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Fresh fruit microstructure, texture and quality
Delilah F. Wood, Syed H. Imam, William J. Orts and Gregory M. Glenn
U.S. Department of Agriculture, Agricultural Research Service, Western Regional Research Center,800 Buchanan St. Albany, CA 94710
ABSTRACT
Fresh-cut produce has a huge following in todays supermarkets. The trend follows the need to decrease preparation timeas well as the desire to follow the current health guidelines for consumption of more whole heart-healthy foods.
Additionally, consumers are able to enjoy a variety of fresh produce regardless of the local season because produce is
now shipped world-wide. However, most fruits decompose rapidly once their natural packaging has been disrupted by
cutting. In addition, some intact fruits have limited shelf-life which, in turn, limits shipping and storage. Therefore, abasic understanding of how produce microstructure relates to texture and how microstructure changes as quality
deteriorates is needed to ensure the best quality in the both the fresh-cut and the fresh produce markets. Similarities
between different types of produce include desiccation intolerance which produces wrinkling of the outer layers,
cracking of the cuticle and increased susceptibility to pathogen invasion. Specific examples of fresh produce and their
corresponding ripening and storage issues, and degradation are shown in scanning electron micrographs.
Keywords: Fresh-cut roduce, fruits and vegetables, microstructure, parsley, carrot, broccoli, kiwifruit, mushroom
1. INTRODUCTIONThe origin of plant tissue components during development and maturation determines the nomenclature used for the
resulting component1. Thus, the edible portion of a plant encompasses a wide array of tissue types
2and constitutes
vegetative and reproductive tissues. Leafy greens (lettuce, spinach, parsley) stems (celery, rhubarb), roots (carrot,
radish), tubers (potatoes), and bulbs (onion, garlic) are all vegetative tissues. Flower buds (broccoli florets, artichokes),fruits (bananas, squash, green beans), seeds (mature beans, almonds), and grains (wheat, corn) are examples of
reproductive tissues.
Intact fruits and vegetables have inherent preservation in their designs. As a first line of defense, a fruit or vegetable is
protected from degradation by a resilient outer layer which forms a skin. The skin which consists of several cell layers, is
covered by a thin, continuous, waxy layer, the cuticle3
The cuticle slows dehydration and serves as an effective barrier topathogens and insects
4and thus, serves as a natural package. Beneath the cuticle lies the epidermis consisting of cells
which may possess thick outer cell walls. The skin may also contain additional cell layers with thickened cell walls. Cell
walls aid in dehydration prevention and provide structure to edible plant tissues and to mushrooms. Unlike plants,
mushrooms are fungi which are non-photosynthetic organisms. Edible mushrooms are comprised of a mass of hyphae
joined together to form a basidiocarp, or fruiting body, the edible portion. The cell walls of both plants and fungi arerigid and rely on appropriate water potential to provide turgor pressure to the cells allowing them to retain their shapes
and their fresh characteristics.
Fruits which are consumed ripe are usually harvested green to allow time for transport to the consumer. Generally,
ripening is due to a loss of firmness in texture due to cell wall depolymerization and the dissolving of the middle
lamella5. The middle lamella, the substance between adjoining, adjacent cell walls, functions as the glue that holds
cells together. Thus, as the middle lamella dissolves, the adhesion between cells decreases and continues to decrease overtime resulting first in ripening and then in aging
6. Fruit firmness is also related to cellular characteristics such as cell wall
strength, cell turgor, the number and sizes of intercellular spaces as well as intercellular adhesion7. Intercellular adhesion
plays a significant role in the texture of produce. The physiological and chemical processes ultimately responsible for
ripening and aging have been well-documented8. Many papers on ripening have been published and these provide insight
into some of the physiological changes that continue to occur as the fruit decays.
Dehydration accelerates fruit decay and is largely due to vapor diffusion through stomata, the gas exchange system of the
plant, and the cuticle9. Therefore, storage in a dry atmosphere can be detrimental and accelerate dehydration, however, a
Scanning Microscopy 2009, edited by Michael T. Postek, Dale E. Newbury, S. Frank Platek, David C. Joy,Proc. of SPIE Vol. 7378, 73781J 2009 SPIE CCC code: 0277-786X/09/$18 doi: 10.1117/12.821351
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moist atmosphere may make it easier for the entry of pathogens into stomata or other openings in the plant surface.
