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© CAB International 2009.

The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 1
1 Introduction: Botany and Importance
S.K. Mukherjee
1
and R.E. Litz
2
1
Calcutta University, Calcutta, India
2
University 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
S.K. Mukherjee and R.E. Litz 2
crops has lagged far behind field crops for several reasons: their heterogene-
ity, polyploidy, lengthy juvenile period, time required for evaluation of trees
in the field, 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 field practices, which include irrigation management, control
of flowering, etc. Agricultural practices are currently undergoing another
revolution, as integrated pest and disease management replaces the earlier
reliance on agrichemicals, and emerging fields 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 flushes. 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 flowers are borne on terminal pyramidal panicles, and are glabrous
or pubescent; the inflorescence is rigid and erect, up to 30 cm long, and is
widely branched, usually tertiary, although the final branch is always cymose.
The inflorescence is usually densely flowered with hundreds of small flow-
ers, which are 5–10 mm in diameter. The flowers are either monoecious or
polygamous, and both monoecious and polygamous flowers are borne
within a single inflorescence (Plate 1). The pistil aborts in male flowers. The
ratio of monoecious to polygamous flowers is strongly influenced by
Introduction: Botany and Importance 3
environmental and cultural factors. The flowers have four or five sepals and
petals that are ovate to ovoid to lanceolate and also thinly pubescent. The
floral disc also is four- or five-lobed, fleshy and large and located above the
base of the petals. There are five large, fleshy 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 filaments arise
from the lobes of the nectaries. The stamens are central. The ovule is anatro-
pous and pendulous. It is believed that the flowers are cross-pollinated by
flies (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, fleshy drupe, containing an edible mesocarp of
varying thickness. The mesocarp is resinous and highly variable with respect
to shape, size, colour, presence of fibre and flavour. The flavour 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 fibrous; the fibres 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).
S.K. Mukherjee and R.E. Litz 4
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 flavour 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
1
and B
2
. Perry and Zilva (1932) determined the vitamin A, C and
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 significant 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 flat, ovoid oblong and surrounded by the
fibrous 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), amplified 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 classification of Mangifera species was based upon floral 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 amplification of
chloroplast DNA (cpDNA), and suggested that the Mangifera species should
be classified 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 flesh, large seeds, abun-
dant fibre 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
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Table 1.1. Classification 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.
M. gracilepes M. nicobarica Kosterm.
M. griffithii Hooker f. M. paludosa Kosterm.
M. merrillii Mukherji M. parvifolia Boerl. & Koorders
M. microphylla Griff. ex Hooker f.
Mangifera Ding Hou M. altissima Blanco. M. mucronulata Bl.
M. applanata Kosterm. M. oblongifolia Hooker f.
M. austro-indica Kosterm. M. orophila Kosterm.
M. austro-yunnanensis Hu M. pedicellata Kosterm.
M. casturi Kosterm. M. pseudo-indica Kosterm.
M. collina Kosterm. M. quadrifida Jack
M. dewildei Kosterm. M. rigida Bl.
M. dongnaiensis Pierre M. rubropetala Kosterm.
M. flava 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. magnifica Kochummen M. timorensis Bl.
M. minor Bl. M. torquenda Kosterm.
M. monandra Merr. M. zeylanica (Bl.) Hooker f.
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Limus (Marchand)
Kosterm.
M. blommesteinii Kosterm. M. leschenaultii Marchand
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
position
M. acutigemma Kosterm. M. persiciformis Wu & Ming
M. bompardii Kosterm. M. subsessifolia Kosterm.
M. bullata Kosterm. M. taipa Buch.-Hamilton
M. campospermoides
Kosterm.
M. transversalis Kosterm.
M. hiemalis Liang Jian Ying M. utana Utana
M. maingayii Hooker f.
S.K. Mukherjee and R.E. Litz 8
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-
rina and M. pentandra are appreciated as salad ingredients (Bompard, 1992d).
In addition, M. griffithii, M. minor, M. monandra, M. quadrifida 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
firmly 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 first 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 conflicting 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 flesh. 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 flesh. 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 influence 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,
S.K. Mukherjee and R.E. Litz 10
Chapter 4, this volume). However, under the Moghul emperor Akbar (1556–
1605), the best selections of seedling mangoes were propagated by approach
grafting and were planted in large orchards. The ‘Lakh Bagh’, a mango
orchard of 100,000 trees, was planted near Darbhanga in Bihar. Perhaps noth-
ing more eloquently attests to the importance of this fruit and the esteem in
which it was held than this vast mango orchard. The Ain-i-Akbari, an ency-
clopedic work that was written during the reign of Akbar, contains a lengthy
account of the mango, and includes information about the quality of the fruit
and varietal characteristics. There was evidently a strong body of informa-
tion about mango cultivation that had accumulated up to that time. Most of
the mango cultivars of India had their origin in those years, and have been
maintained under cultivation for over 400 years by vegetative propagation.
‘Alphonso’, ‘Dashehari’, ‘Langra’, ‘Rani Pasand’, ‘Safdar Pasand’ and other
mango cultivars were selected during that time. Relics of orchards from the
time of Akbar are found in different parts of India, and it has been suggested
that they could still provide valuable material for selection of superior mango
cultivars.
Distribution
Spreading from the centres of domestication
The global spread of mangoes and their cultivation outside their original
centres of domestication probably did not occur until the beginning of the
European voyages of discovery and colonialization in the 15th and 16th
centuries. Because mango seeds are recalcitrant, and cannot survive for more
than a few days or weeks, mango germplasm in the early days must have
been transported as ripe fruit, seedlings or, later on, as grafted plants. It is
believed that the Portuguese transported the mango from their colonies in
India to their African colonies, although Purseglove (1972) suggested that it
might also have been introduced to Africa via Persia and Arabia in the 10th
century by Arab traders. The Portuguese later introduced the mango into
Brazil from their African colonies of Mozambique and Angola. Spaniards,
who encountered a mango-growing civilization in the Philippines after
Magellan’s passage across the Pacific Ocean, introduced polyembryonic
mango types to their New World colonies through the Pacific trading ports
of Mexico and Panama. The most important, traditional mango cultivar in
Mexico remains the ‘Manila’, reflecting its Philippine origin. ‘Carabao’ and
‘Manila’ are probably identical. The mango was introduced to the West Indies
in the mid- to late 18th century, probably from Brazil. The first introductions
of mango into Florida (USA) occurred in 1861, and involved the ‘No. 11’
polyembryonic seedling from Cuba. Seven years later, another polyembry-
onic selection, ‘Peach’ was introduced into the state (Knight and Schnell,
1993). Many of the early introductions into Florida proved to be unproduc-
tive, although ‘Mulgoba’ was planted on a small commercial scale (this culti-
var is referred to as ‘Mulgoa’ in India, ‘Mulgoba’ in the USA and ‘Malgoa’ in
Malaysia).
