The Mango (Cabi)

670 Pages·2009·4.09 MB·English

by  Richard E Litz

1
Introduction: Botany and Importance
S.K. Mukherjee1 and R.E. Litz2
1Calcutta University, Calcutta, India
2University of Florida, Florida, USA
1.1 Introduction 1
1.2 Description of Mango 2
The tree 2
Flowers 2
The fruit 3
The seeds and polyembryony 4
1.3 History of Cultivation 5
Origin of Mangifera indica 5
Domestication of mango 9
Distribution 10
1.4 Germplasm Conservation 11
Genetic erosion 11
Collection and documentation of Mangifera germplasm 12
Relevance of germplasm resources to mango improvement 12
1.5 Importance of Mango 12
Cultivars 12
1.6 Production and Uses 14
1.1 Introduction
Mango has become a major fruit crop of the tropics and subtropics, particu-
larly in Asia, where the mango has always been the most important fruit crop
and where it has been considered the ‘king of fruits’ (Purseglove, 1972). A
generation ago, the Green Revolution culminated, creating surpluses of sta-
ple and horticultural crops in many developing countries. The Green Revo-
lution was the result of nearly a century of effort of applying Mendelian
genetics to crop improvement (i.e. conventional breeding) together with the
optimization of agronomic and horticultural practices and the successful
management of insect pests and diseases. However, improvement of tree
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 1
2 S.K. Mukherjee and R.E. Litz
crops has lagged far behind fi eld crops for several reasons: their heterogene-
ity, polyploidy, lengthy juvenile period, time required for evaluation of trees
in the fi eld, and the relatively high cost of maintaining tree plantings. For the
most part, fruit cultivars continue to be ancient selections, many of which
have serious problems, including alternate bearing, lack of disease resistance,
low yields, etc. The rapid growth of mango production in recent years has
been due to its expansion into new growing regions of the New World, China
and parts of Africa; the planting of regular bearing selections; and the adop-
tion of modern fi eld practices, which include irrigation management, control
of fl owering, etc. Agricultural practices are currently undergoing another
revolution, as integrated pest and disease management replaces the earlier
reliance on agrichemicals, and emerging fi elds within biotechnology begin to
impact cultivar development.
1.2 Description of Mango
The tree
The mango tree is believed to have evolved as a canopy layer or emergent
species of the tropical rainforest of South and South-east Asia (Kaur et al.,
1980; Bompard, Chapter 2, this volume). Mature trees can attain a height of
40 m or more, and can survive for several hundred years. Mango trees that
have been domesticated by selection from openly pollinated seedling popu-
lations show variation in tree architecture (i.e. shape and size). The tree is an
arborescent evergreen. Leaves are simple and alternate, with petioles that
range in length from 1 to 12.5 cm. Leaf morphology is highly variable, de-
pending on the cultivar: leaves can be lanceolate, oblong, ovate and interme-
diate types involving these forms. Leaf length ranges from 12 to 38 cm and
width can be between 2–13 cm. Young leaves are copper-coloured, changing
gradually to light and then dark green with age. The leaves are spirally
arranged in whorls and are produced in fl ushes. The canopy is normally
oval, elongated or dome shaped. The juvenile period of seedling trees can
range from 3 to 7 years. The root system consists of a long, vigorous taproot
and abundant surface feeder roots.
Flowers
Mango fl owers are borne on terminal pyramidal panicles, and are glabrous
or pubescent; the infl orescence is rigid and erect, up to 30 cm long, and is
widely branched, usually tertiary, although the fi nal branch is always cymose.
The infl orescence is usually densely fl owered with hundreds of small fl ow-
ers, which are 5–10 mm in diameter. The fl owers are either monoecious or
polygamous, and both monoecious and polygamous fl owers are borne
within a single infl orescence (Plate 1). The pistil aborts in male fl owers. The
ratio of monoecious to polygamous fl owers is strongly infl uenced by
Introduction: Botany and Importance 3
environmental and cultural factors. The fl owers have four or fi ve sepals and
petals that are ovate to ovoid to lanceolate and also thinly pubescent. The
fl oral disc also is four- or fi ve-lobed, fl eshy and large and located above the
base of the petals. There are fi ve large, fl eshy stamens, only one or two of
them being fertile; the remaining stamens are sterile staminodes that are sur-
mounted by a small gland. In addition, two or three smaller fi laments arise
from the lobes of the nectaries. The stamens are central. The ovule is anatro-
pous and pendulous. It is believed that the fl owers are cross-pollinated by
fl ies (see Davenport, Chapter 5, this volume).
