Mango

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Mango (Mangifera indica L.), “The King

of Fruits”—An Overview
a a a
R.N. Tharanathan , H.M. Yashoda & T.N. Prabha
a
Department of Biochemistry and Nutrition , Central Food
Technological Research Institute , Mysore, India
Published online: 06 Feb 2007.
To cite this article: R.N. Tharanathan , H.M. Yashoda & T.N. Prabha (2006) Mango (Mangifera
indica L.), “The King of Fruits”—An Overview, Food Reviews International, 22:2, 95-123, DOI:
10.1080/87559120600574493
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Food Reviews International, 22:95–123, 2006
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ISSN: 8755-9129 print / 1525-6103 online
DOI: 10.1080/87559120600574493
Mango (Mangifera indica L.), “The King of

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Food Reviews International
International, Vol. 22, No. 02, February 2006: pp. 0–0
Fruits”—An Overview
R.N. THARANATHAN, H.M. YASHODA, AND T.N. PRABHA

Mango
R.N. Tharanathan,
(Mangifera H.M.
indicaYashoda,
L.) and T.N. Prabha
Department of Biochemistry and Nutrition, Central Food Technological

Research Institute, Mysore, India
Mango (Mangifera indica L.) is commercially the most important fruit crop of India,
Downloaded by [Iowa State University] at 14:11 08 October 2013
accounting for > 54% of the total mango produced worldwide. Over 30 different vari-
eties of mango are grown, the most important one is Alphonso, which is rated best in
the world. It is known for its strong aroma, intense peel coloration, delicious taste, and
high nutritive value (due to its high content of vitamin C, β-carotene and minerals).
The chemical composition of mango pulp varies with the location of cultivation, vari-
ety, and stage of maturity. There is an increase from 1 to 14% in the starch content
during fruit development, and towards the end of maturity, both reducing and non-
reducing sugars are found to be increasing. The fruit ripening process involves a
series of physiological, biochemical, and organoleptic changes that lead to the devel-
opment of a soft, edible, ripe fruit with desirable qualities. Ethylene, a plant growth
hormone, regulates many aspects of fruit development and cell metabolism, including
initiation of ripening and senescence, particularly in climacteric fruits. Textural soft-
ening, an integral part of ripening of almost all fruits, is a major quality attribute that
determines consumer acceptance. Fruit softening is thus accompanied by molecular-
structural changes in cell wall constituents, which have been studied at both substrate
(polysaccharides) and enzyme (glycanases and glycosidases) levels. Several lines of
evidence have enumerated on compositional and structural modifications in pectic and
hemicellulosic polysaccharides, especially of xyloglucan-type polymers during mango
fruit ripening. Of late, modern biotechnological approaches are paving the way for
healthy propagation and rapid multiplication of valuable geno types and improved
plants, which augment advantages such as non-seasonal, almost year-round produc-
tion and conservation of germplasm for better international exchange. Somatic hybrid-
ization via protoplast fusion could be an alternative to overcome problems such as
difficulties in establishing aseptic mango cultures from mature explants associated
with phenolic browning. In this direction, further biotechnological approaches may be
worth pursuing for sustained mango cultivation.
Keywords Mangifera indica, mango, fruit ripening, textural softening, enzymes,

pectin, carbohydrate polymers, biotechnology
Introduction
Morphologically, the fruit is a seed receptacle developed from an ovary. This definition
encompasses a very wide range of fruit types, which are generally classified into simple,
aggregate and composite fruits. Being commercially valuable food crops, fresh as well as
processed fruits form an important part of our diet. Fruits provide useful food reserves and
Address correspondence to R.N. Tharanathan, Department of Biochemistry and Nutrition,

Central Food Technological Research Institute, Mysore 570 020, India. E-mail: tharanathan@
yahoo.co.uk
95
96 R.N. Tharanathan, H.M. Yashoda, and T.N. Prabha
are an important source of essential micronutrients, vitamins, and other phytochemicals,

but they are generally low in protein and fat. The quality of a fruit is influenced by variety,
nutritional status, and environmental conditions during growth of the parent plant. Fruits
play a very important role in human nutrition, by providing a source of energy, necessary
growth factors, carbohydrates, dietary fiber and antioxidants, which are essential for main-
taining normal health. Fruits also contain a very high percentage of water as their fresh
weight.
Fruits are classified into tropical, subtropical, and temperate fruits based on their geo-
graphical distribution in different climates. Fruits are classified as either climacteric or
non-climacteric based on the pattern of respiration and ethylene biosynthesis during ripen-
ing. Fruits, when harvested at optimum mature stage, are self-sufficient with their own
reserve food material to maintain a short post-harvest life when detached from the parent
plant.
In climacteric fruits, the rate of respiration and ethylene production are minimal at
post-harvest maturity stage and rise dramatically to a climacteric peak, at the onset of rip-
ening, after which it gradually declines.(1) Unlike the non-climacteric fruits, which are
incapable of continuing the ripening process after their detachment from the parent plant,
the climacteric fruits, when harvested at mature stage, can be ripened off the parent plant.
Non-climacteric fruits produce very small quantity of endogenous ethylene, do not
respond to external ethylene treatment, and show a gradual decline in their respiration pat-
tern and ethylene production throughout the post-harvest ripening process.(2)
Mango—“The King of Fruits”

Mango (Mangifera indica Linn.), a commercially important tropical fruit, morphologi-
cally belongs to the subtype indeliquescent drupe, and contains a single large seed sur-
rounded by fleshy mesocarp. Mango, a dicotyledonous fruit of the family Anacardiaceae,
originated in the Indo-Burmese region.(3,4) Mango is known to be the most important trop-
ical fruit of Asia, grown commercially in more than 87 countries. Mango currently ranks
fifth in total production among major fruit crops world wide.(5) The world production of
mangoes is estimated to be over 23.4 × 106 MT per anum. India ranks first among world’s
mango producing countries, accounting for 54.2% of the total mangoes produced world-
wide. Other prominent mango producing countries are China, Thailand, Indonesia, Philip-
pines, Pakistan, and Mexico (Table 1). Between 1971 and 2001, the production of
mangoes, worldwide has increased by nearly 50%. Much of this increased production has
occurred outside the traditional centers of mango cultivation, viz., southeast of USA, Cen-
tral and South America, Africa, Australia, South-east Asia, Hawaii, Egypt, and Israel; and,
a significant proportion of the increased mango production is for export markets. At
present, mangoes are widely available not only in tropics and sub-tropics, but also in
North America, Japan and Europe, both in fresh as well as processed forms.(6)
Mango is commercially the most important fruit crop of India, with more than a thou-
sand varieties known so far.(7) The cultivation of mango in India covers an area of 12.2 ×
106 hectares. The state, Andhra Pradesh, has the highest production, with 0.24 × 106 hectares
dedicated to mango cultivation, and producing 2.9 × 106 MT per year. Uttar Pradesh,
Bihar, Karnataka, Himachal Pradesh, Maharastra, Orissa, Tamil Nadu, and West Bengal
are the other major mango producing states. Mango cultivars may be classified into two
groups, Indian and Indo-Chinese, based mainly on peel pigments and sensory characteris-
tics of the pulp. Most of the Indian varieties possess strong aroma and intense peel colora-
tion, characterized by attractive fragrance, delicious taste, and high nutritial value, owing
Mango (Mangifera indica L.) 97
Table 1
Production status of mango in the world
Mango production statistics (1000 MT)
Countries 1981 1991 1999
World 13,454 15,700 23,852
India 8516 9500 12,000
China 341 595 2150
Mexico 561 800 1538
Thailand 509 903 1250
Philippines 367 348 950
Pakistan 547 760 917
Nigeria ⎯ ⎯ 731
Indonesia 444 640 605

Brazil 600 546 600
Egypt 123 208 231
Source: FAO year book.(5)
to high amounts of vitamin C, β-carotene, and minerals.(8) In 2001–2002, India exported