Introduction: Botany and Importance 11
Secondary ‘centres of diversity’
In 1910, a seedling of ‘Mulgoba’ came into production in Florida. Its fruit had
a highly attractive red blush, and appeared to bear more heavily than its
parent(s) (Wolfe, 1962). This selection was named ‘Haden’. Although ‘Haden’
was not superior with respect to fruit quality in comparison to the imported
germplasm from India, its genetic base was much wider. During the 20th
century, more introductions of mango germplasm into Florida occurred from
South-east Asia (the Philippines, Cambodia), India and elsewhere. It was at
one time believed that these introductions of mango germplasm created a
secondary centre of diversity of the species (Knight and Schnell, 1993).
‘Eldon’, ‘Glenn’, ‘Lippens’, ‘Osteen’, ‘Parvin’, ‘Smith’, ‘Springfels’, ‘Tommy
Atkins’ and ‘Zill’ are progeny of ‘Haden’. ‘Saigon’ seedlings were selections
made from ‘Cambodiana’, a polyembryonic introduction from Indochina.
From ‘Saigon’ seedlings, ‘Alice’, ‘Herman’ and ‘Florigon’ were selected.
Based upon more recent genetic analysis involving microsatellite markers, it
is now estimated that the majority of Florida cultivars are descended from
only four monoembryonic Indian mango cultivar accessions, i.e., ‘Mulgoba’,
‘Sandersha’, ‘Amini’ and ‘Bombay’, together with the polyembryonic ‘Tur-
pentine’ from the West Indies (Schnell et al., 2006). The Florida mango culti-
vars have been found to be highly adaptable to many agroecological areas
and bear regularly, whereas many of the outstanding Indian cultivars have
been unproductive outside their centre of domestication, and are alternate
bearing. These selections also have a highly attractive red blush at maturity,
firm flesh, a high flesh to seed ratio and a regular bearing habit. Some of the
Florida cultivars, for example ‘Tommy Atkins’, ‘Keitt’, etc. are also moder-
ately resistant to anthracnose, the most important production and posthar-
vest problem of mango in many areas. In the latter half of the 20th century,
plantings of Florida cultivars have been established in many countries and
now form the basis of international trade of mangoes.
Current distribution
The mango is cultivated commercially throughout the tropics and in many
subtropical areas. It is grown at the equator and at a latitude of 35–37 in
southern Spain. According to Knight and Schnell (1993), ‘The process that
began in Florida – introduction of superior germplasm from abroad followed
by selection of improved cultivars adapted to local conditions – is now
underway in many areas.’
1.4 Germplasm Conservation
Genetic erosion
The Mangifera species have their centre of diversity and origin in South-east
Asia, a region that has experienced great economic development in recent
years. Vast wooded areas have been completely or partially deforested either
for expanding agriculture or for removal of tropical hardwoods for export.
S.K. Mukherjee and R.E. Litz 12
This has caused great genetic erosion within many species and genera. The
Mangifera species, like many other tropical fruit trees, are canopy and emer-
gent trees of the tropical rainforest (Kaur et al., 1980). These trees are widely
scattered in the tropical rainforest, flower erratically and reproduce from
large seeds that deteriorate rapidly. As such, they are particularly vulnerable
and in danger of extinction.
Collection and documentation of Mangifera germplasm
The International Plant Genetic Resources Institute (IPGRI), formerly known
as the International Board for Plant Genetic Resources (IBPGR), commis-
sioned an ecogeographical study of known Mangifera genetic resources (Muk-
herjee, 1985). Based upon this documentation, a joint IBPGR-International
Union for the Conservation of Nature (IUCN)-World Wildlife Fund (WWF)
project was initiated to collect wild mangoes on the island of Borneo and in
the Malay Peninsula (Bompard, 1989), the regions that held the highest con-
centrations of Mangifera species. Kostermans and Bompard (1993), in the lat-
est revision of the taxonomy of Mangifera, recognized 69 species, many of
which were collected during the course of this project (Table 1.1). Because of
the loss of natural habitat, the establishment of in situ and ex situ germplasm
collections of Mangifera species was considered to be imperative.
Relevance of germplasm resources to mango improvement
The genetic improvement of mango hitherto has depended on the utilization
of the genetic variability found within a single species, M. indica. According
to Mukherjee (1985), ‘A concerted sampling strategy should be devised for ex
situ samples to meet urgent needs for use in research for improvement of the
crop through breeding or as rootstocks. Sources of resistance to mango mal-
formation, anthracnose, powdery mildew, gall midge are urgently needed.’
1.5 Importance of Mango
Cultivars
A partial list of the principal mango cultivars has been provided in Table 1.2.
This list includes many cultivars that were identified in a survey of world
mango production compiled by Watson and Winston (1984). The distribution
of mango cultivars outside their centres of domestication can be attributed
primarily to three historical events: (i) the movement of Indian varieties
(monoembryonic) along the trade routes of the Portuguese to Africa and
South America; (ii) the spread of South-east Asian varieties (polyembryonic)
across the Pacific Ocean to Central and South America by the Spaniards; and
(iii) the identification of improved mango cultivars initially in Florida and
Introduction: Botany and Importance 13
Table 1.2. Most important mango cultivars in major producing countries.