Produce shows an increase in susceptibility to postharvest diseases and infestation during prolonged storage partly due to
ongoing physiological changes that enable pathogen development on or in the fruits10
. Thus, protective mechanisms fail
over time and when any one of the protective systems are compromised, degradation and a loss of quality will occur.
Therefore, an understanding of each produce system is essential in developing methods of handling fresh produce.
A few selected plant materials are included by discussing a brief survey of the literature and by showing scanning
electron micrographs of specific commodities which were purchased at the local supermarket. We hope to acquaint thereader with a general idea of structure as it relates to the changing of microstructure of produce over time. This paper is
excerpted, in part, from a book chapter11
.
2. MATERIALS AND METHODSFresh material was prepared for microscopy as soon as it was purchased. Aged material was purchased fresh and then
allowed to age at ambient conditions for a week or more prior to preparation for microscopy. Plant material was fixed informalin-acetic acid-ethanol
12or in a solution containing 3% glutaraldehyde, 2% formaldehyde in sodium cacodylate
buffer, pH 6.9. The aldehyde-fixed tissue was then post-fixed in aqueous 1% osmium tetroxide13
and dehydrated in a
graded series of ethanol12, 13
. The samples were then either cryofractured in liquid nitrogen14
or sliced under 100%ethanol, critical point dried in a Tousimis Autosamdri 815 (Tousimis, Rockville, MD), sputter coated with gold-
palladium in a Denton Desk II sputter coating unit (Denton Vacuum, Moorestown, NJ), and observed and photographed
in a Hitachi S-4700 field emission scanning electron microscope (Japan).
3. RESULTS AND DISCUSSION3.1 Leafy greens Italian parsleyThe turgor of plant cells and cell walls is essential in providing the characteristic textural crispness attributes to leafygreens such as parsley. Fresh, Italian parsley has a characteristic, triangular stem (Fig. 1a) which carries water up to the
leaves and can be placed in water post-harvest to increase the shelf life. The lower (abaxial) leaf surface of fresh parsley
(Fig. 1b) reveals plump, epidermal cells indicating that they have adequate turgor and are fully hydrated. The vascular
bundles are distinct and stomata are evident. In contrast, the abaxial surface of the aged leaf (Fig. 1c) has an irregular,
wrinkled surface and the vascular bundle is not distinct as it blends with the extensive folds in the epidermis. The fresh
parsley leaf has a relatively flat epidermis, the vascular bundle has plump, rounded cells (Fig. 1d) and the stomata are in
evidence (Fig. 1e). However, early signs of degradation are also apparent in the fresh leaf in areas where the epicuticular
wax has started to peel and crack (Fig. 1f). The epidermis of the aged leaf (Fig 1g) has shriveled and wrinkled due todehydration also has surface lesions (Fig 1g, h) and other disruptions which allow the proliferation of bacteria (Fig 1i, j).
The bacterial colony was in association with a fibrous substance, probably a biofilm. Biofilms, exudates of some types of
bacteria, provide adherence properties to bacteria allowing them to colonize tissues. Biofilms are typically composed of
highly hydrated polysaccharides15, 16
. The polysaccharides are difficult to preserve using aqueous fixation techniquesused in preparation for scanning electron microscopy because portions of the polysaccharide dissolve in the aqueous
fixative. The insoluble remains of the polysaccharides lose definition and tend to clump together forming fiber-like
structures. The larger fibrous components are part of the parsley leaf which disintegrated from the microbial activity.
Roots Carrot
Carrots have a long shelf life but the shelf life can be shortened by root-rotting pathogens, such as Chalara elegans, a
pathogen common in the Fraser Valley of British Columbia, Canada17
. Carrots may be harvested at any size,however, carrots harvested too late tend to be woody and are unacceptable to consumers. Small, ready-to-eat carrots
were chosen for this study because of their convenience to consumers. Ready-to-eat carrots are peeled, scrubbed,packaged and shipped in bags. The surfaces of prepackaged carrots, whether fresh or aged, consist of crushed cells
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Figure 1. Italian parsley. A cross section of fresh, triangular-shaped stem (a). Lower leaf surfaces show the outer epidermis of fresh
(b, d, e, f) and aged (c, g, h, i, j) leaves. A fresh leaf has distinct vascular bundles (b), epidermal cells and stomata. An aged leaf hasextensive wrinkling or folding, epidermal cells have lost turgor pressure, and the vascular bundle is indistinct (c). A fresh leaf showingplump vascular bundle cells (d) and open stomata (e). Vascular bundle cells that have adequate turgor (f); the "flaky" appearance isdue to epicuticular wax which has cracked. Aged leaf showing folds and an irregular surface blemish ( g, h). The surface of an agedleaf showing a vascular bundle and wrinkled epidermal cells and stomata ( i). The dark, fibrous area is a bacterial colony enlargementshown inj associated with possible biofilm (arrows) (j). Large fibers (*) are decomposing plant material. VB, vascular bundle; S,stomata. Scale bars: a-c, 500 m; d, 300; e, 100 m; f, 50 m; g, 1 mm; h, 200 m; i 50 m; j, 10 m.