Mukherjee (1951a, 1953) investigated the pollen morphology of mango
and 12 other Mangifera species. Their pollen grains were tricolpate of
almost the same size. Mondal et al. (1982, cited in Kostermans and Bom-
pard, 1993) attempted to correlate pollen morphology with taxonomic
relationships of 17 Mangifera species based upon different characteristics
of the exine and sporoderm. They demonstrated that all of the species of
section II (subgenus Limus) possess coarse exine; whereas there was no
clear correlation with pollen type in species within section I (subgenus
Mangifera).
The fruit
Description
The mango fruit is a large, fl eshy drupe, containing an edible mesocarp of
varying thickness. The mesocarp is resinous and highly variable with respect
to shape, size, colour, presence of fi bre and fl avour. The fl avour ranges from
turpentine to sweet. The exocarp is thick and glandular. There is a character-
istic beak that develops laterally on the proximal end of the fruit. A sinus is
always present above the beak. Fruit shape varies, including elongate,
oblong and ovate or intermediate forms involving two of these shapes. Fruit
length can range from 2.5 to > 30 cm, depending on the cultivar. The endo-
carp is woody, thick and fi brous; the fi bres in the mesocarp arise from the
endocarp.
The mango fruit is climacteric (see Brecht and Yahia, Chapter 14, this
volume), and increased ethylene production occurs during ripening. Chloro-
phyll, carotenes, anthocyanins and xanthophylls are all present in the fruit.
The skin is generally a mixture of green, red and yellow pigments, although
fruit colour at maturity is genotype dependent. During ripening the chloro-
plasts in the peel become chromoplasts, which contain yellow and red pig-
ments (Krishnamurthy and Subramanyam, 1970; Akamine and Goo, 1973;
Salunkhe and Desai, 1984; Mitra and Baldwin, 1997). Peel colour obviously is
cultivar dependent (see Knight et al., Chapter 3, this volume). Fruit of ‘Bom-
bay Green’ is green; ‘Carabao’, ‘Manila’, ‘Mulgoa’ and ‘Arumanis’ are greenish-
yellow; ‘Dashehari’ and ‘Alphonso’ are yellow; and ‘Haden’, ‘Keitt’ and
‘Tommy Atkins’ have a red blush. The red blush is due to the presence of
anthocyanins (Lizada, 1991). The pulp carotenoids in ripe fruit also vary with
respect to cultivar (Mitra and Baldwin, 1997).
4 S.K. Mukherjee and R.E. Litz
Flavour
Flavour of the mango mesocarp is a function of carbohydrates, organic acids,
lactones, monoterpene hydrocarbons and fatty acids (Mitra and Baldwin,
1997). During fruit maturation, starch that accumulates in the chloroplasts is
hydrolysed to sucrose, glucose and fructose (Medlicott et al., 1986; Selvaraj
et al., 1989; S. Kumar et al., 1994); sucrose is present in slightly higher concen-
trations than either fructose or glucose. Organic acid content decreases dur-
ing ripening (Krishnamurthy and Subramanyam, 1970). The dominant
organic acid is citric acid, but glycolic acid, malic acid, tartaric acid and oxalic
acids are also present (Sarker and Muhsi, 1981; Medlicott and Thompson,
1985). The peach-like fl avour of mangoes is attributed to the presence of lac-
tones (Lakshminarayana, 1980; Wilson et al., 1990).
Nutrition
Mango fruit contain amino acids, carbohydrates, fatty acids, minerals, organic
acids, proteins and vitamins. During the ripening process, the fruit are ini-
tially acidic, astringent and rich in ascorbic acid (vitamin C). Ripe mangoes
contain moderate levels of vitamin C, but are fairly rich in provitamin A and
vitamins B and B . Perry and Zilva (1932) determined the vitamin A, C and
1 2
D content of the fruit of three Indian mango cultivars, and found that the
pulp of mangoes is a concentrated source of vitamin C. The pulp of mango
fruit contains as much vitamin A as butter, although vitamin D is not present
in a signifi cant quantity. Fruit acidity is primarily due to the presence of malic
and citric acids. In addition, oxalic, malonic, succinic, pyruvic, adipic, galac-
turonic, glucuronic, tartaric, glycolic and mucic acids are also present (Jain et al.,
1959; Fang, 1965). Acidity is cultivar related; for example, immature Florida
cultivars have low acidity (0.5–1.0%) in comparison with ‘Alphonso’ (3%).