45.41 × 103 MT of mango as fresh fruits valued at 791.3 million rupees, and 455.49 × 103
MT of sliced and dried mango, valued at 18.2 million rupees. This accounted for 41.5% of
the income due to fresh and processed fruit export.(9) India exports fresh mangoes to more
than 50 countries. Over 90% of exports are directed to seven countries viz., UAE, Saudi
Arabia, Kuwait, UK, Singapore, Netherlands, and Bangladesh.(10) Processed products are
likely to have better prospects than fresh fruit in the export market mainly because of the
low shelf life of the fruit and also because its availability is restricted to one season.
Mango fruits have been utilized for a long time at every stage of growth. The raw
fruits are utilized for products like pickles, chutney or mango sauce, amchoor (raw mango
powder), and green mango beverage (panna), whereas ripe fruits are used in making pulp,
juice, nectar, squash, mango leather, frozen and canned slices, jam, ready-to-serve bever-
ages, mango puree, mango cereal flakes, mango powder, mango toffee, and mango fruit
bars.(11) Mango nectar constitutes the major item, followed by mango chutney, pickles,
and mango slices in brine. Since India is the richest source of quality mango varieties in
the world, India could potentially increase its export of mangoes through proper storage,
packaging, and marketing practices.
Mango is one of the most commonly eaten fruits in tropical countries around the
world. About 30 varieties are grown on a commercial scale in different states in India. The
most important mango varieties cultivated are Alphonso (Badami), Banganapalli (Banes-
han), Bangalora (Totapuri), Bathua, Bombay Green (Bhojpuri), Chousa (Khajari), Dashe-
hari (Dasheri), Fajri, Gulabkhas, Himsagar, Kesar, Krishnabhog, Langra (Langarhi),
Jamadar, Mallika, Mankurad, Mundappa, Mulgoa (Mulgoba), Neelam, Pairi (Paheri),
Rajapuri, Suvarnarekha (Swarnarekha), and Vanraj.(12,13) Alphonso is the leading com-
mercial variety and rated best in the world; it is known by different names in different
regions, viz., Badami, Gundu, Khader, Appas, Happus, and Kagdi. The fruit of this variety
is medium to large in size, ovate oblique in shape with a prominent ventral shoulder and
orange yellow in color (Fig. 1), with each fruit weighing between 225–325g. The fruits,
Figure 1. Mango tree bearing fruits.
attractive to the consumer by their excellent fruit quality and good keeping quality, are
characterized by thin skin, soft flesh with a low fiber content and sweet aroma. The flavor
is captivating and the taste is high quality with an excellent sugar/acid ratio.(14) Maharastra,
Gujarat, Madyapradesh, and Karnataka are the major producing states of this variety in
India.
Botanical Aspects
The genus Mangifera belongs to the order sapindales in the family Anacardiaceae, which is
a family of mainly tropical species with 73 genera (C. 850 species). The word “mango” orig-
inated as early as 16th century from the ancient Tamil word ‘mangai‘.(15) Based on taxo-
nomic and recent molecular evidence it is now apparent that mango probably evolved within
a large area including northwestern Myanmar, Bangladesh, and Northeastern India.(6)
The mango tree is erect, 30 to 70 ft (10–40 m) high, an arborescent, evergreen with
symmetrical, round and broad canopy, or more upright with a relatively slender crown. Its
color varies between green through yellow to red. The tree is long-lived and mature speci-
mens can survive for more than one hundred years. The flowers (yellowish or reddish) are
borne in inflorescences, which appear at branch terminals in dense panicles of up to 2000-
minute flowers, glabrous or pubescent; inflorescence is pseudoterminal, rigid, and erect
and is widely branched. The minute flowers (5–10 mm in diameter) are monoecious,
polygamous, and hermaphrodite. Both male and perfect flowers are found in an inflores-
cence. The pistil aborts in male flowers. It is believed that the flowers are cross-pollinated
by flies, wild bees, wasps, moths, beetles, etc.(6,11,16)
The mango fruit is a simple, large, more or less compressed, fleshy, resinous drupe. It
varies in size, shape, color, fiber content, flavor, and taste. The most characteristic feature
of mango fruit is the formation of a small conical projection developing laterally at the
proximal end of the fruit, known as the “beak.” The pericarp is distinguished into smooth
exocarp, fleshy mesocarp, and stony endocarp. The exocarp region develops into a leathery
protective skin that is smooth, green and waxy, and when ripe, changes to a pale green or
yellow marked with red, according to cultivars. The mesocarp provides the fleshy edible
pulp, which is firm and can be fibrous or fiber free, with a flavor ranging from turpentine to
sweet. The quality of fruit is based on the scarcity of the fiber and minimal turpentine taste.
Chlorophyll, carotenes, anthocyanins, and xanthophylls are all present in the fruit, although
chlorophyll disappears during ripening, whereas anthocyanins and carotenoids increase
with maturity.(16) The endocarp develops into a thick, tough, leathery, glandular covering of
the seed. The seed is ex-albuminous. It is solitary, stony, hard, large and flat, ovoid-oblong,
or kidney shaped and is surrounded by the fibrous endocarp at maturity. The testa is thin
and papery. The seed may be monoembryonic producing one seedling, or polyembryonic
with several seedlings that are identical but are not always true to the parent type. Almost
all the Indian varieties are monoembryonic and are less viable than other polyembryonic,
which are abundant in Myanmar, Thailand, Indonesia, and the Philliphines.
Growth and Development of Mango Fruit

Flowering in mango is preceded by the differentiation of the flower bud in the shoots. The
period of differentiation varies from variety to variety and is also governed by the local
climactic conditions. The hermaphrodite flowers of the mango inflorescence, after polli-
nation and fertilization, set fruit. Mango trees have an enormous potential to yield fruits.
Mature trees produce up to 1000 inflorescence each with 500–6000 flowers.(17) Fruit set is
usually less than 10% and only 0.1–0.25% reach the harvesting stage.(18) The development
of the mango fruit can be divided into 4 different stages:
1. The juvenile stage (up to 21 days from the day after fruit set) leads to rapid cellular
growth.
2. Stage of maximum growth (21–49 days) results in cell enlargement and maturity.
3. Maturation and ripening stage (49–77 days) represents respiration climacteric and rip-
ening process, and
4. Senescence stage (77th day onwards) is the post-ripening stage, which is prone to
microbial attack followed by death and decay.(19)
The growth patterns of mango, unlike other fruits, appear to take the form of a simple,
rather than double, sigmoid curve.(20) Coloring and softening of the flesh is from seed out-
wards; at this stage, the latter has become surrounded by a cartilaginous and finally, strong
endocarp. These readily observable changes have been used as a means of assessing the
optimal picking date for immediate consumption or for storage.
Mangoes normally reach maturity in 4 to 5 months from flowering. They are har-
vested at a mature green stage, and are kept for normal ripening. When the fruit is fully-
grown and ready for picking, the stem will snap easily with a slight pull. If strong pull is
necessary, the fruit is somewhat immature and should not be harvested. In general,
depending on the variety and environmental conditions, mangoes take 6–10 days to ripen
under ambient temperature and become over-ripe and spoiled within 15 days after har-
vest.(21) As a climacteric fruit, the period of ripening is characterized by a series of bio-
chemical changes initiated by the autocatalytic production of ethylene and increase in the
rate of respiration.(22) Ripening results in the development of characteristic color, aroma,
and taste of the fruits with desirable softening.
Figure 2. An overview of fruit ripening with particular emphasis on textural softening, control
points are at ethylene (1) and post-ethylene (2) levels.
The maturity of mango fruit has been correlated with various physical characteristics
such as surface color, shape, size, shoulder growth and specific gravity, and chemical
parameters such as total soluble solids, titratable acidity, starch, phenolic compounds, and
carotenoids. On the basis of external color and growth, four maturity stages are catego-
rized during mango fruit development: (a) shoulder in-line with the stem end and green-
olive color; (b) shoulders outgrowing the stem end and olive-green color; (c) shoulders
outgrowing the stem end and the light green color; and (d) gradual softening of the flesh.
The stages (b) and (c) were denoted to be the best for harvesting, as the fruits of these
stages developed good flavor and taste upon ripening (see Fig. 2).
Compositional Changes During Development