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I h
from the outer cortex (Fig. 2a, b). Peeled and stored carrots often have a white translucent appearance. The whiteappearance is due the dehydration of the shredded cells on the carrot surface
18. Treatment of carrots by slicing or peeling
can cause physical damage, stress and the increased risk of microbial growth. The risk and severity of damage was lesswith gentle handling and cutting with sharp knives
19and was further reduced following vacuum
packaging20
. Fresh (Fig. 2c) and aged (Fig. 2d) carrots have similar microstructure even though the aged carrot hadwilted. Carrots are resilient and the aged carrot was almost completely rehydrated during aqueous fixation for scanningelectron microscopy, therefore, most of the cell walls in the aged carrot appear turgid. However, some aged carrot cellshad broken cell walls. Thus, large gaps in the tissue (Fig. 2e) and collapse were apparent (Fig. 2f). Broken cell walls andcollapsed cells were not apparent in the fresh sample (Fig. 2g).
Figure 2. Prepackaged peeled and washed baby carrot. Outer surface of fresh carrot (a). Cross section of the outer layers of freshcarrot showing crushing of the cells due to initial processing steps of peeling and washing (b). Cross section of fresh carrot (c). Crosssection of aged carrot showing cell wall breakage and cell collapse (d, arrows). Aged carrot, close view showing the effects of cellwall breakage (e). Aged carrot, close view showing cell wall breakage and cell collapse (f). Fresh carrot cross section showing no cellwall breakage or collapse (g). Scale bars: a, 200 m; b, 50 m; c, d 1 m; e-g, 200 m.
3.3 Flower buds Broccoli
Broccoli is harvested when the immature flowering heads are growing rapidly21
. Broccoli should be consumed within amonth if stored at 0 C or 3 days if stored under ambient conditions
22. The floret buds (Fig. 3a, b) are more susceptible to
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decay than the stalk and quickly show dehydration and superficial mold growth (Fig. 3a-c) upon aging. Undesirable
changes in color is accelerated by microbial growth as the pH decreases resulting in the conversion of chlorophyll to
pheophytin23
. Fresh broccoli epidermis is smooth because the cells are turgid (Fig. 3c). The aged epidermis is dehydrated
and moldy (Fig. 3d). Cross sections of fresh (Fig. 3e) and aged (Fig. 3f) stems are similar at low magnification and show
minor differences at higher magnifications (Fig. 4a, b). Intercellular spaces appear smaller and cell walls appear thickerin the fresh (Fig. 4a) and than those in the aged (Fig. 4b) sample. Cell walls in the aged carrot were wavy, indicative ofdehydration and less turgidity.
Figure 3. Scanning electron micrographs of broccoli florets. Aged broccoli flower buds showing dehydration and fungal growth onthe epidermis (a, b).Epidermis of a fresh stalk (c) showing smooth, hydrated cells and an aged stalk (d) showing dehydration andfungal growth. Cross section of a fresh stalk showing tissue organization (e). Cross section of an aged stalk (f). C, cortex; CW, cellwall; IS, intercellular space; P, pith; VB, vascular bundles. Scale bars: a, e, f, 1 mm; b, 500 m; c, d, 200 m.
3.4 Fruit Kiwifruit
Kiwifruit is harvested mature and unripe when the fruit is firm and softening occurs during the ripening process.