Continent Country Cultivars
Africa Cote d’Ivoire ‘Amelie’, ‘Kent’
Egypt ‘Alphonso’, ‘Bullock’s Heart’, ‘Hindi be Sennara’,
‘Langra’, ‘Mabrouka’, ‘Pairie’, ‘Taimour’, ‘Zebda’
Kenya ‘Boubo’, ‘Ngowe’, ‘Batawi’
Mali ‘Amelie’, ‘Kent’
South Africa ‘Fascell’, ‘Haden’, ‘Keitt’, ‘Kent’, ‘Sensation’, ‘Tommy
Atkins’, ‘Zill’
Asia Bangladesh ‘Aswina’, ‘Fazli’, ‘Gopal Bhog’, ‘Himsagar’, ‘Khirsapati’,
‘Langra’
China ‘Gui Fei’, ‘Tainong No. 1’, ‘Keitt’, ‘Sensation,’ ‘Zill’, ‘Zihua’,
‘Jin Huang’
India ‘Alphonso’, ‘Banganapalli’, ‘Bombay’, ‘Bombay Green’,
‘Chausa’, ‘Dashehari’, ‘Fazli’, ‘Fernandian’,
‘Himsagar’, ‘Kesar’, ‘Kishen Bhog’, ‘Langra’, ‘Mallika’,
‘Mankurad’, ‘Mulgoa’, ‘Neelum’, ‘Pairi’, ‘Samar
Behisht’, ‘Suvarnarekha’, ‘Totapuri’, ‘Vanraj’, ‘Zardalu’
Indonesia ‘Arumanis’, ‘Dodol’, ‘Gedong’, ‘Golek’,
'
Madu’,
'
Manalagi’
Israel ‘Haden’, ‘Tommy Atkins’, ‘Keitt’, ‘Maya’, ‘Nimrod’, ‘Kent’,
‘Palmer’
Malaysia ‘Apple Rumani’, ‘Arumanis’, ‘Golek’, ‘Kuala Selangor 2’,
‘Malgoa’
Myanmar ‘Aug Din’, ‘Ma Chit Su’, ‘Sein Ta Lone’,
'
Shwe Hin Tha’
Pakistan ‘Anwar Ratol’, ‘Began Pali’, ‘Chausa’, ‘Dashehari’,
‘Gulab Khas’, ‘Langra’, ‘Siroli’, ‘Sindhri’,
‘Suvarnarekha’, ‘Zafran’
The Philippines ‘Carabao’, ‘Manila Super’, ‘Pico’
Taiwan ‘Irwin’, ‘Jin-hwung’, ‘Keitt’, ‘Tommy Atkins’, ‘Tainong
No. 1’, ‘Tsar-swain’
Thailand ‘Nam Doc Mai’, ‘Ngar Charn’, ‘Ok Rong’, ‘Keow Savoey’,
‘Pimsen Mum’
Australia ‘Calypso’, ‘Kensington Pride’
North and
Central
America
Costa Rica ‘Haden’, ‘Irwin’, ‘Keitt’, ‘Mora’, ‘Tommy Atkins’
Dominican
Republic
‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
Guatemala ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
Haiti ‘Francine’, ‘Madame Francis’
Mexico ‘Ataulfo’, ‘Haden’, ‘Keitt’, ‘Kent’, ‘Manila’, ‘Palmer’,
‘Sensation’, ‘Tommy Atkins’, ‘Van Dyke’
USA ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
South
America
Brazil ‘Bourbon’, ‘Coite’, ‘Coquinho’, ‘Coracao’, ‘Espada’,
‘Haden’, ‘Itamaraca’, ‘Keitt’, ‘Mamao’, ‘Palmer’, ‘Rosa’,
‘Tommy Atkins’, ‘Uba’, ‘Van Dyke’
Colombia ‘Vallenato’
Ecuador ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
Peru ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
Venezuela ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
S.K. Mukherjee and R.E. Litz 14
later in other new mango-producing areas, as a result of open and controlled
pollination among local and introduced mango germplasm from India and
South-east Asia.
Further information about many of the mango cultivars, including their
fruit characters, is available in Knight et al. (Chapter 3, this volume), and in
publications by Burns and Prayag (1921) for mangoes of Maharashtra, Naik
and Gangolly (1950) for south Indian mangoes, Singh and Singh (1956) for
Uttar Pradesh mangoes, Mukherjee (1948) for Bengal mangoes and Camp-
bell (1992) for Florida mangoes.
Because many clonally propagated mango cultivars have unique local
and/or regional names, there is considerable confusion in nomenclature. The
Indian Agricultural Research Institute (IARI), New Delhi, has been recog-
nized by the International Society for Horticultural Science (ISHS) as the
International Registration Authority for Mango, whose mission is to consoli-
date superfluous names of mango cultivars. The potential for molecular, for
example randomly amplified polymorphic DNA (RAPD), markers, to resolve
much of this confusion has been demonstrated by Schnell and Knight (1992),
Degani et al. (1993), Schnell et al. (1995), Eiadthong et al. (1999a), Kashkush et al.
(2001) and Gonzalez et al. (2002) (see Bompard, Chapter 2 and Iyer and
Schnell, Chapter 4, this volume).
There is little variation among seedlings derived from polyembryonic
mangoes. None the less, a certain amount of variability does occur, probably
as a result of somatic mutation. Thus, in Indonesia there are several ‘Aru-
manis’ selections that are denoted numerically, for example ‘Arumanis 1’,
‘Arumanis 2’, etc. In addition, although Philippine mango cultivars are dis-
tinguished by different names, for example ‘Carabao’, ‘Manila’, ‘Philippine’,
etc., the differences among them are quite subtle.
1.6 Production and Uses
The mango is the most important fruit of Asia, and currently ranks fifth in
total production (in metric tonnes) among major fruit crops worldwide, after
Musa (bananas and plantains) (105,815,354 t), Citrus (all types) (105,440,168
t), grapes (65,584,233 t) and apples (59,444,377 t) (FAOSTAT, 2006). According
to the Food and Agriculture Organization of the United Nations (FAO)
database (FAOSTAT, 2006), world mango production has increased from
16,903,407 t in 1990 to 28,221,510 t in 2005. Much of this new production has
occurred outside the traditional centres of mango culture of South and
South-east Asia. In 1990, India produced approximately 51% of the world’s
mangoes, but by 2005, India’s share had declined to approximately 38%,
despite the substantial increase in mango production since 1990 (from 8,645,405
to 10,800,000 t between 1990 and 2005). The current leading producing nations
after India include (in metric tonnes) China (3,450,000), Thailand (1,800,000),
Pakistan (1,673,900), Mexico (1,600,000), Indonesia (1,478,204), Brazil (1,000,000)
and the Philippines (950,000). Although world production has increased by
67% between 1990 and 2005, mango exports have increased almost sixfold
Introduction: Botany and Importance 15
from 158,030 to 907,782 t, with total export value estimated to be
US$583,763,000 (FAOSTAT, 2006). The major exporting countries are (in met-
ric tonnes) Mexico (212,505), India (156,222) and Brazil (111,181). As a result,
mangoes are widely available as fresh fruit and as processed products (i.e.
dried fruit, dairy products, juice, pickles, etc.).