The chemical composition of mango pulp varies with the location of cultivation, variety, and
stage of maturity. The major constituents of the pulp are water, carbohydrates, organic acids,
fats, minerals, pigments, tannins, vitamins, and flavor compounds (Table 2). During growth
and maturation of mangoes, the period of rapid growth is characterized by an increase in
alcohol-insoluble solids; principally starch accumulation is the main chemical change in the
pulp-tissue. The rate of starch accumulation is rapid at the beginning of fruit growth and
slows down later, but it continues to increase up to maturity. There is an increase from 1 to
14% in starch content in Alphonso mango during development.(23) At initial stages of fruit
development, no systematic trend was observed in the sugar content, but towards the end of
maturity, both reducing and non-reducing sugars were found to be increasing.(24)
Table 2
Food value per 100 g of ripe mango pulp
Calories 62.1–63.7 Cal
Moisture 78.9–82.8 g
Protein 0.36–0.40 g
Fat 0.30–0.53 g
Carbohydrates 16.20–17.18 g
Fiber 0.85–1.06 g
Ash 0.34–0.52 g
Calcium 6.1–12.8 mg
Phosphorus 5.5–17.9 mg
Iron 0.20–0.63 mg
Vitamin A (carotene) 0.135–1.872 mg
Thiamine 0.020–0.073 mg
Riboflavin 0.025–0.068 mg
Niacin 0.025–0.707 mg
Ascorbic Acid 7.8–172.0 mg
Tryptophan 3–6 mg
Methionine 4 mg
Lysine 32–37 mg
Source: Taken in part from Gopalan et al.(39)
The soluble sugars of the fruit pulp consisted mainly of glucose, fructose, and
sucrose.(25) The presence of glucose, fructose, maltose, and xylose was also reported in
ripening mangoes.(26) The total sugar content of mangoes varied between 11.5 and 25%
(fresh weight) and up to 15% of the fresh pulp of green, mature fruits was starch. In the
developing mango fruits, acidity increased at the early growth phase, reached a peak, and
then declined gradually until harvest. In the Alphonso mango, the acidity reached maxi-
mum (4.2–4.4%) in about 7 weeks and declined slowly to around 2.7–2.8% at the time of
harvest.(27) Pectin increases from the fifth week of fruit set until the stone is formed; there-
after, the pectin content falls.(28) In the case of the Dashehari mango cultivar, water-solu-
ble pectins showed a steep rise after 70 days, reaching a maximum at 101 days of fruit
growth.(29) The ammonium oxalate-soluble fraction showed a similar increase during fruit
growth. The alkali soluble fraction (protopectin) increased up to 70 days after the fruit set
but decreased thereafter until harvest. During ripening of the fruit, sucrose rose from 5.8 to
14.2% of the fresh weight, while the pH rose from 3.0 to 5.2. In the post-climacteric stage,
the content of non-reducing sugars fell to 0.6% and total acidity (as citric acid) varied
from 0.13 to 0.71%. Oxalic, citric, malic, succinic, pyruvic, adipic, galacturonic, glucu-
ronic, and mucic acids, together with two unidentified acids, were reported; citric acid was
the major organic acid present in mango fruit.(30)
Mango fruit contains 0.5–1% protein on a fresh weight basis.(16) A decrease in the sol-
uble protein content up to 44 days after fruit set, which increased again until 96 days was
reported.(27) A Peruvian variety showed an exceptionally high content of 1.57–5.42% of
protein.(30) The skin of Java grown fruit contains 1–2% and the pulp 0.6–1% of protein.
Twelve amino acids were reported during fruit growth.(29) At the peak stage, only alanine,
arginine, glycine-serine, and leucine-isoleucine were detected, while others were present
in traces. At maturity, their levels were predominant, which decreased during ripening,
with the exception of alanine.
The lipid content in peel and pulp of five mango varieties ranged from 0.75 to 1.7%
and 0.8 to 1.36%, respectively.(31) The total lipid in seven commercial cultivars ranged
between 0.26 and 0.67% at harvest.(32) A major component of the pulp was reported to be
a triglyceride, while mono- and di-glycerides were minor components.(33) The characteris-
tic odor that appeared in the fruits during ripening is due to components of ester and car-
bonyl types, which are varietal specific. The major volatile components of mango are
terpenes, although several other hydrocarbons, esters, and alcohols were also found to be
present in ripe mango fruit.(34,35) Vitamin C content was maximum (300 mg /100 g) in
Pairi variety in the early stages of growth although the ripe mango is an excellent source
of this vitamin.(36) A downward trend from 88 to 22 mg% within 5–10 weeks after fruit set
in Mulgoa, Pico, Amini, and Turpentine varieties of mango during growth was
observed.(37) Gosh(38) reported 36 mg of folic acid in 100 g of green fruit and Gopalan
et al.(39) found 0.08 mg of thiamine (Vitamin B1) and riboflavin (Vitamin B2) and 0.09
mg of niacin per 100 g of ripe mangoes.
At the initial stages, there was a steep fall in peel chlorophyll, which slowed down at
later stages of development, and the pulp chlorophyll became negligible as the fruit
approached maturity. Total carotenoids and β-carotene remained very low initially and
increased gradually as the fruits approached maturity and ripening. Sixteen different caro-
tenoids were identified, of which β-carotene was found to account for 60%.(40) The
increase in carotene was accompanied by a decrease in acid and an increase in sugar con-
tent. Some of the phenolic compounds identified in mango are gallic acid, indigallic acid,
gallotannin, quercetin, isoquercetin, mangiferin, and ellagic acid.(22,41) Ash content
decreased during development with some increase near maturity, while crude fiber
remained more or less constant.(19)
Physical and Chemical Indices of Mango Fruit Maturity

Mango fruits traded commercially for consumption as ripe fruits are harvested green and
ripened after harvest. If picked immature, however, fruits develop white patches or air
patches and show lower amounts of brix or total soluble solids (TSS) to acid ratio, taste
and flavor, whereas over-mature fruits lose their storage life. Such fruits pose a lot of
problems during handling. Mango fruits attained physiological maturity in about 90 days
and the increase in size and weight almost stopped 4 to 5 weeks before harvest in Dashe-
hari, Langra, Fazli, Zafrani, Alphonso, and Kishanbhog varieties.(21) In Alphonso, the
titrable acidity increased from the sixth to the tenth week after fruit set and steadily
declined thereafter as the fruit matured. Although endogenous ethylene and the induction
of the ripening process appear to be involved in the later stages of maturation and enhance
uniformity of the process, it may diminish, depending on the handling the fruit undergoes.
Some of it persists through ripening.
Harvesting of Mango
Mangoes are generally harvested at a physiologically mature stage and ripened for opti-
mum quality. Generally, in India, fruits are handpicked or plucked with a harvester, or
branches are vigorously shaken to drop them. Fruits not reached by hand are most often
retrieved with poles adapted with a severing blade and a bag. Recently, somewhat modified
mango harvesters have been developed, for use mainly in developed countries. Hydraulic
driven lifts are used in the United States for picking mango fruits. It was further observed
that the decay loss, particularly due to stem end rot, and the rates of respiration were min-
imum in fruits harvested with the stalk.(12) After harvesting, fruits should be heaped under
shade to avoid direct sunlight. Damaged, diseased, immature, and ripe fruits are sorted
out, and the fruits of similar maturity are packed together. The fruits should not be allowed
to fall on the ground as the damaged fruits cause spoilage to other healthy fruits during
packaging and storage.
Packaging of the Fruit

Fruits are graded according to their size, weight, color and maturity. It has been observed
that bigger size fruits take 2–4 more days for ripening than smaller fruits.(11) Hence, pack-
aging of smaller fruits with larger fruits should be avoided to achieve uniform ripening.
Proper packaging is an essential prerequisite. The formation of brown spots on the lenti-
cles has been problematic and is attributed to post-harvest handling procedures. In India,
baskets of bamboo, pigeon pea (Cajanus) or mulberry with paddy straw as cushioning
material have been used because of their easy availability and low cost. This type of pack-
aging was found to be unsatisfactory because of uneven ripening of fruits, excessive
shrinkage and bruising. Moreover, stacking was also a problem with the use of baskets.
However, more uniform ripening and better-quality mangoes were observed when fruits
were packed in ventilated wooden boxes.(42) Too much ventilation also affects the quality
of fruits due to shrinkage, loss in weight, and color changes. To overcome these problems,
corrugated fiber board (CFB) boxes are being used extensively. Alphonso mangos packed
in CFB boxes with partitions showed less bruising, slower ripening, reduced shriveling,
and less spoilage, as compared to fruits packed in wooden boxes. Various cushioning
materials such as newspaper, paddy straw, paper scraps, tissue paper, dry, and soft
grasses, neem leaves, or wood wool have been used as packaging of mangoes.(28,43)
Mango fruits are transported in various packings or loose in carts, trucks, and by rail. Ven-
tilated low-density polyethylene (LDPE) linings have also been found to be beneficial, as
this material maintains humidity, which results in less shrinkage during storage.(44) In a
recent study, it was reported that mango fruits stored in wax-lined cartons sealed with chi-
tosan films have a longer shelf life and retain a higher level of desirable quality attributes
than fruits stored in wax-lined cartons sealed with LDPE films or in perforated plastic
boxes.(45) Wrapping of individual fruit (unipack system) in tissue or newspaper was also
found to be suitable for uniform ripening during storage of Dashehari and Langra.(46,47)
Post-harvest Physiology of Mango
Preservation
Mango fruit is vulnerable to post-harvest losses due to its high perishable nature. Stor-
age and ripening of mango are beset with many problems. Many factors, such as culti-
var, stage of maturity, size grading, method of harvesting, handling, packaging, and
mode of transport affect the storability of mango fruits. Most losses are caused by pre-
ventable transportation and inappropriate storage methods. The post harvest life of man-
goes usually does not exceed 2–3 weeks and is limited by the physiological
deterioration of the fruit related to over-ripening and by pathogen infection and devel-
opment leading to decay. Various methods of post-harvest handling have been
employed to extend the shelf life of mango fruit and reduce losses, through inhibition of
respiration and ethylene production, which slows deterioration and senescence. These
can be classified as physical and chemical methods, which include refrigeration or cold
storage, polythene film packaging, wax coating, sub-atmospheric pressure storage, con-
trolled atmosphere and modified atmospheric storage, irradiation, heat treatment, and
the use of various chemicals. A combination of these can also be adopted to extend the
shelf life of the fruit.(48)
The current storage techniques are expensive, and also not fully satisfactory. Fur-
ther, a variety of disorders including development of off-flavor can result if fruits are
exposed to O2/CO2 concentrations below or above certain threshold levels.(49) There-
fore, storage and ripening of mangoes continue to be challenging problems that need
attention. In recent years, with molecular-biology studies, “Tomato biotechnology” took
a new turn of events, wherein fruit ripening was manipulated at the gene level—can
approach considered to be very promising. Control of ripening has been very successful,
as shown by the antisense RNA technology, where firmer tomatoes were grown with an
extended shelf life by individually suppressing the ACC synthase, ethylene-forming
enzyme (EFE), polygalacturonase (PG), and pectin methyl esterase (PME). One or more
genes were identified and used in the “sense” or “antisense” orientation to extend the
shelf life of commercially important fruits.(50) To control the post-harvest life of any
fruit with a molecular approach, a basic understanding of the events occurring during
fruit ripening is essential.
Processing
Mango fruit is processed and used at almost every stage of its growth. The range of products
includes those derived from both raw as well as ripe fruit. Mango fruits during early stages of
growth are commonly used for sweet or sour chutney (mango sauce). As the fruit attains the
stone hardening stage, they become suitable for some other useful products like amchoor,
pickles, and green mango beverages. Raw mango slices are also preserved for use as a basic
material for pickle and chutney manufacturing.(19) Ripe mango fruit has a characteristic com-
bination of taste and flavor. Due to the comparatively shorter storage life of mango fruits, it is
essential to make these products immediately. A large number of products are prepared from
ripe mango fruit, the methodology for which has been variously described for frozen and
canned slices, pulp, jam, squash, juice, nectar, ready-to-serve (RTS) beverages, mango-cereal
flakes, mango fruit bars or Aam papad, mango powder, and mango toffee.(51)
Diseases and Insect Pests