Microscopy has been used to document changes occurring during ripening at the microstructure level and correlated with
penetrometer tests to measure the hardness of the whole fruit28
. During ripening, the angular cells become more rounded
and starch granules degrade. Kiwifruit consists of distinct regions including the outer and inner pericarp and
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a
Figure 4. Scanning electron micrographs of broccoli florets. Cortex of the fresh stalk showing thickened parenchyma cell walls (a),turgid cells, and small intercellular spaces. Aged cortex cells of the stalk (b) showing somewhat wavy cell walls, less apparent turgor,thin cell walls and large intercellular spaces. CW, cell wall; IS, intercellular space. Scale bars: a, b, 10 m.
the core. The rate of change and composition of cell wall components differs in each tissue region. Cell walls in all
tissues change considerably in structure and become thicker during ripening24, 25
. The thickening of the cell walls
coincides with the solubilization of pectin26
. Pectin hydrolysis and the modification of hemicelluloses are also involved
in ripening and fruit softening in papaya27
.
Intercellular air space volumes were shown to increase with ripening using morphometric studies. The volume increase
correlated with the disintegration of the middle lamella, thus, the middle lamella was replaced by air. However, as fruit
became very ripe, intercellular spaces decreased probably due to the loss of tissue integrity and collapse. Cellularstructure changes are, however, tissue dependent. Intercellular air spaces did not increase in the locular region in the
inner pericarp with ripening. Cell walls exhibit less change, i.e., cell wall thickening was less, in the locular region than
those of other tissues. Intercellular space did increase, however in the locular wall28
.
The desirable soft texture of kiwifruit is due to swelling of the cell walls and weakening of the middle lamella24
. The
changes that occur in kiwifruit during ripening are similar to those in other fruit. The ripening changes in kiwifruit,
however, occur before the start of ethylene production as is common in climacteric fruits. Tensile strength measurementswhich show structure changes indicated that cell walls in unripe kiwifruit ruptured. Whereas, as the tissue got softer, the
stress breaks occurred around cells, indicating that the adhesive substance between cells, the middle lamella, was weaker
than the cell walls in ripe fruit29
.
Kiwifruit skin has a multitude of trichomes, which is responsible for the fuzzy appearance and feel to the fruit. Thetrichomes are of two size classifications. The large trichomes are about 2.5 mm long; the twisted short trichomes are
about 200 m long (Fig. 5a-c). Platelets also occur on the surface of the skin and might be flattened trichomes or
outgrowths of the epidermis rather than wax1. In fresh kiwifruit, the platelets appear to be closely appressed to the
surface of the fruit (Fig. 5a) whereas those in the aged fruit are more at an angle from surface of the fruit (Fig. 5b,c).
The outer pericarp of fresh kiwifruit (Fig. 5d) contains a mixture of large and small cells. The outer cells, just beneath the
skin are radially compressed in the fresh (Fig. 5d) and in the aged (Fig. 5e) fruits. Cells further into the outer pericarp ofthe aged fruit show increased compression (Fig. 5e). Starch granules in the fresh (Fig. 5d) kiwifruit have virtually
disappeared in the aged (Fig. 5e) fruit in agreement with Hallett et al in their observations of starch degradation30
.
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(
) Figure 5. Scanning electron micrographs of kiwifruit. Fresh fruit showing the various types of trichomes covering the skin surface (a)without evidence of fungal growth. Aged fruit showing the short, twisted trichomes and platelets (b) covering the skin surface; theplatelets might be flat trichomes since they do not appear to be wax and appear to be outgrowths of the epidermis. An extensivegrowth of fungal hyphae and the lifting of the surface platelets is also evident. Aged fruit showing the base of a long trichome andthe growth of fungi (c). Fresh cross section of the outer pericarp showing a mixture of large and small cells in the flesh and the radiallycompressed cells near the surface (d). Aged cross section showing areas of further compression resulting from aging; note the ovals(e). S, starch granules. Scale bars: a, 1 mm; b, c, 200 m; d, e, 1 mm.
3.5 Fungi mushroomA fresh, white button mushroom of high quality has a tightly closed cap (pileus), the gills are covered by a membrane
(velum), the stalk is short and the mushroom is white. Browning, which might occur during storage, of the mushroom is
unacceptable. Mechanisms associated with browning range from rough handling to microbial activity. Low-dose
irradiation was shown to be effective in the prevention of browning and microbial growth31
. As the mushroom ages, it
continues to grow and creates undesirable quality changes. The stipe grows, the velum stretches and breaks, and thepileus opens to expose and release the spores on the surfaces of developing gill tissue. The broken velum leaves a ring of
tissue on the stipe, the annulus32
. Part of the pileus structure is shown in Figure 6a. Spores form on the inner portions of
the gills shown by the arrows (Fig. 6a); the remaining pileus and stipe are composed of hyphal masses (Fig. 6b-d). Freshmushroom hyphae are tightly compacted, have smooth, plump, hydrated cell walls and evident cytoplasm (Fig. 6b, d).