Mangoes are an important component of the diet in many less developed
countries in the subtropics and tropics. In regions of the world that have
experienced low living standards and serious nutritional deficiencies, their
attractiveness and flavour have also enhanced the quality of life. Surplus
production has increasingly been processed and fruit of certain cultivars is
destined for export as fresh fruit. Approximately 1% of mango production is
utilized for processing for juice, nectars, preserves (including chutney), fruit
leather, dried fruit slices, frozen pulp and as a flavouring for baked goods,
ice cream, yoghurt, etc. (see Raymundo et al., Chapter 17, this volume). No
part of the fruit is wasted. In India and the subcontinent, the seed is used for
extraction of starch ‘amchur’, and the peels (skin) have been used as a source
of anacardic acid. Mango wood is a low quality timber, and the bark of the
tree is an important source of tannins for curing leather.
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© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 19
2 Taxonomy and Systematics
J.M. Bompard
Les Mazes, Montaud, France
2.1 Introduction 19
2.2 The Genus Mangifera L. 20
Distribution 20
Ecology and habitat 20
2.3 Taxonomy and Systematics 22
Taxonomic history 22
2.4 Phytogeography 28
Species distribution 28
Subgenera and section distribution 29
2.5 Interspecific Molecular Characterization 30
2.6 Region of Origin of the Genus 31
2.7 Origin of the Common Mango 32
The common mango in South-east Asia 32
2.8 Conclusion 35
Potential contribution of wild species to mango cultivation 35
Source of rootstock 35
Hybridization 36
Potential of wild species 36
2.1 Introduction
The genus Mangifera is one of the 73 genera (c.850 species) belonging to the
family of Anacardiaceae, in the order of Sapindales. Anacardiaceae is a fam-
ily of mainly tropical species, with a few representatives in temperate regions.
Malesia, which is the phytogeographic region extending from the Malay
Peninsula south of the Kangar-Pattani line to the Bismarck Archipelago east
of New Guinea (Whitmore, 1975) contains more species in the Anacardiaceae
than any other area. Within Malesia occurrence is mainly in Western Malesia
(Ding Hou, 1978b).
J.M. Bompard 20
Apart from mango, several other cultivated fruit trees belong to the fam-
ily, for example the ambarella or Otaheite apple (Spondias dulcis Forst.) prob-
ably from Melanesia, and the yellow and purple mombins (Spondias mombin
L. and S. purpurea L., respectively) from tropical America, the Bouea species
from IndoMalesia, dragon plums (Dracontomelum spp.) from IndoMalesia
and the Pacific region, kaffir plum (Harpephyllum caffrum Bernh. ex K. Krause)
and the marula plum (Sclerocarya caffra Sond.) of southern Africa. The cashew
(Anacardium occidentale L.) is from tropical America and the pistachio (Pistacia
vera L.) from Iran and Central Asia. Anacardiaceous species also yield other
valuable products: wood (several genera), gums and resins (Pistacia spp.),
varnishes (Rhus spp. and Melanorrhoea spp., ‘lacquer trees’) and tanning
materials (Rhus spp. and Schinopsis spp.). It is also a family well known for
the dermal irritation produced by some of its members, such as the poison
ivies and oaks (Rhus spp.) in North America, rengas (Gluta spp.) in South-
east Asia and other species including some Mangifera species whose resinous
sap may induce a mild to strong allergic reaction.
2.2 The Genus Mangifera L.
Distribution
The range of natural distribution of the 69 Mangifera species is mainly
restricted to tropical Asia, and extends as far north as 27° latitude and as far
east as the Carolines Islands. Wild mangoes occur in India, Sri Lanka, Ban-
gladesh, Myanmar, Sikkim, Thailand, Kampuchea, Vietnam, Laos, southern
China, Malaysia, Singapore, Indonesia, Brunei, the Philippines, Papua New
Guinea and the Solomon and Carolines Islands. The highest species diver-
sity, c.29 species, occurs in western Malesia, especially in peninsular Malay-
sia and in Borneo and Sumatra, which represent the heart of the distribution
range of the genus (Fig. 2.1).
Ecology and habitat
The majority of Mangifera species occur as a rule as scattered individuals in
tropical lowland rainforests on well-drained soils. The species are distrib-
uted mostly below 300 m, but can occur up to c.1000 m above sea level, on
well-drained soils (44 species), in periodically inundated areas (ten species)
and in certain types of swamp forest (i.e. M. gedebe, M. griffithii and M. parvi-
folia). Three species are mainly found in sub-montane forests above 1000 m
and occasionally up to 1700 m above sea level (M. bompardii, M. dongnaiensis
and M. orophila). There are also species that are adapted to seasonally dry
climates in deciduous or semi-deciduous forests (e.g. M. caloneura, M. collina,
M. timorensis and M. zeylanica). A few species occur north of the Tropic of
Cancer, for example M. austro-yunnanensis and M. persiciformis in China, M.
Taxonomy and Systematics 21
sylvatica Roxb. in Sikkim and southern China, at altitudes of 600–1900 m above
sea level; apparently wild M. indica can also be found outside the tropics.
Wild mangoes are large trees, 30–40 m (occasionally 54 m) in height, with
tall columnar boles. Several species are exploited for their timber. The major-
ity of wild mangoes occur as scattered individuals at very low densities in
lowland forests on well-drained soils. Some of these are very rare; there are
normally one to three trees above 40 cm in diameter/10 ha. Only a few spe-
cies (M. gedebe, M. griffithii and M. parvifolia) are gregarious in certain types of
swamp forest. Most species are evergreen although a few are deciduous in
the rainforests following a dry period, and stand bare for a short time before
flushes of new leaves appear. A deciduous habit that is not linked to a sea-
sonal climate also occurs in other genera of Anacardiaceae (Ding Hou, 1978b).
In the rainforest of western Malesia, Mangifera species flower and fruit
very irregularly. As with many other genera in the region, mast or general
fruiting at intervals of 3–8 years is the dominant pattern. In mast years, the
ground beneath the trees can be covered with mangoes, whose strong smell
attracts many animals. Isolated flowering may occur after a dry period and
is generally followed by a poor fruit crop. The occurrence of flowering of a
few species, for example the ‘lanjut’ (M. lagenifera) is only once every 5–10
years. There seem to be clear reproductive barriers between species in the
wild, although limited hybridization among cultivated species has been
reported (see section 2.8, Conclusion, this chapter).
