Peel Russeting in ‘Apple’ Mango (Mangifera indica L.): Characterization, Mechanisms and Management

The aesthetic value and marketability of table fruits are greatly reduced by russeting, a disorder that severely affects the ‘Apple’ mango (Mangifera indica L.) cultivar in Kenya. Despite its prevalence, the underlying mechanisms and prevention strategies for russeting in mangoes are unknown. To address this gap in knowledge, this project aimed to: (1) review existing literature on russeting, (2) characterize the disorder in ‘Apple’ mango, (3) identify its mechanistic basis in comparison to a non-russet susceptible cultivar, (4) investigate the role of moisture and (5) lenticels on russeting, and (6) develop strategies to prevent the disorder. To achieve these objectives, russeting was quantified in ‘Apple’ mango within fruit and in different geographic locations in Kenya. Fruit skins and cuticles from russet susceptible ‘Apple’ and russet tolerant ‘Tommy Atkins’ mangoes were examined during fruit development, and cuticular strain was partitioned into its reversible and irreversible components. Mechanical properties of isolated cuticles from both cultivars were tested. The role of moisture in microcrack development and russeting was studied by partially wetting the fruit surface. Lenticels were characterized microscopically across cultivars and locations. Field studies were also conducted to establish the effect of pre-harvest bagging on russeting and postharvest performance. The results showed that russeting in ‘Apple’ mango increased with fruit development particularly in the stem end region. Russeting was triggered by rainfall and low temperature. The skin’s permeance to water vapor was larger in russeted than in non-russeted skin. The cuticle of ‘Apple’ mango was thinner than that of ‘Tommy Atkins’. Strains released on excision and isolation and wax extraction were higher in ‘Apple’ than in ‘Tommy Atkins’. Stiffness, fracture force, and strain at fracture were consistently lower in ‘Apple’ than in ‘Tommy Atkins’. Surface wetness induced microcracking and increased the skin's water vapor permeance, and moisture-treated fruit skins later developed russet symptoms in ‘Apple’ mango. Russeting began at lenticels and then spread across the surface, ultimately forming a network of rough, brown patches over the skin. Cross-sections of russeted areas revealed stacks of phellem cells. Pre-harvest bagging of mangoes effectively prevented russeting and lowered the rates of transpiration postharvest

Russeting of Fruits: Etiology and Management

The skin of a fruit protects the vulnerable, nutrient-rich flesh and seed(s) within from the hostile environment. It is also responsible for the fruit’s appearance. In many fruitcrop species, russeting compromises fruit appearance and thus commercial value. Here, we review the literature on fruit russeting, focusing on the factors and mechanisms that induce it and on the management and breeding strategies that may reduce it. Compared with a primary fruit skin, which is usually distinctively colored and shiny, a secondary fruit skin is reddish-brown, dull and slightly rough to the touch (i.e., russeted). This secondary skin (periderm) comprises phellem cells with suberized cell walls, a phellogen and a phelloderm. Russeted (secondary) fruit skins have similar mechanical properties to non-russeted (primary) ones but are more plastic. However, russeted fruit skins are more permeable to water vapor, so russeted fruits suffer higher postharvest water loss, reduced shine, increased shrivel and reduced packed weight (most fruit is sold per kg). Orchard factors that induce russeting include expansion-growth-induced strain, surface wetness, mechanical damage, freezing temperatures, some pests and diseases and some agrochemicals. All these probably act via an increased incidence of cuticular microcracking as a result of local concentrations of mechanical stress. Microcracking impairs the cuticle’s barrier properties. Potential triggers of russeting (the development of a periderm), consequent on cuticular microcracking, include locally high concentrations of O2, lower concentrations of CO2 and more negative water potentials. Horticulturists sometimes spray gibberellins, cytokinins or boron to reduce russeting. Bagging fruit (to exclude surface moisture) is also reportedly effective. From a breeding perspective, genotypes having small and more uniform-sized epidermal cells are judged less likely to be susceptible to russeting