Mango suffers from several diseases at all stages of its life. All the parts of the plant,
trunk, branch, twig, leaf, petiole, flower, and fruit may be attacked by a number of patho-
gens, including fungi, bacteria, and algae; they may also suffer from infestation by various
insect pests. Those diseases affecting flowers and fruits are the most serious and may
sometimes result in the loss of a crop. Some of the infections of mango are anthracnose
caused by Colletotrichum gloeosporioides, alternaria rot (Alternaria alternata), and Pow-
dery mildew (Oidium mangiferae Berthet and Oidiopsis spp.).(52) The primary insects that
damage mango are the mango hopper (Idioscopus clypeatu), the mango mealy bug
(Drosicha mangiferae), the mango fruit fly (Daccus dorsalis, Strumata ferrugineus), the
mango seed borer (Nozorda albizonalis), and the mango bud mite (Aceria mangiferae);
weevils and ants also may cause extensive damage to crops.(53,54)
Disorders of Mango
Like any other tropical and sub-tropical fruit, mangoes are also susceptible to various dis-
orders during post-harvest handling and storage. Post-harvest disorders primarily consist
of three kinds: pathological, entomological, and physiological disorders. Among patho-
logical and entomological disorders, mango scab and sooty mould are the major diseases
that affect the mango fruit. The primary post-harvest pests of mango are fruit flies of the
genus Dacus, mango stone weevil (Sternochetus mangiferae), and the mango weevil
(S. gravis F). Control measures by chemical treatments can be applied either as pre-harvest
spray or post-harvest dip. Pre-harvest factors that predispose mango to physiological dis-
orders include growing location, orchard condition, tree nutrition, and conditions at har-
vest, whereas post-harvest storage conditions such as temperature, oxygen and CO2 levels,
packaging, and surface coating treatments are contributing factors to the occurrence of
other disorders. Conditions such as soft-nose, tip-pulp, internal breakdown, soft-center or
spongy-tissue have been described in the literature.(43) Some of the physiological disorders,
like chill injury, are caused by prolonged storage of fruits below a temperature of 10°C.
Physiology of Mango
Since mangoes are generally harvested at a mature-green stage, post-harvest changes are
principally concerned with events associated with ripening and senescence and with the
effects of post-harvest handling techniques. Accordingly, the post-harvest life of mangoes
can be divided into three phases.(55)
(a) Storage life (or Transportation life), which encompasses the period from harvest dur-
ing which the fruit remains unripe and in a condition resistant to physical damage dur-
ing normal handling.
(b) Ripening period, which designates the period from harvest until the fruit attains the
stage of maximum consumer acceptability (10–12 days). This period encompasses the
storage life period plus the final stage of ripening.
(c) Shelf life, which starts when the fruit is fully ripe and is the period in which fruit
remains in an edible condition.
In general, mangoes take 6–14 days to ripen under ambient conditions, depending on
the variety and environmental conditions and gradually become overripe. As a climacteric
fruit, the period of ontogeny is characterized by a series of biochemical changes initiated
by the autocatalytic production of ethylene and increase in respiration. Ripening results in
the characteristic color, taste, and aroma with desirable softening.
Much work dealing with biochemical changes during ripening has been done to study
the post-harvest physiology of mango fruit, specifically the organic acid metabolism,(56–57)
overall composition, and gross changes in cell wall and total pectin during ripening. (58–62)
However, considerable differences exist between the cultivars of same species.
Fruit Ripening
The ripening process is concerned mainly with alterations in biochemical components
already existing in the organ. Fruit ripening is a genetically programmed, highly coordi-
nated, and irreversible phenomenon involving a series of physiological, biochemical, and
organoleptic changes that lead to the development of a soft, edible, ripe fruit with desirable
qualities. A spectrum of biochemical changes such as increased respiration, chlorophyll
degradation, biosynthesis of carotenoids, anthocyanins, essential oils and flavor compo-

nents, increased activity of cell wall degrading enzymes, and a transitory increase in ethyl-
ene production are instrumental for changes involved during fruit ripening.
Ripening changes involve multiple biochemical pathways that affect all the cell com-
partments. The ripening of fruit is accompanied by a change in color, which is caused by
degradation of the chlorophyll leading to unmasking of previously present pigments(63)
and dismantling of the photosynthetic apparatus, synthesis of different types of anthocya-
nins and its accumulation in vacuoles,(64) and accumulation of carotenoids such as β-caro-
tene, xanthophyll esters, xanthophylls, and lycopene in the plastids.(65) The increase in
sweetness, as a result of gluconeogenesis, hydrolysis of polysaccharides, especially starch,
decreased acidity and accumulation of sugars and organic acids with an excellent sugar/
acid blend are responsible for the taste development. The increase in flavor and aroma
during fruit ripening is owing to the production of a complex mixture of volatile com-
pounds viz., ocimene and myrcene(65) and degradation of bitter principles, flavanoids, tan-
nins, and related compounds. The metabolic changes during fruit ripening include an
increase in biosynthesis and an evolution of ethylene, de novo synthesis of enzymes cata-
lyzing ripening specific changes, and an increase in respiration mediated by mitochondrial
enzymes.(63)
The ripening phenomenon is associated with loss of firmness, hydration of cell wall,
changes in cell wall thickness, decrease in the structural integrity, and increase in the
intracellular spaces. The major textural changes resulting in the softening of fruits are due
to enzyme-mediated alteration in the structure and composition of cell wall, partial or
complete solubilization of cell wall polysaccharides (pectins and celluloses), and hydroly-
sis of starch and other polysaccharides.(66) An increase in the soluble, but decrease in
insoluble, proteins was reported during ripening of mango fruits.(67) The changes in gene
expression during ripening involves the appearance of new “ripening-specific” mRNAs as
well as tRNA, rRNA, and poly (A+) RNA, and the disappearance of some mRNAs and
proteins.(64) However, some mRNAs are found to remain constant throughout ripening,
and these changes are known to be activated by plant hormones. An overview of biochem-
ical events during fruit ripening, depicted in Fig. 2, shows several control points at ethylene (1)
and post-ethylene (2) levels.
Role of Ethylene in Fruit Ripening