30° N
4
2
5
4
3
6
13
27
27
28
9
5
7
4
3
5
2
20° N
10° N
0° N
10° N
30° N
20° N
10° N
0° N
10° N
80° 100 120 160
80° 100 120 160 140
Fig. 2.1. Distribution of Mangifera species in the range of the genus. Numbers shown indicate
the number of wild species in each area: Sri Lanka, India and Sikkim, Andaman and Nicobar
Islands, Myanmar, Thailand, Indochina, China, peninsular Malaysia, Sumatra, Borneo, Java,
Lesser Sunda Islands, Sulawesi, Moluccas, the Philippines, New Guinea, and Solomon Islands
(the Caroline Islands not represented).
J.M. Bompard 22
These widely scattered towering tree species, often with an inaccessible
crown, are undercollected and poorly represented in herbarium collections
(Bompard, 1995). Because of their irregular flowering, the flowers and fruits
of a few species are still unknown. Collecting plant material is consequently
very difficult, and plant explorations are still yielding new records or new
species. Many species have been recently recorded for the first time, even
from peninsular Malaysia, a country that has already been rather well combed
by botanists, having one of the highest collecting indices in the Malesian
region. Other species still await to be discovered. Sadly some species of very
limited range may already have been lost to posterity by deforestation.
Our very meagre knowledge of the wild mangoes is due to the fact that
identification at the species level from leaves only is often difficult because of
intraspecific variation in vegetative characters. Moreover, many of the origi-
nal species were based on very poor specimens. Consequently, frequent mis-
identification of herbarium material has resulted in much confusion, requiring
a critical revision of all the specimens in these collections. It is not uncommon
that the same species has been described from different places under differ-
ent names. For instance, M. inocarpoides described by Merrill and Perry from
New Guinea in 1941, M. camptosperma and M. reba (recorded by Pierre in
South Vietnam in 1897) are now recognized to be a single species M. gedebe
Miquel, a species initially named in 1861 from a specimen collected in Suma-
tra. Mangifera longipes Griffith is now treated as M. laurina Blume, because
this name takes precedence as it was validly published 4 years earlier.
After thorough study of herbarium collections and field collections, a
number of species have been newly described. Sixty-nine species are now
recorded, including 13 species of uncertain affinities, in contrast with the
49 species recognized by Mukherjee (1949). As more collections are made,
there will doubtless be further taxonomic adjustments made to the genus
Mangifera.
2.3 Taxonomy and Systematics
Taxonomic history
Subdivision of the genus
An historical review of the subdivisions of the genus Mangifera shows that
two major groups have been rather consistently recognized in taxonomic
treatments. Hooker (1862) was the first to recognize two sections based on
the characters of the flower disc: section I with a disc broader than the ovary,
and section II with a disc stalk-like or wanting. These sections were later
named by Marchand (1869) Amba, an Indian name for the common mango,
and Limus, a Sundanese name for M. foetida in West Java, respectively. He
also added a section Manga for M. leschenaultii, which in fact belongs to the
section Limus.
In his monograph of the Anacardiaceae, Engler (1883) maintained Hook-
er’s sections, and subdivided group A (Hooker’s section I) into two groups,
Taxonomy and Systematics 23
one group with four or five petals and the other group with four petals. He
considered the following sequence of morphological characters to be impor-
tant for classification: (i) texture of the leaves; (ii) number of fertile stamens;
(iii) prominence of veins; (iv) pilosity of inflorescences; and (v) leaf shape.
Pierre (1897) further divided the genus Mangifera into five sections based
on flower characters, i.e. number of stamens, the attachment of stamens to
the disk, and the style. Two of these five sections – namely section I Euan-
therae, with a short thick flower disc and 4–12 fertile stamens, and section V
Marchandora then consisting of M. camptosperma (currently considered a syn-
onym of M. gedebe) are still maintained as they form clear-cut sections.
In his monograph, Mukherjee (1949) recognized two unnamed sections,
conserving Hooker’s subdivision. Ding Hou (1978a) adopted the same
method in his revision of the Malesian Anacardiaceae recognizing only
Hooker’s two original sections and providing them with proper names and
synonyms: section Mangifera (section I Hooker, section Amba Marchand,
group A Engler, sections Euantherae and Marchandora Pierre) and section
Limus (section II Hooker, sections Limus and Manga Marchand, group B
Engler, and sections Eudiscus and Microdiscus Pierre).
Most recent classification of the genus
The taxonomic classification referred to herein follows that proposed by Kos-
termans and Bompard (1993). This treatise includes the results of collections
and surveys carried out between 1986 to 1998 in Borneo and peninsular
Malaysia, which were initiated and sponsored by the International Institute
for Genetic Resources (now Biodiversity International) and the World Wide
Fund for Nature.
1
It was published under the auspices of the International
Board for Plant Genetic Resources (now International Plant Genetics Resources
Institute) and the Linnean Society of London.
The most recent treatment of Mangifera reflects the current status of what
is still fragmentary knowledge. It can provide a basis for further studies
involving all aspects of the wild relatives of mango, but particularly their
potential in mango breeding. Determining phylogenetic affinities based upon
molecular markers could change our thinking about relationships among
Mangifera species and among the cultivated forms of M. indica (see Interspe-
cific Molecular Characterization section, this chapter).
The morphological characters used for identification have been placed in
the following sequence of importance:
Shape of the floral disc (see section Subdivision of the genus). 1.
Number of fertile stamens. 2.
Seed labyrinthine or not. 3.
Shape of secondary branches of the inflorescences: open or lax panicle, 4.
flowers glomerulate or sub-glomerulate, the ramifications racemoid or
spike like.
Pubescence of the inflorescence. 5.
Shape, number and attachment of the nerves (ridges or fingers) at the 6.
inner surface of the petals.
J.M. Bompard 24
Shape and size of the petals. 7.
Flowers tetra- or pentamerous (not a very constant character and often 8.
overlapping).
Reticulation of the leaves, especially of the lower surface. 9.
Shape of the leaf (only fully grown leaves of sterile branches can be used). 10.
Texture of the leaves. 11.
Deciduous or non-deciduous trees. 12.