During ripening, acidity decreases to 0.1–0.2%. Following fruit set, starch
accumulates in the mesocarp. Free sugars, including glucose, fructose and
sucrose, generally increase during ripening; however, the sucrose content
increases three- to fourfold due to the hydrolysis of starch. Sucrose is the
principal sugar of ripe mangoes. The sucrose content of ripe fruit of three
Indian cultivars, ‘Alphonso’, ‘Pairie’ and ‘Totapuri’, ranges from 11 to 20%
representing 15 to 20% of the total soluble solids (Popenoe, 1932).
The seeds and polyembryony
Mango seeds are solitary, large and fl at, ovoid oblong and surrounded by the
fi brous endocarp at maturity. The testa and tegumen are thin and papery.
Embryos are dicotyledonous. Seeds of monoembryonic mango types contain
a single zygotic embryo, whose cotyledons can be unequal in size or lobed in
shape. The seeds of polyembryonic mango types contain one or more embryos
(Plate 2); usually one embryo is zygotic, whereas the remaining embryos are
derived directly from the nucellus, a maternal tissue. Nucellar embryos
apparently lack a suspensor. Polyembryony has also been reported in
Mangifera casturi, M. laurina and M. odorata (Bompard, 1993). Certain
Introduction: Botany and Importance 5
polyembryonic cultivars reportedly can produce seeds with adventitious
nucellar embryos only, for example ‘Strawberry’ (Juliano, 1934), ‘Carabao’
and ‘Pico’ (Juliano and Cuevas, 1932) and ‘Olour’ and ‘Cambodiana’
(Maheshwari et al., 1955). Early studies suggested that polyembryony
appeared to be a polygenic trait (Juliano, 1934; Sturrock, 1968), segregating as
a recessive character in the progeny of controlled crosses. Recent studies,
however, have demonstrated that the polyembryony trait is inherited as a
dominant character (Aron et al., 1998). Several studies have shown that nucel-
lar seedlings can be distinguished from the single zygotic seedling of poly-
embryonic seeds by isozymes (Schnell and Knight, 1992; Degani et al., 1993)
and DNA markers, for example single sequence repeats (SSRs) (Eiadthong et al.,
1999a), amplifi ed fragment length polymorphisms (AFLPs) (Kashkush et al.,
2001) and inter-simple-sequence-repeats (ISSRs) (Gonzalez et al., 2002). Mango
seeds are considered to be recalcitrant, and cannot survive for more than a
few days or weeks at ambient temperatures (Parisot, 1988). This important
characteristic of mango seeds would have inhibited the long distance dis-
persal of mango by seed until recent times.
1.3 History of Cultivation
Origin of Mangifera indica
The largest number of Mangifera species occurs in the Malay Peninsula, the
Indonesian archipelago, Thailand, Indochina and the Philippines (Mukher-
jee, 1985; Bompard, 1989; see Bompard, Chapter 2, this volume). The most
recent classifi cation of Mangifera species was based upon fl oral morphology
(Kostermans and Bompard, 1993) and included 69 species, most of which are
included in two subgenera Mangifera and Limus with another 11 species
occupying an uncertain position (Table 1.1). Eiadthong et al. (1999b) described
the phylogenetic relationships among Mangifera species using genomic
restriction fragment length polymorphisms (RFLPs) and amplifi cation of
chloroplast DNA (cpDNA), and suggested that the Mangifera species should
be classifi ed using molecular data. In the next few years, it is likely that
molecular biology will have a major impact on phylogenetic studies involving
mango and its relatives.
Mangifera species with a single fertile stamen are distributed in north-
eastern India, Myanmar, Thailand and the Malay Peninsula. Many of the
mango relatives have small fruits with thin, acidic fl esh, large seeds, abun-
dant fi bre and astringent resinous substances that are localized near the skin.
In addition to M. indica, edible fruit is produced by at least 26 other species
in the genus, primarily species found in South-east Asia (Gruezo, 1992).
Mangifera caesia, known as ‘binjai’ or ‘kemang’ in South-east Asia, is culti-
vated in Java, where it bears fruit in the mango off-season (Bompard, 1992a).