Ethylene, a simple olefin, is a plant hormone, specifically a fruit-ripening hormone, that
exists in the gaseous state under normal physiological conditions and regulates many aspects
of plant growth, development and cell metabolism including initiation of ripening, and
senescence, particularly in climacteric fruits. It is biologically active in trace amounts and its
effects are physiologically very important. Ethylene regulates its own biosynthesis in various
plant organs.(68) Fruit ripening is controlled by ethylene, which is autocatalytically synthe-
sized in small concentrations prior to the initiation of ripening, which in turn triggers the
entire array of changes during ripening. Thus, synthesized ethylene activates the crucial
enzymes for ripening and inactivates the inhibitors present in unripe fruits.(69) Ethylene, as
low as 0.01 μL L−1 and 0.05–0.25 μL L−1 triggered the ripening process in mango and
banana, respectively.(70) Alphonso mangoes generate ethylene in the range of 0.02–0.18 ppm
during ripening, and a three-fold increase in ethylene production was observed in ripening
fruit slices with added methionine.(71) Ethylene evolution in ripening tomato indicated that
production follows a sigmoid curve. A number of reviews have been published on the role of
ethylene in fruit ripening, particularly in mangoes, as well as its biogenesis.(72)
The pathway of ethylene biosynthesis was first established in apple fruit.(72) Since
then it has been shown to operate in other climacteric fruits such as avocado, banana, and
tomato. The two key enzymes in the pathway are those catalyzing the conversion of S-
adenosyl methionine (SAM) to 1-aminocyclopropane-1-carboxylic acid (ACC) and ACC
to ethylene, called ACC synthase and ACC oxidase (ethylene-forming enzyme, EFE),
respectively. At the onset of fruit ripening, expression of multiple ACC synthase genes are
activated, resulting in increased production of ACC. ACC is then oxidized to ethylene by
ACC oxidase. In most cases, it is the ACC synthase activity that determines the rate of
ethylene biosynthesis. Inhibition of ethylene biosynthesis by antisense RNA to ACC syn-
thase and ACC oxidase was very well established in tomato fruit.(73,74) The resulting trans-
genic fruit do not overripen as normal controls, though some color change occurs, and a
mere ethylene boost triggers back all the ripening related biochemical changes in similar
way as normal fruit. Deamination of ACC to α-ketobutyrate by over-expressing ACC
deaminase enzyme also suppressed ethylene formation and fruit ripening.(75) Recently, the
cDNA encoding for ACC oxidase enzyme has been isolated and characterized from
mango.(76) The mango ACC synthase and ACC oxidase genes are being used for trans-
genic work in mango, for the extension of shelf life.(76) Besides ethylene, abscisic acid is
known to enhance fruit ripening, while indole acetic acid (auxin), gibberlic acid and cyto-
kinin delay ripening by antagonizing the stimulatory effect of ethylene.(77) Thus, the cru-
cial role of ethylene in fruit ripening is clear. An overview of fruit ripening, with special
reference to textural softening, has been diagrammatically represented in Fig. 2.
Textural Softening During Fruit Ripening

The softening process is an integral part of ripening of almost all fruits. Fruit ripening is
associated with textural alterations and extensive softening of the tissue, which is dramatic
in climacteric fruits. Texture is a major quality attribute that determines the acceptance of
a fruit. Textural changes during ripening arise from a loss of turgour and/or enzymatic
degradation of structural as well as storage polysaccharides, especially starch. Depending
upon their nature, inherent composition and organization, different fruits soften at differ-
ent rates and to varying extents. Fruits like mango, papaya, avocado, sapota, and banana
undergo drastic and extensive textural softening from hard to soft ripe stage, while fruits
like apple and citrus fruit do not exhibit such drastic softening though they undergo tex-
tural modifications during ripening.
Fruit Texture
The texture of fruit can be attributed mainly to the structural integrity of the cell wall and
middle lamella, as well as to the turgor pressure generated within cells by osmosis and
accumulation of storage polysaccharides.(78) Changes in turgor pressure, and degradation of
starch and cell wall polysaccharides determine the degree of fruit softening. In citrus fruits,
softening is mainly associated with change in turgor pressure, a process associated with the
post-harvest dehydration and/or loss of dry matter. Starch is the major polysaccharide
present in some fruits like mango and banana, and its enzymatic hydrolysis results in pro-
nounced loosening of cell structure with development of sweetness, due to sugar accumula-
tion. However, textural changes during ripening of most of the fruits are largely due to
changes in the physicochemical properties of the cell wall, which is mainly due to the deg-
radation of cell wall polysaccharides, endogenously controlled and catalyzed by various
carbohydrate hydrolases.(79) Subtle structural changes of the constituent polysaccharides
may occur during fruit softening, without affecting much of the gross cell wall composi-
tion.(80,81) Cell wall polysaccharides may undergo (partial) hydrolysis or solubilization
resulting in changes in their molecular mass, solubility and degree of substitution of the
individual polysaccharides. Non-covalent changes in the cell wall are detected by the
localized alteration in pH or ionic concentration, whereas covalent modification of the cell
wall (polysaccharides) generally results from the enzymatic processes.
The major classes of cell wall polysaccharides that undergo modifications during rip-
ening are pectins, cellulose and hemicelluloses. In fruits, which are known for excessive
softening, the cell wall is thoroughly modified by deesterification and depolymerization,
accompanied by an extensive loss of neutral sugars and galacturonic acid, followed by sol-
ubilization of the depolymerized oligo- and polysaccharides.(82) Softening is normally
accompanied by an increase in the concentration of soluble pectic polysaccharide. It
appears that pectin polymers become less tightly bound in the cell wall during ripening,
and the cell wall loosening involves hydrolysis of galactose-containing polysaccharides.
Brinson et al.(58) reported net loss of arabinose, galactose, and galacturonic acid during
cell wall degradation of mango.
Textural softening is of commercial importance as it directly dictates fruit shelf life
and its keeping quality, which should be considered to avoid mechanical damage during
harvesting and transportation. In general, the textural properties of fruits play a significant
role in consumer acceptability. The increased interest in controlling the textural qualities
of fruit has stimulated further research on cell wall biochemistry, with particular reference
to cell wall polysaccharides and their degradation.(77,83,84) The textural qualities of the fruit
are attributed to its inherent composition, particularly the cell wall composition. Attempts
to understand the molecular mechanism of fruit softening have directly led to the investi-
gation of cell wall polymers, their compositional changes and the related cell-wall degrad-
ing enzymes during ripening.(85)
Structural Polysaccharides and Textural Changes

Plant polysaccharides, in general, are an extremely diverse set of biopolymers, which play
a very important role as structural elements. These include celluloses, hemicelluloses, pec-
tins, reserve polysaccharides such as, starch and galactomannan, gel formers, such as
gums and mucilages, and physiological information carriers such as antigens.(86–88) Fruit
polysaccharides, upon their degradation in situ, play a crucial role in textural softening
during ripening.
Plant polysaccharides play a major role in storage, mobilization of energy, and in
maintaining cell and tissue integrity due to their structural and water binding capacity.
Polysaccharides from different sources vary in their chemical-biological, physico-chemi-
cal, and structure-functional characteristics. Cell wall polysaccharides differ widely in
their physical and nutritional properties; they regulate utilization of other dietary compo-
nents in the food. Recently, plant polysaccharides have emerged as important, bioactive
natural products exhibiting a number of biological properties, such as regulating gene
expression and host-defense mechanism, which includes release of enzymes and proteins
involved in the generation of elicitor-active oligonucleotide fragments from the cell wall.
Carbohydrate Hydrolysis During Ripening

Fruit softening is determined by the molecular changes occurring in cell walls during rip-
ening, which can be analyzed by either polymers released during ripening or changes in
total wall material. The observed changes can then be correlated with the enzyme activi-
ties present in the tissue and with the action of these enzymes on isolated cell walls.
Polysaccharides are polymers in which many monosaccharides of various types are
covalently linked by glycosidic bonds. The glycosidic bond is formed between hemiacetal
or hemiketal of one monosaccharide (glycosyl donor) and a hydroxyl group of the suc-
ceeding monosaccharide (glycosyl acceptor or aglycone) with the elimination of water.
Polysaccharides are classified as homo- or heteropolymers, based on the type of constitu-
ent monosaccharide units present. Homopolymers are made up of only one type of
monosaccharide unit (e.g., starch, cellulose), whereas heteropolymers have more than one
type of monomeric unit (e.g., pectins, hemicelluloses, gums, etc.). In addition, another
group called “glycoconjugates” is also included, usually under the broad definition of car-
bohydrates. Glycoconjugates include glycolipids, glycoproteins, and proteoglycans.
Pectins and Pectic Substances

