the factors affecting cracking. Cracking is more common in necked than in normal-shaped fruit. Most macroscopic cracks ('macrocracks') occur in the seedless neck. Large fruit is more cracking susceptible than medium size or small fruit. Macrocrack orientation is predominantly latitudinal in the proximal region of the neck and longitudinal in the mid and distal regions of the neck. The neck region of necked fruit has a thicker cuticle than the body of necked or normal-shaped fruit. The vascular bundles in the neck (seedless) are orientated longitudinally, while those in the body (with seeds) are both longitudinal and radial. Epidermal cells in the neck region are elongated longitudinally, with those in the proximal region of the neck being more elongated than those in the mid or distal regions of the neck. Cuticular microcracking was more severe in necked fruit than in normal-shaped fruit. The orientations of the microcracks matched those of the macrocracks, i.e., latitudinal in the proximal neck and longitudinal in the mid and distal neck regions. Following artificial incisions (blade), gaping was significantly more pronounced in necked than in normal-shaped fruit. Incubation of fruit in deionized water induced macrocracks in about 75% of fruit. Necked fruit cracked more than normal-shaped fruit. Most macrocracks were oriented latitudinally in the proximal neck and longitudinally in the distal neck regions. The results indicate cracking results from excessive growth strains which are further increased by surface water uptake.

Detached, wetted strawberries take up substantial water in the calyx region.

In strawberry, surface disorders like 'water soaking', 'cracking' and 'shrivel' impair fruit quality of this high value crop. Water movement through the fruit surface is implicated a role in these disorders. The objective was to identify the pathways of water uptake and water loss (transpiration) and to identify factors affecting these flows. Water movement was quantified gravimetrically in detached fruit. Cumulative transpiration and uptake increased linearly with time. During ripening, fruit osmotic potential and water potential became slightly more negative. Rates of transpiration and water uptake and their corresponding permeances were constant during early ripening but increased as the fruit turned red. The permeance for osmotic water uptake was more than 10-times that for transpiration. Sealing selected regions of the fruit surface with silicone rubber allowed identification of the petal and staminal abscission zones in the calyx region and cuticular microcracks of the calyx region and receptacle as high flux pathways particularly for water uptake (osmotic). These results were confirmed by acridine orange infiltration and fluorescence microscopy. Increasing the relative humidity (RH) decreased the rate of transpiration, while increasing temperature increased both transpiration and water uptake. There was no effect of storing fruit (2 °C, ~ 80% RH) for up to 10 days. Our results identify petal and staminal abscission zones and cuticular microcracks as high flux pathways for water uptake.

Calcium ions decrease water-soaking in strawberries.

Water soaking is a common disorder of field-grown strawberries (Fragaria × ananassa Duch.). It develops when ripe fruit is exposed to rain. Here we investigate the effects of Ca on water soaking. Fruit was incubated in solutions of various Ca salts and the extent of water soaking quantified using a simple rating scheme. Exposure to CaCl2 (10 mM) decreased water soaking and anthocyanin leakage but had no effect on water uptake. The decrease in water soaking due to CaCl2 was not limited to a single cultivar but occurred in all cultivars examined. Incubating fruit in a chelating agent (EGTA) increased water soaking compared to the water control. Calcium salts of different acids varied in their effects on water soaking. Only CaCl2 reduced water soaking significantly. The chlorides of different cations, also varied in their effects on water soaking. Those of the monovalent cations had no effects on water soaking, while those of the divalent cations (CaCl2, BaCl2 and SrCl2) and of the trivalent cations (FeCl3 and AlCl3) were all effective in decreasing water soaking. Overall, CaCl2 decreased microcracking of the strawberry cuticle as compared to deionized water. Furthermore, CaCl2 also reduced the leakage of anthocyanins from flesh discs, irrespective of the osmotic potential of the incubation solution. Our results indicate that CaCl2 reduced water soaking by decreasing cuticular microcracking, by decreasing leakage of plasma membranes and, possibly, by increasing the crosslinking of cell wall constituents.

Water Soaking Disorder in Strawberries: Triggers, Factors, and Mechanisms.

Water soaking is an important surface disorder of strawberries that limits unprotected field production. The objective was to identify the mechanism(s) of water soaking. Symptomatic fruit show pale, deliquescent patches of skin. This damage extends into the flesh. Numerous cuticular microcracks occurred in water-soaked areas. Water soaking occurred only if the skin was exposed to liquid water. Water soaking was more rapid when the cuticle had been abraded. Water soaking, anthocyanin leakage, and water uptake all increased with incubation time. There was a lag phase for water soaking and anthocyanin leakage, but not for water uptake. Susceptibility to water soaking increased with fruit ripening and mass. Incubation in isotonic PEG 6000 increased cuticular microcracking but decreased water soaking and water uptake. Incubation in hypotonic fruit juice (natural and artificial) increased water soaking incidence and severity but reduced water uptake. Incubation in dilute citric and malic acids increased plasma membrane permeability as indexed by anthocyanin leakage and increased water soaking. Thus, water soaking involves cuticular microcracking, localized water uptake, bursting of cells, and the release of organic acids into the apoplast. The damage propagates from cell to cell.

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.