More space is apparent between hyphal strands in the aged mushroom (Fig. 6c) and cell walls are somewhat collapsedand wrinkled (Fig. 6c, e), due to dehydration stress. Cytoplasm is no longer evident in the aged mushroom (Fig 6e)
agreeing with findings of Braaksma et al.32
where growth of the pileus was reported to occur by
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Figure 6. Scanning electron micrographs of white button mushroom. Cross section of the cap showing the cap body and the gills ( a),on the surfaces of which spores are formed (arrow). Fresh mushroom hyphae (b) showing the close appression of the hyphae withlittle air space. Aged mushroom hyphae (c) showing the separation of hyphae and resulting air spaces. Fresh mushroom hyphaeshowing evident cytoplasm and plump cell walls (d). Aged hyphae showing collapsed, wrinkled hyphae (e) indicative of dehydration.
Scale bars: a, 500 m; b, c, 100 m; d, e, 10 m; f, g, 20 m.
expansion of the vacuoles which would push the cytoplasm to the limits of the cell walls. Fresh mushroom spores are
smooth, rounded and immature (Fig. 7a, d) and the majority are covered by a smooth substance (Fig 7d). Spores in the
aged mushroom are ovoid and areas of mesh-like material are apparent (Fig 7c). Close examination of individual spores
shows that they are covered with irregular deposits or precipitates on their surfaces (Fig 7d), probably the remnants ofthe digested and dehydrated smooth, membranous substance apparent in Figure 7c.
4. CONCLUSION4.1 Textural Characteristics
Produce texture is determined by tissue structure and physiology. Microscopy has been used to relate tissue
microstructure and texture by measuring tissue failure under tension followed by observation of the areas of failure using
various microscopy methods. Texture is largely dependent on cell structure and the relationships between cells.
Microstructure studies can show many of the characteristics giving rise to texture in any given produce commodity.
All fruits and vegetables are composed of cells. Cell characteristics include cell structure, composition, size, shape, type,
water content; air spaces between cells; cell wall structure, composition and thickness; and adhesion between cells.Adhesion is determined mainly by the state of the middle lamella. Fruits and vegetables are dynamic, living systems,
thus, structural and textural changes occur continuously. Textural diversity at the cellular level has been studied invarious tissues using tensile strength measurements and correlating the measurements with observation by microscopy.
Examination of the fracture surfaces allowed the detection of differences in tissue strength and juiciness and provided
insight into the cellular basis of plant texture. It also helped to identify specific cell characteristics which influence the
sensory texture attributes of hardness and juiciness33
.
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Figure 7. Scanning electron micrographs of white button mushroom. Immature spores at low magnification (a) from a fresh
mushroom. Mature spores from an aged mushroom at low magnification (b). Immature spores at high magnification from a freshmushroom (c) showing the smoothly covered spores. Mature spores from an aged mushroom ( d) showing the precipitate which isprobably the digested and dehydrated formerly smooth covering seen in h. Scale bars: a, b, 20 m; c, d, 5 m. Soft texture is a desirable characteristic in kiwifruit. In contrast to apple fruit, kiwifruit cells remain in close apposition
whereas apple cells have more intercellular spaces and have very little cell-to-cell contact, thus the intercellular spaces
account for part of the reason the two fruits have such differing textures28
.
4.2 Shelf life extension
Considerable time might elapse between harvest and the consumers table because fresh produce is shipped world-wide.
Thus, cold storage and commodity-specific packaging is essential for the protection and preservation of fruits and
vegetables. Minimally processed, i.e., those that are cut into bite-sized pieces or ready-to-eat salads, require modified
atmosphere packaging and controlled atmosphere storage34
.
At the consumer level, dehydration can be controlled by applying water sprays and using open refrigeration in the
supermarket. Refrigeration alone is insufficient to meet the demands of todays marketing techniques. Therefore, otherpost-harvest treatments are needed to provide an increase of storage life. Edible coatings, such as waxes, and those which
also include anti-browning agents form a semi-permeable barrier to air to control respiration and prolong shelf-life 35, 36.
Due to increasing concern of unacceptable residues on fruit surfaces following chemical treatments, a number of
chemical-free post-harvest techniques have been investigated to increase shelf life.