Pectins are the prominent structural constituents of primary cell wall and middle lamella.
Along with cellulose microfibrils, they contribute to fruit texture, while they may be virtu-
ally absent in secondary walls. Pectin content varies from fruit to fruit and fruit pectins are
used for commercial purposes.(89)
Tissue softening is attributed to enzymatic degradation and solubilization of pro-
topectin, the insoluble, high molecular weight parent pectin, into soluble polyu-
ronides.(79,90) Protopectin increases before physiological maturity but decreases during
mango fruit ripening.(31) Native pectin plays an important role in the consistency of fruit
and also in textural changes during ripening, storage, cooking or irradiation, and other pro-
cessing operations. Pectins are likely to be the key substances involved in the mechanical
strength of the primary cell wall and are important to the physical structure of the plant.(91)
The edible portion of most plant foods, i.e., fruit pulp or the mesocarp, is composed of
parenchymatous tissue consisting of cells that are ∼50–500 μm across and polyhedral or
spherical in shape. Parenchymatous tissues are thought to consist principally of calcium
salts of pectic substances, which are deposited in early stages of the cell growth, specifi-
cally when the area of cell wall is increasing.(82) Middle lamella are heat labile, and their
dissolution results in separation of plant cells. Ultrastructural studies in ripening fruits
have also shown that cell wall breakdown was accompanied by dissolution of middle
lamella and gradual dissolution of fibrillar network of primary cell wall.(78) Deesterified
pectins in the middle lamella are associated with calcium ions, and its removal usually
leads to cell separation. The association involves binding of two or more polymeric
chains, in the form of corrugated egg-box, with interstices in which calcium ions are
packed and coordinated.(92)
Pectins are structurally diverse heteropolysaccharides containing partially methylated
galacturonic acid residues, methyl esterified pectins, deesterified pectic acids and their
salts, pectates and the neutral polysaccharides, which lack galacturonan backbone, i.e.,
arabinans, galactans and arabinogalactans.(93,94) The primary pectin chain, i.e., α-D-galac-
turonans, consists largely of D-galacturonic acid linked by α (1→4) linkages, wherein the
carboxyl groups are partially esterified with methanol and the hydroxyl groups are par-
tially acetylated with acetic acid. The degree of polymerization, degree of esterification,
and the proportion of neutral sugar side chains are the principal factors contributing to
heterogeneity.
Arabinogalactans (AG) are heteropolymers of D-galactose and L-arabinose residues. Two
structurally different forms of AGs are found in plants.(95) AG-I is a simple polysaccharide
composed of chains of β-(1→4) linked D-galactose residues with single L-arabinose resi-
dues linked to O-3 of the galactose residues. They have been isolated from different fruits
including apple, kiwi, tomato, and pineapple.(96–99) AG-II are complex and branched
polysaccharides, consisting of chains of β-(1→3) linked D-galactose residues linked to
chains of β-(1→6) linked D-galactose residues at the O-6 position of the main chain. The
O-3 and O-6 positions of the side chains are in turn linked to terminal L-arabinose resi-
dues.(95,96) Plant arabinogalactans are known for their multifaceted physiological and
functional characteristics,(100) such as freeze-inhibition, water holding, and adhesive prop-
erties. Due to their specific carbohydrate binding properties, they may possibly affect cell-
cell interaction.
The primary cell wall pectic polysaccharides have a relatively higher proportion of
neutral oligosaccharide chains on their backbone, and these side chains are much longer
than those of the pectins of middle lamella.(101) The side chains are concentrated in “hairy
regions.” Highly esterified and slightly branched rhamnogalacturonan, the “smooth
regions,” are present in middle lamella, whereas highly branched rhamnogalacturonan, the
“hairy regions,” are present in primary cell wall. In plant cell wall, the side chains of the
pectin molecules link to protein, hemicellulose, and cellulose.
The nonsugar substituents of acidic and neutral pectins are essentially methanol, ace-
tic acid, phenolic acids and amide groups, and contribute to their structural diversity. Che-
lator-soluble pectins have a high degree of methylation, as well as a high degree of
acetylation than those extracted with alkali. Phenolic acids, especially ferulic acid and p-
coumaric acid are found esterified to the non-reducing ends of the neutral arabinose/galac-
tose residues. Ferulic acid facilitates oxidative cross-linking between pectins or with other
polysaccharides in the cell walls, by the formation of diferuloyl bridges, which would
limit wall extensibility(102) and plays a significant role in growth regulation and defense
mechanism.(103)
Changes in Pectic Polysaccharides During Ripening

During ripening, fruits lose firmness, and unless the fruit is dehydrated, osmotic properties
of the cell and the turgor pressure usually remain constant. In plant tissues, it is assumed
that turgor pressure alone is not contributing to the loss of firmness, instead it is the result
of changes in the cell wall polysaccharides.(83) Much work concerning chemical changes
in cell walls to fruit softening has been focused towards the characterization of changes in
pectic substances.(104) Being soluble in water, pectins can be deesterified and depolymer-
ized mostly by enzymatic reactions. The retardation of textural softening by the addition
of Ca++ ions is related to the ability of divalent cations to form calcium bridges between
the pectic polysaccharide chains. In some, limited degradation of the pectic polymers
might be due to the methylation of galacturonic acid groups. Loss of firmness during heat
treatment of acid fruit has been attributed to hydrolysis of glycosidic bonds in cell wall
polysaccharides.(105) Arabinofuranosyl linkages are most labile, whereas the glycosidic
linkages are most stable.
Changes in the proportion and characteristics of pectic substances are reported in
many fruits. During ripening, the progressive loss of firmness is the result of a gradual sol-
ubilization of protopectin in the cell walls to form pectin and other products.(106) Solubili-
zation followed by depolymerization and deesterification of pectic polysaccharides is the
most apparent change occurring during ripening of many fruits.(107) Pectins from ripe fruit
exhibited a lower degree of esterification, a lower average molecular weight, and
decreased neutral sugar content compared to pectins from unripe fruits.
The loss of neutral sugar side chains from the pectin is another important feature
occurring during ripening. Out of 17 types of economically important fruits, 14 types
showed a net loss of galactose and arabinose from the cell wall during ripening, but no
such loss was observed in ripening plum and cucumber fruits.(108) A net loss of neutral
sugars during ripening of pear, apple and tomato was reported.(109–111) The mutant tomato
fruit “rin” containing little or no PG activity also showed substantial loss of galactose sug-
gesting that this loss is not due to the action of PG.(111) This evidence suggests that other
cell wall hydrolases, especially glycosidases, play an important role in textural softening
during ripening.
Cellulose and Hemicelluloses

The plant cell wall is made up of cellulose fibrils imbedded in a matrix consisting largely
of pectic substances, hemicelluloses, proteins, lignins, lower molecular weight solutes,
and water. Cellulose occurs naturally as an insoluble material, which is not easily
degraded by enzymes. Cellulose is a linear polymer of (1→4) β-D-glucosyl residues that
form the skeletal scaffolding of the cell wall through the formation of microfibrils, ∼5–15
nm in diameter consisting of several thousand units.(78) The glucan chains are assembled
into a paracrystalline array and are arranged in parallel to each other to form a
microfibril.(95) An apparent dissolution of the middle lamella and cell wall fibrillar net-
work due to cellulolytic activity in ripening of avocado, pear and apple was demon-
strated.(112,113) Ripening associated changes involving dramatic decrease in the content
and molecular size of hemicellulose are reported in several fruits The amount of hemicel-
lulose decreased steadily during ripening of many fruits including mango.
Hemicelluloses are a distinct group of structural (neutral sugar) polysaccharides.
They are the only cell wall polysaccharides that have the capacity to strongly bind cellu-
lose microfibrils through hydrogen bonding. Xyloglucans, mannans, xylans, galactan, glu-
cans, arabinoxylans, and glucuronoarabinoxylans are the major hemicelluloses found in
plant cell wall. The common structural features of hemicelluloses are a main chain with a
structural resemblance to cellulose and either short side chains that form a “pipe-cleaner-
brush” shaped molecule, or a different sugar interpolated in the main chain. The extraction
of hemicellulose is achieved by strong choatropic agents like 6 M urea, 6 M guanidium,
4.5 M thiocyanate guanidine or 1–6 M sodium hydroxide or potassium hydroxide. Nor-
mally, hemicelluloses are extracted by treating holocelluloses with aqueous alkali, which
may saponify any hemicellulosic ester linkages either between polysaccharides or
between hemicelluloses or non-carbohydrate components.(114) The extracted hemicellu-
losic materials are precipitated on neutralization or mild acidification and further by the
subsequent addition of an excess of acetone or ethanol. Alkaline extracted hemicelluloses
have been characterized from apple and sugar beet pulp.(115)
Xyloglucans are the major hemicelluloses found in dicotyledonous primary cell wall
(20%). The basic structure consists of a backbone of β-(1→4) linked D-glucosyl residues
with α-D-xylosyl chains linked to 0-6 of glucosyl residues. Xylans are the major hemicel-
lulosic components of monocots. They consist of a backbone of β-(1→4) linked D-xylosyl
residues. Xylans are strongly associated with cellulose through hydrogen bonding. Ripen-
ing associated changes involving dramatic decrease in the size of hemicelluloses are
reported in tomato, pepper, strawberry, and muskmelons. Degradation of arabinans by
fungal arabinases and arabinofuranosidase has been reported.(116)
Other important hemicellulosic components include mannans, which serve as food
reserve polysaccharides and confer plasticity and the ability to stretch. In plant systems,
mannans exist mostly in the β-configuration, either as hetero- or homopolymers. The β-D-
mannan polymers (linear homoglycan of β-(1→4)-linked-D-mannopyranosyl unit) are
documented in higher plant systems, whereas α-D-manno-oligomers (oligosaccharides of
α(1→4)-linked-D-mannose) exist as homo- or heterooligomers in oligosaccharides, gly-
coproteins, and glycolipids. The main group of mannose-containing polysaccharides from
cell walls of higher plants are glucomannans, galactomannans, or galactoglucomannans.
In fruits, it was noted that mannan type of substrates may undergo endogenous
hydrolysis during ripening, thus contributing to textural softening either directly or indi-
rectly. Mannose is an important component of the cell walls in fruits such as apples and
tomatoes.(117) A polysaccharide fraction from depectinated apple cell walls that contained
a very high proportion 1,4-linked mannosyl residues has been reported.(117)
Cell Wall Degrading Enzymes in the Context of Fruit Ripening