Russeting in ‘Apple’ Mango: Triggers and Mechanisms

Russeting is an important surface disorder of many fruitcrop species. The mango cultivar ‘Apple’ is especially susceptible to russeting. Russeting compromises both fruit appearance and postharvest performance. The objective was to identify factors, mechanisms, and consequences of russeting in ‘Apple’ mango. Russeting was quantified on excised peels using image analysis and a categorical rating scheme. Water vapour loss was determined gravimetrically. The percentage of the skin area exhibiting russet increased during development. Russet began at lenticels then spread across the surface, ultimately forming a network of rough, brown patches over the skin. Cross-sections revealed stacks of phellem cells, typical of a periderm. Russet was more severe on the dorsal surface of the fruit than on the ventral and more for fruit in the upper part of the canopy than in the lower. Russet differed markedly across orchards sites of different climates. Russet was positively correlated with altitude, the number of rainy days, and the number of cold nights but negatively correlated with minimum, maximum, and mean daily temperatures, dew point temperature, and heat sum. Russeted fruit had higher transpiration rates than non-russeted fruits and higher skin permeance to water vapour. Russet in ‘Apple’ mango is due to periderm formation that is initiated at lenticels. Growing conditions conducive for surface wetness exacerbate russeting

Low cuticle deposition rate in ‘Apple’ mango increases elastic strain, weakens the cuticle and increases russet

Russeting compromises appearance and downgrades the market value of many fruitcrops, including of the mango cv. ‘Apple’. The objective was to identify the mechanistic basis of ‘Apple’ mango’s high susceptibility to russeting. We focused on fruit growth, cuticle deposition, stress/strain relaxation analysis and the mechanical properties of the cuticle. The non-susceptible mango cv. ‘Tommy Atkins’ served for comparison. Compared with ‘Tommy Atkins’, fruit of ‘Apple’ had a lower mass, a smaller surface area and a lower growth rate. There were little differences between the epidermal and hypodermal cells of ‘Apple’ and ‘Tommy Atkins’ including cell size, cell orientation and cell number. Lenticel density decreased during development, being lower in ‘Apple’ than in ‘Tommy Atkins’. The mean lenticel area increased during development but was consistently greater in ‘Apple’ than in ‘Tommy Atkins’. The deposition rate of the cuticular membrane was initially rapid but later slowed till it matched the area expansion rate, thereafter mass per unit area was effectively constant. The cuticle of ‘Apple’ is thinner than that of ‘Tommy Atkins’. Cumulative strain increased sigmoidally with fruit growth. Strains released stepwise on excision and isolation (εexc+iso), and on wax extraction (εextr) were higher in ‘Apple’ than in ‘Tommy Atkins’. Membrane stiffness increased during development being consistently lower in ‘Apple’ than in ‘Tommy Atkins’. Membrane fracture force (Fmax) was low and constant in developing ‘Apple’ but increased in ‘Tommy Atkin’. Membrane strain at fracture (εmax) decreased linearly during development but was lower in ‘Apple’ than in ‘Tommy Atkins’. Frequency of membrane failure associated with lenticels increased during development and was consistently higher in ‘Apple’ than in ‘Tommy Atkins’. The lower rate of cuticular deposition, the higher strain releases on excision, isolation and wax extraction and the weaker cuticle account for the high russet susceptibility of ‘Apple’ mango

Lenticels are sites of initiation of microcracking and russeting in ‘Apple’ mango

The mango cultivar ‘Apple’ is an important fruitcrop in Kenya, but it is highly susceptible to russeting. The objective was to establish whether lenticels predispose cv. ‘Apple’ mango to russeting. Fruit mass and surface area increased in a sigmoidal pattern with time. The frequency of lenticels per unit surface area decreased during development. The number of lenticels per fruit was constant. Lenticels were most frequent in the apex region and least common in the cheek and nak (ventral) regions. The cheek region also had lenticels with the largest core areas, whereas the lenticel core areas in the apex region were significantly smaller. Microscopy revealed stomata became covered over with wax deposits at 33 days after full bloom (DAFB). By 78 DAFB, periderm had formed beneath the pore. At 110 and 161 DAFB, cracks had developed and the periderm had extended tangentially and radially. The presence of lenticels increased the strain released upon excision of an epidermal segment, further strain releases occurred subsequently upon isolation of the cuticle and on extraction of the cuticular waxes. The number of lenticels per unit surface area was negatively correlated with the fruit surface area (r2 = 0.62 **), but not affected by fruit size. Mango cv. ‘Apple’ had fewer, larger lenticels and more russet, compared with ‘Ngowe’, ‘Kitovu’ or ‘Tommy Atkins’ mango. In cv. ‘Apple’, the lowest lenticel frequency, the largest lenticels and the most russeting occurred at a growing site at the highest altitude, with the highest rainfall and the lowest temperature. Moisture exposure of the fruit surface resulted in enlarged lenticels and more microcracking of the cuticle. Our results establish that russeting in ‘Apple’ mango is initiated at lenticels and is exacerbated if lenticels are exposed to moisture