Colour of the flowers. 13.
Shape, colour and smoothness of the fruit. 14.
Number and size of the stone fibres. 15.
Kostermans (Kostermans and Bompard, 1993) raised the sections to the
rank of subgenus, i.e. subgenus Limus (Marchand) Kosterm., having a disc
narrower than the base of the ovary, stalk-like or even lacking and subgenus
Mangifera (Ding Hou) Kosterm., having a disc broader than the base of the
ovary, cushion-like, often divided in four or five lobes.
SUBGENUS LIMUS (MARCHAND) KOSTERM. Mangifera species of the subgenus Limus
are quite distinctive and show only remote affinity with the common mango.
This taxon is more primitive than the subgenus Mangifera and may be ances-
tral to it, although the two subgenera may have originated from two different
ancestors. The subgenus Limus consists of 11 species, which are native to the
rainforests of western Malesia (peninsular Thailand, Malay Peninsula, Suma-
tra, West Java and Borneo), with the exception of M. foetida, which extends to
the east, possibly as far as New Guinea, and M. odorata which is only known
in cultivation.
Kostermans divided the subgenus Limus into two sections: (i) section
Deciduae for deciduous trees (i.e. M. caesia, M. kemanga, M. pajang, M. superba
and possibly M. blommesteinii, M. decandra and M. lagenifera); and (ii) section
Perennes for non-deciduous species (i.e. M. foetida, M. leschenaultii, M. macro-
carpa and M. odorata) (Kostermans and Bompard, 1993). In deciduous trees,
the bracts enclosing the buds leave a characteristic collar of dense, narrow
scars, which persist on old twigs and are especially prominent in M. caesia
and M. kemanga.
Mangifera lagenifera and M. decandra have ten stamens, five of which are
fertile. The other nine species have only one (and rarely two) fertile stamen(s)
and two to four staminodes. The two species with five fertile stamens (M.
decandra and M. lagenifera) and M. superba, M. caesia, M. kemanga and M. blom-
mesteinii, whose leaves are apically aggregated into rosettes at the end of mas-
sive twigs are particularly distinctive. The fruits of these species are broadly
ellipsoid or pear shaped, not compressed, and have dirty whitish or pinkish
mesocarp and lanceolate, and fibrous, non-ligneous leathery endocarp.
Mangifera subsessilifolia shows some affinity with M. lagenifera and M.
blommesteinii; however, it has been placed among the species of uncertain
taxonomic position due to a lack of complete study material. This is not a
very rare species, but flowering and fruiting seem to occur at intervals of, or
> 5 years, similar in this respect to M. lagenifera, which can be found growing
Taxonomy and Systematics 25
in old orchards in peninsular Malaysia. The flowers and fruits of M. sub-
sessilifolia are still unknown.
Mangifera foetida, M. odorata, M. caesia and M. kemanga are widely cultivated
in the humid lowlands of the Malay Peninsula, Sumatra, Borneo, Java and Bali.
They have also been introduced elsewhere in South-east Asia; M. caesia, M. foe-
tida and M. odorata are grown in the southern part of the Philippines, M. foetida
is grown in Myanmar, and M. odorata is found in Indochina. They have been
described in general reviews of tropical fruit (Ochse and Bakhuizen, 1931; Ochse
et al., 1961; Molesworth, 1967; Verheij and Coronel, 1991).
Mangifera pajang, an endemic and commonly cultivated species in Bor-
neo, is hardly known outside its native island. This deciduous tree has very
stout twigs, with leaves more or less aggregate at the apices. The globose
fruits, up to 20 cm in diameter, are the largest known fruits in the genus. The
rough, potato-brown rind (0.5–1 cm thick) can be peeled off like that of a
banana. Its bright, deep yellow, thick and fibrous flesh is sweet with a dis-
tinctive taste (Kostermans, 1965; Bompard, 1991a). In orchards in Borneo
where M. foetida and M. pajang are both cultivated, forms with leaves and
fruits having intermediate characters are occasionally found.
Mangifera caesia, M. foetida, M. pajang and especially M. odorata are impor-
tant in tropical humid regions where the common mango cannot be grown
satisfactorily. Mangifera pajang has potential as an ornamental tree, having
brilliant rose-red blossoms (Philipps et al., 1982).
SUBGENUS MANGIFERA. The subgenus Mangifera contains most of the species (47),
and is divided into four sections: (i) section Marchandora Pierre; (ii) section Euan-
therae Pierre; (iii) section Rawa Kosterm.; and (iv) section Mangifera Ding Hou.
Section Marchandora Pierre. This section has only one species, M. gedebe Miquel
(syn. M. camptosperma Pierre, M. inocarpoides Merr. and Perry, M. reba Pierre).
The labyrinthine seed is unique to this species, wherein the inner integu-
ments penetrate the cotyledons and form numerous irregular folds. The flat,
discus-like fruit has only a very thin mesocarp. Mangifera gedebe grows in
inundated places along rivers or lakes. The seed floats in water and is dis-
persed during periods of high water, and this may explain its wide distribu-
tion, from Myanmar through Malesia to New Guinea and the Bougainville
Island.
Section Euantherae Pierre. The three species in this section (M. caloneura Kurz
(syn. M. duperreana Pierre), M. cochinchinensis Engler and M. pentandra Hook.
f.) appear to be the most primitive among the species of the subgenus
Mangifera. The flowers are characterized by the presence of five fertile sta-
mens. The three species are mainly confined to Myanmar, Thailand, Indo-
china and the north of the Malay Peninsula. The region is in the transition
zone from the humid tropical rainforest to monsoon forest, and these species
show an adaptation to low rainfall. Mangifera cochinchinensis, which occurs in
south-eastern Thailand and in Vietnam, has small oblong fruits with a thin
seed; the fruits are much relished by local people in southern Vietnam,
although they are very acidic. Mangifera caloneura and M. pentandra are closely
J.M. Bompard 26
related, and can be mistaken for M. indica. However, their leaves are more
leathery, have a more conspicuously dense reticulation, and the panicles are
much more hirsute than the common mango. Mangifera caloneura occurs from
Myanmar through Thailand to Indochina, in lowland evergreen forests, as
well as in semi-deciduous forests. It is cultivated for its acidic-sweet fruit,
and has been planted along the streets of Vientiane and Ho Chi Minh City
(Saigon). Mangifera pentandra, apparently native to the northern Malay Pen-
insula close to the Kra isthmus transition zone, is found in old orchards, in
scattered locations, especially in Kedah and possibly also in peninsular Thai-
land. It is also grown in the Anambas Islands and in Sabah, where it might
have been introduced in early times. It is a prolific bearer, with small man-
goes, c.8 cm length, and ripening green or yellow. The pale orange, watery
pulp has a sweet taste and few fibres.