Plant cell wall polysaccharides are degraded by the plant’s own enzymes during physio-
logical developments such as fruit ripening, seed germination, and cell wall extension. In
avocado, tomato, apple and pear, the middle lamella becomes dissoluted and the fibrillar
structures disappear during softening.(118) These changes are considered to result from the
action of cell wall degrading enzymes, essentially the hydrolases. Most of these enzymes,
present in low levels, are constitutive throughout fruit development and ripening. But dur-
ing ripening, generally all the hydrolases increase in activity, particularly cell wall hydro-
lases, showing maximum activity at climacteric stage.
A wide range of cell wall hydrolases are identified in fruit tissues. The major carbo-
hydrate hydrolases involved in polysaccharide dissolution in vivo can be broadly classi-
fied into two types; viz, glycanases and glycosidases. Glycanases (glycanohydrolases) by
definition are a class of enzymes cleaving high molecular weight polymers (polysaccha-
rides) into shorter chains, whereas glycosidases (glycohydrolases) generally act on shorter
chain oligosaccharides, which may be homo- or hetero-oligomers, glycoproteins or gly-
colipids. The carbohydrate hydrolases may also be involved in signal transduction path-
way by way of deglycosylation.
Hydrolases
Among cell wall hydrolases, pectin-degrading enzymes are mostly implicated in fruit soft-
ening. Pectic enzymes are classified based on their mode of action on pectin and pectic
substances into PG, PME, pectate lyase, and pectin lyase.(119) The other enzymes, such as
arabinanase, galactanase, and β-galactosidase, act on the side chains of the galacturonide
backbone, degrading the entire pectic substance. Increased solubilization of the pectic
substances, progressive loss of tissue firmness and rapid rise in the PG activity accompany
normal ripening in many fruits including tomato and banana.(120,121)
Little is known about the enhancement of cellulase or hemicellulase activity in con-
nection with fruit softening. Cellulase is a multienzyme system composed of several
enzymes; endoglucanase (EC 3.2.1.4), exoglucanase (EC 3.2.1.91), and glucosidase (EC
3.2.1.21).(122) Endoglucanase hydrolyses the β-1,4-linked glucose residues at random
positions. Exoglucanase breaks the bonds at non-reducing end of the chain, producing glu-
cose or cellobiose (dimers of β-1,4-linked glucose), whereas β-glucosidase splits cellobi-
ose into glucose molecules. Cellulose levels in unripe fruit are generally low and increase
dramatically during ripening.(123) The loss of firmness, climacteric rise of respiration, and
ethylene evolution in ripening fruit was directly correlated with marked increase in cellulase
activity.(124) Cellulase has been implicated in softening process in tomato.(125) Cellulase

activity was reported in several Indian mango cultivars, which increased during ripening
especially in Alphonso.(126) The progressive textural loss in “keitt” mango was attributed
to marked increase in cellulase activity.(61)
Mannanase is a glycanase that catalyses the hydrolysis of β-1,4-mannans, a polymer
of mannose. The role of an endo-β-1,4-mannanase in fruit softening is suggested by its
ripening-specific accumulation. Pressy has characterized endo-β-mannanase in ripening
tomato.(127) McCleary and Matheson reported that the degradation of galactomannan
involves endo-β-mannanase.(128) It has been proposed that β-endomannanase is the rate-
limiting enzyme controlling germination timing.
Xylanases (EC 3.2.1.8) catalyze the hydrolysis of β-1,4-xylan. β-1,4-D-endoxylanase
and β-1,4-D-exoxylanase are reported as cell wall degrading enzymes in fruits such as avo-
cado(129) and banana.(130) Xylanase, arabinanase and mannanase are localized both in soluble
and bound form, which showed an increase during ripening. It was interesting to note that
arabinanase, galactanase and mannanase, and to some extent laminarinase were very promi-
nent enzymes in mango fruit giving activity peaks at climacteric stage of ripening.(131)
α-Amylase (EC 3.2.1.1) and β-amylase (EC 3.2.1.2) are the two amylases in plant tis-
sues capable of metabolizing starch. α-Amylases hydrolyse the α-1,4-linkages of amylose
at random to produce a mixture of glucose and maltose. β-Amylase attacks only the penul-
timate linkages and thus, releases only maltose. Amylase activity increases to some extent
during ripening.(63) These enzymes are unable to degrade the α-(1→6) branch points of
amylopectin, which are catalysed by amyloglucosidases. Mango and banana are the major
starch-containing fruits (∼15 to 20%), where the starch was almost completely hydrolysed
during ripening to free sugars, thus contributing to loosening of cell structure and textural
softening.
Glycosidases
Among glycosidases, the prominent enzymes found in ripening fruit were α-mannosidase,
β-hexosaminidase and α- and β-galactosidases.(132,133) α-Mannosidase (EC 3.2.1.24), docu-
mented in very few fruit systems, acts on short chain oligomers of ∼8–10 mannose units
present either as glycoprotein, glycolipid, or hetero-/homopolysaccharides. This enzyme
partially degrades the pectic and hemicellulosic components of the cell wall and is possibly
related to the breakdown of polysaccharides. Increased activity of α-mannosidase during
ripening has been reported. Interestingly, it not only showed an activity peak during ripen-
ing/softening, but it was also the most active enzyme amongst the glycosidases examined.
The physiological role of most of the glycosidases during ripening is not known.
One novel approach to elucidate the role of enzymes in cell wall degradation and soft-
ening is to employ antisense RNA technology. This technology was one of the first molec-
ular approaches used for delaying fruit ripening.(50) It has been possible to obtain firmer
tomatoes with longer shelf life by specific suppression of PG gene expression with anti-
sense RNA.(134) Pectin methyl esterase (PME) suppression resulted in increased solid con-
tent in tomato.(135) The genes coding for PG, PME and other enzymes have been cloned in
tomato and other fruits.(136)
Biotechnology
As the traditional methods of breeding have almost reached a stagnation point in terms of
productivity, there is a need to supplement with biotechnological methods. Propagation of
plants through tissue culture, including sophisticated techniques of meristem culture and
disease indexing, is of immense use in making available healthy propagation materials.
Besides this, tissue culture techniques have direct application in large-scale production of
plants in relatively smaller space, shorter times, as well as rapid multiplication of valuable
genotypes and improved plants. It also has added advantages such as the nonseasonal,
almost year-round production of plantlets and conservation of germplasm for convenient
international exchange.
Plant transformation and gene cloning are becoming important tools in plant improve-
ment via genetic engineering. However, development of an efficient and reproducible tis-
sue culture regeneration protocol is the first step in utilizing the power and potential of this
new technology. The plant organism is composed of many single cells, which organized
as tissues, form the different organs of the differentiated plant. Isolated cells, tissues or
organs of the plant can be cultivated on synthetic media under sterile conditions.
The essential cell material in most plant cell culture systems is known as callus,
which is a mass of undifferentiated cells. The explant derived from the whole plant is
surface-sterilized and transferred aseptically onto a complex semi-solid medium supple-
mented with plant growth regulators (usually an auxin or cytokinin). Subsequently, cell
proliferation occurs and callus forms. Callus can be removed from the explant and main-
tained in vitro by routine subculture or by various storage techniques. Alternatively, the
callus can be manipulated to cause cell differentiation into organized tissue and hence,
whole plants can be regenerated.
One of the unsolved problems in biology is the process by which single cell (e.g.,
germ cells), or small populations of seemingly identical cells, undergo the coordinated
divisions and development that result in the formation of a complex, highly structured
mature organism in which a great many different cell types may be present. This is the
process of differentiation, and clearly involves the differential expression of genetic infor-
mation. The main type of differentiation seen in plant cell and tissue cultures are root for-
mation and shoot production, collectively termed organogenesis. In a few cases, these
processes occur simultaneously in an apparently coordinated manner, and this phenome-
non is called somatic or adventive embryogenesis. Numerous plant species have been
reported to be capable of forming somatic embryo from diverse explants.(137,138) Somatic
embryogenesis offers distinct advantages over regeneration via organogenesis.
Much of the research into biotechnology has been carried out in industrialized coun-
tries, resulting in the comparative neglect of regional crops in developing countries, where
many of the tropical and subtropical fruit species are grown. In addition, since the task of
establishing collections of world’s plant genetic material began, most activity has
occurred quite naturally, with the major food crops. Very little effort has been applied to
the minor crops and their wild relatives especially tropical and subtropical fruit species.
Many of these fruit crops are vital to local communities in terms of variety of available
food, nutrition (particularly vitamins), and therefore quality of life. In addition, there
exists potential to bring some of these crops to an expanded international market by apply-
ing new biotechnologies, particularly those that can enhance flavor and post-harvest char-
acteristics. In recent years, some of the minor tropical fruits have already become more
popular resulting in increased exports to countries such as USA, EC, Singapore, and Hong
Kong. Tropical and subtropical fruit species are currently somewhat neglected as far as
both germplasm collection and application of biotechnology are concerned. From a posi-
tive perspective, they represent a wealth of as yet untapped potential for valuable R&D
efforts. There is much that can be done using biotechnology techniques, not only to con-
serve germplasm of these species, but also to develop many of them as major fruit crops.
Protoplast Culture
Exploitation of the variation that exists in populations has led to the development of many
commercial varieties and hybrids. Although protoplasts can be isolated from a range of tis-
sues of almost any plant species, regeneration of plants from protoplasts is one of the most
difficult in vitro techniques. Reports of success with recalcitrant woody species are limited
and applications to tropical and subtropical fruit species are rare. The only exception is with
citrus species. There have been reports of protoplast isolation of non-woody tropical fruit spe-
cies, such as mango (139) and banana,(140) although sustained cell division was not achieved.
The principal uses of protoplasts are as targets for direct methods of gene transfer and
to facilitate interspecific hybridization. Direct methods of gene transfer requiring proto-
plasts are becoming obsolete. Although, protoplast fusion may provide a method of inter-
specific hybridization, the difficulties in achieving sustained cell division and plantlet
regeneration in many species generally outweigh the advantages. In reality, protoplast
fusion is only an advantage when prozygotic barriers prevent interspecific hybridization.