Mangifera foetida is less commonly cultivated due to its highly astringent
fruit; however, the fruit is widely used for pickling and as a substitute for
tamarind (Bompard, 1992b). Mangifera kemang and M. altissima are consumed
6
Table 1.1. Classifi cation of Mangifera species according to Kostermans and Bompard (1993).
Genus Subgenus Section Species
Mangifera Mangifera Marchandora Pierre M. gebede Miq
Euantherae Pierre M. caloneura Kurz
M. cochinchinensis Engler
M. pentandra Hooker f.
Rawa Kosterm. M. andamanica King M. minutifolia Evard. S.K
M. gracilepes M. nicobarica Kosterm. . M
M. griffi thii Hooker f. M. paludosa Kosterm. u
k
M. merrillii Mukherji M. parvifolia Boerl. & Koorders h
e
M. microphylla Griff. ex Hooker f. rje
e
Mangifera Ding Hou M. altissima Blanco. M. mucronulata Bl. a
n
M. applanata Kosterm. M. oblongifolia Hooker f. d
M. austro-indica Kosterm. M. orophila Kosterm. R
.E
M. austro-yunnanensis Hu M. pedicellata Kosterm. . L
M. casturi Kosterm. M. pseudo-indica Kosterm. itz
M. collina Kosterm. M. quadrifi da Jack
M. dewildei Kosterm. M. rigida Bl.
M. dongnaiensis Pierre M. rubropetala Kosterm.
M. fl ava Evard. M. rufocostata Kosterm.
M. indica L. M. similis Bl.
M. lalijiwa Kosterm. M. sulauesiana Kosterm.
M. laurina Bl. M. sumbawaensis Kosterm.
M. linearifolia (Mukherji) Kosterm. M. sylvatica Roxb.
M. longipetiolata King M. swintonioides Kosterm.
M. magnifi ca Kochummen M. timorensis Bl.
M. minor Bl. M. torquenda Kosterm.
M. monandra Merr. M. zeylanica (Bl.) Hooker f.
Limus (Marchand) M. blommesteinii Kosterm. M. leschenaultii Marchand
Kosterm. M. caesia Jack M. macrocarpa Bl.
M. decandra Ding Hou M. odorata Griff.
M. foetida Lour. M. pajang Kosterm.
M. kemanga Bl. M. superba Hooker f.
M. lagenifera Griff.
Species of uncertain M. acutigemma Kosterm. M. persiciformis Wu & Ming
position M. bompardii Kosterm. M. subsessifolia Kosterm.
M. bullata Kosterm. M. taipa Buch.-Hamilton
M. campospermoides M. transversalis Kosterm.
Kosterm. In
M. hiemalis Liang Jian Ying M. utana Utana tro
d
M. maingayii Hooker f. u
c
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8 S.K. Mukherjee and R.E. Litz
as fresh fruit or used green as a salad (Angeles, 1992; Bompard, 1992a).
Mangifera pajang has the largest fruit in the genus, and is an attractive fruit.
Mangifera odorata is grown in the Philippines and Indonesia, and has occa-
sionally been used as a rootstock for mango (Ochse, 1931; Bompard, 1992c).
Mangifera odorata is widely grown in the humid lowlands of South-east Asia
in areas that are unsuitable for mango as a mango substitute. Mangifera lau-
rinaand M. pentandra are appreciated as salad ingredients (Bompard, 1992d).
In addition, M. griffi thii, M. minor, M. monandra, M. quadrifi da and M. similis
have palatable fruit that are considered to have great potential (Gruezo,
1992). All mango cultivars belong to the species M. indica.
According to De Candolle (1884), ‘It is impossible to doubt that it (the
mango) is a native of south Asia or of the Malay Archipelago, when we see
the multitude of varieties cultivated in those countries, the number of ancient
names, in particular a Sanskrit name, its abundance in the gardens of Bengal,
of Deccan peninsula, and of Ceylon even in Rheede’s time (i.e. 1683).’
Although the centre of origin and diversity of the genus Mangifera is now
fi rmly established as being in South-east Asia, the origin of M. indica has been
a matter of speculation for many years. The fossil record provides few clues,
as only a single fossil bearing the imprint of a leaf of M. pentandra has ever
been found (Seward, 1912). Mangifera indica is believed to have fi rst appeared
during the Quatenary period (Mukherjee, 1951b). Blume (1850) considered
that mango might have originated from several related species, primarily
located in the Malay archipelago.