Section Rawa Kosterm. This group, consisting of nine species, is not well
delimitated. Most species have thick twigs and rather coriaceous leaves
seated on protruding pedestals. The small, hardly flattened ovoid or ellip-
soid fruits that are black or partly red at maturity in several species are also
characteristic. ‘Rawa’ is the Malay word for marsh, indicating that these spe-
cies usually are found in periodically or permanently inundated areas. The
five species that occur in west Malesia (M. gracilipes, M. griffithii, M. micro-
phylla, M. paludosa and M. parvifolia) grow primarily in the swamps of south
peninsular Malaysia, in central coastal areas of east Sumatra and western
Borneo, and occasionally in peripheral uplands. It has also been reported
from the Andaman Islands and from Thailand (Sreekumar et al., 1996; Eiad-
thong et al., 2000a).
Mangifera andamanica and M. nicobarica are endemics from the Andaman
and Nicobar Islands, respectively. Mangifera merrillii is a rare species endemic
to the Philippines and M. minutifolia is known solely from a single collection
from southern Vietnam. Mangifera griffithii and M. microphylla are the only
cultivated species within section Rawa. The former species is considered to
be representative of the section, and is cultivated along the eastern coast of
peninsular Malaysia and in western Borneo, and rarely in Sumatra. The fruits
are small (3–5 cm long) and oblong or ovoid; the skin is rose-red, turning
purplish black at maturity. The rind is thin and easily removed from the
orange-yellow pulp, which is juicy and pleasantly sweet. Different forms are
recognized by local people, according to the size and taste of fruits. Mangifera
microphylla is a related, but less well-known species, having thinner leaves
and a rather similar fruit.
Section Mangifera Ding Hou. With more than 30 species, section Mangifera is
by far the largest. The common mango and the related M. laurina belong
here. Species within the section have the same distribution range as the
genus. The section may be divided into three groups based on floral structure
and organ number variation: (i) those having pentamerous flowers; (ii) those
having tetramerous flowers; and (iii) an intermediate group of species hav-
ing both pentamerous and tetramerous flowers. Within these three groups, it
is possible to distinguish species with either puberulous or glabrous panicles.
Taxonomy and Systematics 27
Only characteristics of representative species within each group, especially
those found in cultivation, are described below.
Pentamerous flowers (14 species): Three species, M. laurina, M. minor and
M. sylvatica, show affinity with the common mango. Mangifera laurina is a
species of the lowland forests of Malesia, where it is also under cultivation in
old orchards. It can be distinguished from the common mango by having lax
and widely pyramidal, glabrous or sparingly puberulous panicles. The flow-
ers are smaller and are not glomerulate; the petals have a different shape,
texture and colour. The fruit resemble those of a small common mango, with
orange-yellow pulp, which is almost liquid at maturity. It is generally con-
sumed when unripe. Several forms are in cultivation; however these are now
becoming rare. Mangifera laurina is well suited to the humid tropical lowlands,
fruiting well in areas where the common mango cannot be grown satisfacto-
rily; moreover, it appears to be highly resistant to anthracnose (Bompard,
1991b).
Mangifera minor occurs east of Wallace’s line, from Sulawesi to New
Guinea (east Malesia) and to the Carolines Islands in the east. It is adapted to
a wide range of ecological conditions, growing equally well in dry savannahs
and in tropical rainforests up to 1300 m. The fruit is obliquely oblong, 5–10 cm
long, much narrowed, the tip obtuse, with a distinct beak and sinus. It is
found in cultivation, although the yellowish fruit pulp is acidic and scant.
Mangifera sylvatica is found from Sikkim (up to 1200 m) to northern Myan-
mar and Thailand, and apparently also in Yunnan up to 1900 m. The fruit is
obliquely ovate, 8–10 cm long, much compressed distally forming a hook,
has scanty whitish-yellow pulp which is almost fibreless. Other species are
occasionally found in cultivation, for example M. rufocostata, which is esteemed
by the Banjarese people of South Kalimantan for its very sour fruits that are
used to prepare a spicy condiment with chilli.
Tetramerous flowers (15 species): Mangifera altissima is apparently endemic
to the Philippines, where it occurs mainly at low elevations in the forests
from northern Luzon to Mindoro (Brown, 1950). The fruit is mango shaped,
ovoid or ellipsoid, slightly compressed, up to 8 cm length, green or some-
what yellow when ripe, with whitish, sweetish-acidic flesh. It is commonly
found in dooryards, and thrives in regions with distinct wet and dry seasons
(Angeles, 1991).
Mangifera torquenda occurs wild in west Malesia, and is cultivated in
south Sumatra and in Borneo, where it is common in the forests and orchards
of eastern Kalimantan. The sub-globose fruit, c.7.5 cm long and 6.5 cm in
diameter, is yellow-green with darker spots at maturity, and has a thin rind.
The pale yellow pulp has a rather pleasant sweet-acid, slightly resinous taste
and a light turpentine smell. Short fibres are attached to the seed. It is closely
related to M. longipetiolata.
Mangifera magnifica is a common species in the rainforests of western
Malesia, occasionally cultivated in central Sumatra and in West Kalimantan,
where it has a special importance in the myths of Land Dayak peoples. The
fruit is ovoid oblong, up to 12 cm long, 10 cm in diameter, only slightly
compressed, greyish green with brown spots. The pulp is whitish, soft at
J.M. Bompard 28
maturity, sweetish acid. Sweeter forms are reported in central Kalimantan
(J.J. Afriastini, personal communication). The stone is unique in the genus in
that it lacks fibres adhering to it.
Mangifera quadrifida is found from peninsular Malaysia to the Moluccas.
The fruit is ellipsoid-globose, 6–8 cm long, green covered with black dots turn-
ing completely black at maturity, and has a pale yellow, sweet-acid pulp.
Another form is recognized by its more coriaceous leaves, smaller fruits, c.4 cm
long, having dark yellow pulp, purplish around the stone, and a sweet, palat-
able taste, somewhat like prunes. Both forms are cultivated in old orchards.