Available literature shows that application of biotechnology to tropical and subtropical
fruit species is limited. There has been some research effort directed to this field in the 1990s
however, before this period little information was available, with the exception of a few spe-
cies (citrus, banana, papaya, pineapple). Relative to the effort that has been applied to research
of temperate species, biotechnology of tropical and subtropical species is a vastly unexplored
frontier. A notable exception is citrus species, which have been the subject of considerable in
vitro research, and are a model for application to other tropical and subtropical species.
Micropropagation
Regeneration via callus is prone to somaclonal variation, whereas the meristematic tissue
in buds is inherently the most genetically stable tissue in plants. Genetic stability in a
micropropagation system is very important when applied to elite genotypes, particularly if
the aim is germplasm conservation.
Vegetative propagation of trees is an effective way to capture gain and produce large
amounts of plant material. Micropropagation and somatic embryogenesis are currently
being used with commercial tree species. Conventional breeding approaches with woody,
perennial fruit crops have been complicated by generational cycles as long as 6 to 7 years,
and by the absence of useful genetic markers. The advantages of efficient generation of
tropical, perennial fruit trees from cell and tissue cultures in crop improvement programs
have been reviewed.(141) The propagation in vitro of superior, disease-indexed selections
that are otherwise hard to propagate clonally would have an immediate effect on the pro-
duction of many tree crops. Mutant selection, the recovery of horticulturally useful soma-
clonal variants from cell and protoplast cultures, and the use of recombinant DNA may
alter the breeding strategies for many tropical fruit crops. Unfortunately, the application of
cell culture techniques to the improvement of woody, crop plants has been limited due to
the absence of methods of regeneration from tissues of mature origin. Mango, an impor-
tant crop, is a woody perennial. Regeneration/propagation of mango in vitro has been dif-
ficult due to inherent problems of contamination and recalcitrance.
Agrobacterium Tumefaciens Mediated Mango Transformation

Genetic transformation, developed for many plant species, is usually based on the delivery
of defined foreign genes into plant cells, obtaining integration of the genes into plant
genomes and observing expression of the genes in the regenerated plants. Numerous DNA
delivery systems have been reported. The preferred method of gene for dicots is Agrobac-
terium-mediated transfer. Cocultivation of callus, suspension cultures or leaf discs with
Agrobacterium has been used to successfully transform many species; for example, effi-
cient systems have been described for major crop species such as potato,(142) tomato,(143)
and sugar beet.(144) In the 1980s, Agrobacterium tumefaciens-mediated transformation
was limited to dicotyledonous plants and thought to be not readily applicable as a method
to produce transgenic plants using Agrobacterium. Compared to herbaceous species, there
have been only a few reports of in vitro transformation involving angiosperm tree species.
Transformed cell lines have been reported in peach(145) and papaya.(146) Embryogenic cul-
tures of papaya have also been transformed using microprojectile bombardment.(147)
Transformed somatic embryos and plantlets have also been obtained from embryogenic
cultures of walnut.(148,149) The successful application of plant transformation techniques is
dependent on plant tissue culture protocols to regenerate transformed plants. Transgenic
plants of a number of fruit species have been produced.

Mango tissue seems to be relatively easily amenable for regeneration in vitro. Litz has
reported considerable success with somatic embryogenesis from cultures of mango.(139)
For example, Agrobacterium transformation was successful after cocultivation of an
embryogenic mango culture.(150) Although it has taken a very long generation time, the
prospects for gene transfer in mango seem quite good.
The ripening of most fruits involves cell wall degradation. A large number of genes
involved in membrane trafficking have been identified and characterized in animal and
yeast systems, which appear to be involved in controlling membrane fusion. A full-length
c-DNA clone from mango fruit has homology to the rab1/YPT 3 class of small GTPases.
The corresponding mRNA is expressed in fruit, only during ripening. The likely involve-
ment of this rab X protein in trafficking cell wall modifying enzymes through the trans-
golgi network has been reported.(151) DNA finger print information was used for the iden-
tification of mango cultivars for genetic relatedness analysis of a family structure. Classi-
cal approaches for the identification of mango cultivars are based on morphological traits.
The evaluation of these traits is subjective and their assessment is difficult. Recently,
isozyme analysis has been used to identify mango cultivars and to decipher their
percentage.(152,153) However, isozyme polymorphism is limited, hence limiting its ability
to identify cultivars.
Conclusions
Mango is a tropical fruit tree of great economic importance. Mango fruit is susceptible to
many pathogens, and due to this its production is threatened in both traditional and new
growing areas. Difficulties in establishing aseptic mango cultures from mature explants
associated with phenolic browning, greatly hinders the micropropogation of mango.
Somatic hybridization via protoplast fusion could be an alternative to overcome these
problems. Several positive characteristics have been incorporated into many plants by pro-
toplast fusion. An efficient system for plant regeneration is required for somatic hybridiza-
tion. While protoplast regeneration protocols have been described for a few tree species,
protoplast research involving tropical fruit trees other than citrus is very limited. The
regeneration of plants from protoplasts isolated from pro-embryogenic masses in a sus-
pension culture derived from the nucellar callus of the mango cv. “Amrapali” has been
reported.(154) In this direction, further biotechnological approaches may be worth pursuing
for sustained mango cultivation.
Acknowledgment
HMY thanks CSIR, New Delhi, for a Senior Research Fellowship.
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This article was downloaded by: [Iowa State University]

On: 08 October 2013, At: 14:11

Food Reviews International

Mango (Mangifera indica L.), “The King

To link to this article: http://dx.doi.org/10.1080/87559120600574493

PLEASE SCROLL DOWN FOR ARTICLE

Mango (Mangifera indica L.), “The King of

R.N. THARANATHAN, H.M. YASHODA, AND T.N. PRABHA

Department of Biochemistry and Nutrition, Central Food Technological

Keywords Mangifera indica, mango, fruit ripening, textural softening, enzymes,

Address correspondence to R.N. Tharanathan, Department of Biochemistry and Nutrition,

are an important source of essential micronutrients, vitamins, and other phytochemicals,

Mango—“The King of Fruits”

Indonesia 444 640 605

to high amounts of vitamin C, β-carotene, and minerals.(8) In 2001–2002, India exported

Figure 1. Mango tree bearing fruits.

Growth and Development of Mango Fruit

Compositional Changes During Development

Physical and Chemical Indices of Mango Fruit Maturity

Packaging of the Fruit

Post-harvest Physiology of Mango

Diseases and Insect Pests

degradation, biosynthesis of carotenoids, anthocyanins, essential oils and flavor compo-

Role of Ethylene in Fruit Ripening

Textural Softening During Fruit Ripening

Structural Polysaccharides and Textural Changes

Carbohydrate Hydrolysis During Ripening

Pectins and Pectic Substances

Changes in Pectic Polysaccharides During Ripening

Cellulose and Hemicelluloses

Cell Wall Degrading Enzymes in the Context of Fruit Ripening

activity.(124) Cellulase has been implicated in softening process in tomato.(125) Cellulase

fusion is only an advantage when prozygotic barriers prevent interspecific hybridization.

Agrobacterium Tumefaciens Mediated Mango Transformation

plants of a number of fruit species have been produced.

HMY thanks CSIR, New Delhi, for a Senior Research Fellowship.

the United Nations. Rome, Italy, 1999.

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