Post-harvest heat treatment has been in commercial use for many years37
, however, the technology is still being perfected
for different commodities. Short hot water rinses were investigated as a method of reducing post-harvest losses of
cantaloupe (Galia melon, Cucumis melo cv. reticulatus)38
and sweet peppers39
. Scanning electron microscopy showed
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that the fruit surfaces were free of debris and fungal spores and that superficial cracks in the epidermis were sealed. Hot
water immersion proved to be effective in disinfestations and maintenance of fruit quality of Valencia orange40
. Light
and electron microscopy showed that there was little effect on surface waxes of the orange fruit as a result of heating.
Mandarin oranges were heat-treated for 3 min prior to storage at temperatures ranging from 50 C to 58 C in the
assessment of temperature effectiveness in storage performance. Treated and control fruit surfaces were compared bySEM observation of mandarins. Heat treatment at 50-54 C smoothed the granular waxy surfaces of mandarins, at 56 Cpartly removed the wax and treatment at 58 C completely removed the wax
41. Hot water brushing of organic citrus fruits
reduced postharvest decay and scanning electron microscopy showed that the epicuticular waxes had been smoothed by
the treatment and the cracks and stomata were sealed by the wax, thus, reducing pathogen entry sites42
.
Heat treatment of heat-sensitive mango cultivars was ineffective due to the resulting tissue damage. Such heat-related
injuries were investigated using scanning and transmission electron microscopy where cuticle and exocarp ruptured and
exposed internal cells. Cell walls of the mesocarp were convoluted and thickened and starch granules still remained in
the tissue suggesting that the carbohydrate metabolic enzymes had been disrupted43
.
Other non-toxic shelf life extension methods include treatment with carbon dioxide followed by refrigerated storage.
Firmness was increased and decay susceptibility decreased (compared to those at harvest) in strawberries followingstorage at 0 C. The positive effects were enhanced by treatment with CO2 followed by storage at 0C. Examination ofthe fracture surfaces of strawberry following tensile testing indicated that the primary mode of tissue failure was cell-to-cell debonding, not cell breakage, in both air and CO2-treated fruit
44. An increase in firmness over harvest is common in
fruit that is stored at 0 C. The mechanism is not well understood but is theorized to be due to an increase in viscosity ofpectin because no change in structure or chemistry is observed
45, 46.
4.3 Quality and Specific Disorders
Electron microscopy demonstrated the withered juice sac or dry pulp disorder, where the soluble solids content is
higher in the rind than in the pulp, in Ponkan mandarin orange at the cellular level. Prior to the onset of symptoms of the
disorder, nuclear divisions occurred in the cells of the flavedo (the colored part of the rind). The appearance of symptoms
corresponded to further changes in the flavedo cells, the enlargement of the cells and their vacuoles and a decrease in thecytoplasm content. The tonoplast disappeared in latter stages of the disorder with only a nucleus and small amounts of
cytoplasm remaining. Thus, the disorder was caused by changes occurring in the rind: cell division, growth andsenescence, which caused water to be translocated to the rind due to a water potential gradient. Treatment with
gibberellic acid, a plant hormone which slows senescence, prior to storage delayed the onset of the disorder47
.
Water loss, determined by weight loss, was found to be a non-destructive predictor of chilling injury in grapefruit and
lemon. Chilling injury appears as distinct, swollen areas with pitting and cuticular damage. Prior to the appearance of the
gross symptoms of chilling injury, scanning electron microscopy revealed calcium oxalate crystals growing inside cracks
which had developed around the stomata48
.
Ultrastructure and electrophoresis were useful in determining the reason for degreening inhibition of bananas. In
bananas, degreening is inhibited above 24 C, ripening at higher temperatures results in retaining thylakoid membranesand a delayed breakdown in chlorophyll b as well as a reduced break-down of pigment-protein complexes. Retention ofthylakoid membranes is an important factor in the failure of Cavendish bananas to degreen when ripened at tropicaltemperatures
49. Fruit softening in banana is the result of the coordinated degradation of pectin, hemicelluloses and starch
in the banana pulp50
and was shown using stress-relaxation probe measurements to describe the changes in physical
properties during softening of the banana fruit pulp.
4.4
Microscopy as a tool
Microscopy can be used as an effective tool to understand and explain the physiological, chemical and structural
processes involved in determining parameters that indicate fresh produce quality. Correlating physical measurements
with findings from microscopy may be used to illustrate where improvements are needed in a specific commodity.
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