On the basis of ancient accounts of travellers and the written historical
record, it was believed for many years that mango must have originated in
India and spread outwards from there to South-east Asia and thence to the
New World and Africa. Because north-eastern India is at the northernmost
edge of the distribution of the Mangifera species, Hooker (1876) suggested
that mango might have been naturalized in India. The historical record pro-
vides a sometimes confl icting account of the distribution of mango. Miquel
(1859) did not record it as being wild in the Indonesian archipelago. Accord-
ing to Rumphius (1741), the mango was introduced into certain islands of the
Indonesian archipelago within recent times; however, the mango was in cul-
tivation in Java at least as early as ad 900–1100, when the temple at Borobo-
dur was built and faced with carvings of the Buddha in contemplation under
a mango tree (Plate 3). Based upon taxonomic and recent molecular evidence,
it is now apparent that the mango probably evolved within a large area
including north-western Myanmar, Bangladesh and north-eastern India (see
Bompard, Chapter 2, this volume).
Polyembryonic and monoembryonic M. indica
Within M. indica, there are two distinct types that can be distinguished on the
basis of their mode of reproduction and their respective centres of diversity:
a subtropical group with monoembryonic seed (Indian type) and a tropical
group with polyembryonic seed (South-east Asian). A few polyembryonic
cultivars occur along the west coast of India; however, they may have been
introduced into Goa from South-east Asia, perhaps by the Portuguese from
Introduction: Botany and Importance 9
their colonies of Malacca in the Malay Peninsula or Timor in the Indonesian
archipelago. Kumar et al. (2001) estimated the genetic relatedness among ten
polyembryonic and monoembryonic cultivars from the west coast of south-
ern India using genomic and chloroplast DNA RFLP analysis. The cultivars
could be grouped on the basis of embryo type (i.e. monoembryonic and poly-
embryonic) and had distinctly different genetic backgrounds. They con-
cluded that polyembryonic mangoes could not have originated in India, and
must have been introduced, probably from South-east Asia.
Domestication of mango
Historical record
It is probable that mango cultivation originated in India, where De Candolle
(1884) estimated that mango cultivation appeared to have begun at least 4000
years ago. In the early period of domestication, mango trees probably yielded
small fruit with thin fl esh. Such fruit can be found today in north-eastern
India and in the Andaman Islands (Anonymous, 1992). Folk selections of
superior seedlings over many hundreds of years would have resulted in
larger fruit with thicker fl esh. Mukherjee (1950a, b) described many of these
primitive selections from Orissa in north-eastern India; they demonstrated
great variation in fruit shape and size.
The mango is a very important cultural and religious symbol of India.
Buddhist pilgrims Fa-Hien and Sung-Yun mentioned in their travel notes
that the Gautama Buddha was presented with a mango grove by Amradarika
(c.500 bc) as a place for meditation (Popenoe, 1932). According to Burns and
Prayag (1921), a mango tree is depicted in friezes on the stupa of Bharut,
which was constructed c.100 bc. Other travellers to India, including the Chi-
nese Hwen T’sung (ad 632–645), the Arabs Ibn Hankal (ad 902–968) and Ibn
Batuta (ad 1325–1349) and the Portuguese Lurdovei de Varthema (ad 1503–
1508), all described the mango. The Indian subcontinent was the birthplace
of some of the earliest highly developed civilizations, and over the centuries,
India exerted strong cultural, religious and commercial infl uence over South
and South-east Asia. In successive waves, Hinduism, Buddhism and Islam
were introduced into South-east Asia from India. To this day, many com-
monly used words in Indonesia are derived from both Sanskrit and Tamil.
One of the most widely used words for mango in Malaysia and Java (Indone-
sia) is ‘mangga’, which is derived from the Tamil ‘manga’. Traders and monks
from India possibly introduced superior selections of mango into South-east
Asia; however, vegetative propagation was unknown in India until after the
arrival of the Portuguese in Goa in the 15th century. Moreover, the most im-
portant mango selections of Thailand, Cambodia, Vietnam, Malaysia, Indo-
nesia and the Philippines historically have all been of the polyembryonic
type, and have traditionally been seed propagated. Until the establishment
of Portuguese enclaves on the coast of India beginning in the late 15th century,
mango cultivars did not exist in India, as there was no known method
for vegetatively propagating superior selections (see Iyer and Schnell,