Bagging prevents russeting and decreases postharvest water loss of mango fruit cv. ‘Apple’

In Kenya, the mango (Mangifera indica L) cultivar ‘Apple’ is commercially important but it often suffers excessive russeting, which both compromises its appearance and impairs its postharvest performance. Together, these effects seriously reduce its market potential. Exposure to surface moisture is implicated in russeting of cv. ‘Apple’ mango. The objective was to establish the effect of bagging on russeting. Developing fruit were bagged at the onset of the exponential growth phase, using brown paper bags (Blue star®). Un-bagged fruit served as controls. The brown paper bags were selected because of their high permeance to water vapor. At harvest maturity, bagged fruit were larger, less russeted and had smaller lenticels than un-bagged control fruit. Staining with aqueous acridine orange in conjunction with fluorescence microscopy revealed numerous microcracks and larger lenticels on un-bagged control fruit but these were not evident on bagged fruit. Postharvest mass loss (principally water loss) of bagged fruit was lower than of un-bagged control fruit. In the un-bagged control fruit, the skin's water permeance increased as the russeted surface area increased (r2 = 0.88 **). Fruit skins were less permeable to water vapor than the brown paper bags. The brown paper bags contributed not more than 4.2 to 9.1% of the total in-series diffusion resistance of skin + bag. The masses of isolated cuticular membranes, and of dewaxed cuticular membranes, and of wax per unit surface area were higher for un-bagged control fruit than for bagged fruit. Bagged fruit were also greener and showed less blush. There was little difference in skin carotenoid content between bagged and un-bagged control fruit, but skin anthocyanin content was lower in bagged fruit. The rates of respiration and ethylene evolution of bagged fruit were lower than those of un-bagged control fruit. There were no differences between bagged and un-bagged control fruit in their organoleptic and nutritional properties including titratable acidity, total soluble sugars, sucrose, glucose, fructose, vitamin C and calcium content. In conclusion, bagging decreased russeting and increased postharvest performance of fruit of mango cv. ‘Apple’

Surface Moisture Induces Microcracks and Increases Water Vapor Permeance of Fruit Skins of Mango cv. Apple

Exposure to surface moisture triggers cuticular microcracking of the fruit skin. In mango fruit cv. apple, microcracking compromises postharvest performance by increasing moisture loss and infections with pathogens. This study reports the effects of exposing the fruit’s skin to surface moisture on the incidence of microcracking and on water vapor permeance. Microcracking was quantified microscopically following infiltration with a fluorescent tracer. Water mass loss was determined gravimetrically. Moisture exposure increased cuticular microcracking and permeance. During moisture exposure, permeance increased over the first 4 d, remained constant up to approximately 8 d, then decreased for longer exposure times. Fruit development followed a sigmoid growth pattern. The growth rate peaked approximately 103 days after full bloom. This coincided with the peak in moisture-induced microcracking. There were no increases in water vapor permeance or in microcracking in control fruit that remained dry. When experimental moisture exposure was terminated, microcracking and water vapor permeance decreased. This suggests a repair process restoring the barrier properties of the fruit skin. Histological analyses reveal a periderm forms in the hypodermis beneath a microcrack. Our study demonstrates that surface moisture induces microcracking in mango cv. apple that increases the skin’s water vapor permeance and induces russeting

Surface Moisture Induces Microcracks and Increases Water Vapor Permeance of Fruit Skins of Mango cv. Apple

Exposure to surface moisture triggers cuticular microcracking of the fruit skin. In mango fruit cv. apple, microcracking compromises postharvest performance by increasing moisture loss and infections with pathogens. This study reports the effects of exposing the fruit’s skin to surface moisture on the incidence of microcracking and on water vapor permeance. Microcracking was quantified microscopically following infiltration with a fluorescent tracer. Water mass loss was determined gravimetrically. Moisture exposure increased cuticular microcracking and permeance. During moisture exposure, permeance increased over the first 4 d, remained constant up to approximately 8 d, then decreased for longer exposure times. Fruit development followed a sigmoid growth pattern. The growth rate peaked approximately 103 days after full bloom. This coincided with the peak in moisture-induced microcracking. There were no increases in water vapor permeance or in microcracking in control fruit that remained dry. When experimental moisture exposure was terminated, microcracking and water vapor permeance decreased. This suggests a repair process restoring the barrier properties of the fruit skin. Histological analyses reveal a periderm forms in the hypodermis beneath a microcrack. Our study demonstrates that surface moisture induces microcracking in mango cv. apple that increases the skin’s water vapor permeance and induces russeting