Tetra- and pentamerous flowers (four species, and also M. indica): Mangifera
casturi is related to M. quadrifida, from which it can be distinguished by leaf
and fruit characters. It has never been collected in the wild, and is a favourite
among the Banjarese people in south Kalimantan. The fruits are small, a little
compressed and up to 6 cm in length, becoming completely black at matu-
rity. The orange pulp is very sweet and palatable, and resembles ‘honey
mango’ or ‘mangga madu’ grown in East Java. Although M. casturi bears
heavily, it has a strong- to alternate-bearing habit. It is an excellent fruit for
the humid tropical lowlands, and appears to be resistant to anthracnose. Sev-
eral differently named forms exist; these have polyembryonic seeds. Mangifera
rubropetala is also only known in cultivation, and may be a primitive race of
M. indica.
SPECIES OF UNCERTAIN TAXONOMIC POSITION. There is a group of 11 disparate spe-
cies of uncertain taxonomic position that cannot be placed with certainty due
to the absence of adequate material. There are three species only known in
China.
2.4 Phytogeography
Species distribution
An examination of the present distribution of the genus shows that the larg-
est number of Mangifera species in either subgenera is found in western
Malesia on the Sunda shelf. A decreasing number of species occurs towards
the genus boundary east of Wallace’s line in east Malesia, and in its northern
and western range of distribution. While peninsular Malaysia and the islands
of Sumatra and Borneo have the highest diversity of species, the number of
species becomes gradually lower in east Malesia, especially in the Lesser
Sunda Islands, Moluccas and New Guinea. This is explained by the geologic
and paleogeographic features of the Malesian region which spans two large
partly submerged continental shelves, the Asiatic shelf (Sunda Shelf linking
the Malay Peninsula with the islands of Sumatra, Java, Borneo and Palawan)
and the Australasian shelf (Sahul Shelf linking the Aru islands and New
Guinea with Australia). During the last glaciation period (c.22,500–11,000 bp)
the shelves were regions of land uncovered by the lowering of sea level, and
present day peninsular Malaysia, Sumatra and Borneo were connected by
Taxonomy and Systematics 29
land bridges during the late period of maximal sea lowering. During the cool
periods of glacial maxima, the Malesian forest was reduced in extent, but
there is no evidence that it was reduced to isolated island forests. The Sunda-
land and Papuasian rainforest blocks are therefore comparable to refugia in
terms of species richness and the high degree of endemism (Whitmore, 1981).
Mangifera has undergone major species development in west Malesia, which
has remained relatively stable over a long period of time. The current vegeta-
tion of west Malesia probably differs very little from that at the end of the
Tertiary (van Steenis, 1950). A lower number of Mangifera species is found in
Java and the Philippines, regions less often connected with Asia during the
Pleistocene.
Only three species occur in New Guinea, which is largely covered with
rainforest. These include M. minor, M. mucronulata and the widely distributed
M. gedebe. Mangifera foetida also occurs, but may have been introduced. Mangifera
minor occurs from Celebes and the Philippines to the Solomon Islands; M.
mucronulata is found in the Moluccas, New Guinea and the Solomon Islands.
The distribution of these species suggests a late immigration of a Laurasian
genus from Sundaland via the Philippines, Sulawesi and the Moluccas into
New Guinea, which is supported by the geological history of the region. No
Mangifera species have ever been recorded from northern Australia.
Very few species are found in peninsular India and Sikkim. From present-
day distribution, there is little evidence of migration of species into the sub-
continent of India after its collision with Eurasia in the middle Eocene
(Audley-Charles et al., 1981). According to Mehrotra et al. (1998), fossil leaves
described as Eomangiferophyllum damalgiriensis Mehr. from the Upper Palaeo-
cene in north-eastern India are an analogue of the modern genus Mangifera.
Mangifera sylvatica occurs along the northern limit of the range of
Mangifera, with more or less discontinuity, from Sikkim to northern Thailand
and to the southern part of Yunnan, where it is reported in mountains up to
1900 m above sea level (Anonymous, 1980). The few species that grow in
southern China are very poorly known: M. austro-yunnanensis from western
Yunnan, M. persiciformis from south-eastern Yunnan and southern Guizhou
at latitudes up to 26°N and M. hiemalis, the ‘winter mango’ from Guangxi
near the northern border Vietnam. In the revised Flora of China (Min and Bar-
fod, 2008), M. austro-yunnanensis is considered to be conspecific with M.
indica, M. hiemalis is treated as a synonym of M. persiciformis, and M. laurina
is recorded from the lowland forests of south Yunnan.
Subgenera and section distribution
The species distribution is especially meaningful when the ranges of the spe-
cies of each subgenus and section are considered separately.
Subgenus Limus
All species of the subgenus Limus are restricted to the Malesian area (M. foetida
and M. macrocarpa occurring in peninsular Thailand), whereas all the species
J.M. Bompard 30
with five fertile stamens, considered the most primitive condition, are con-
fined to west Malesia (M. decandra in Sumatra and Borneo; M. lagenifera in the
two latter areas and in peninsular Malaysia). Only M. caesia, M. foetida and
closely related M. leschenaultii occur in east Malesia.
Subgenus Mangifera
In the subgenus Mangifera, M. gedebe is the only species belonging to the sec-
tion Marchandora, and has the widest range within the genus, extending from
Myanmar through Malesia to New Guinea and Bougainville Island. Section
Euantherae is centred in the region from Myanmar to Vietnam. Mangifera
pentandra is only known from peninsular Malaysia, the Anambas Islands and
Borneo. Section Rawa is mainly in western Malaysia and shows notable
diversification in the swamps and peripheral uplands in the south of penin-
sular Malaysia, east central Sumatra (notably the Riau province) and west
Borneo. During the glacial period this area, termed the ‘Riouw pocket’ (Cor-
ner, 1978), formed a vast plain connecting the Malay Peninsula, Sumatra and
Borneo, and is believed to have been filled with swamps. Mangifera merrillii
is an endemic of the Philippines, M. minutifolia is an endemic of Vietnam, M.
andamanica and M. nicobarica are endemics of the Andamans and Nicobar
Islands. None of the species of section Mangifera occurring in mainland
South-east Asia, north of the isthmus of Kra, are found in eastern Malesia;
however, it would be interesting to assess the genetic relatedness of M. syl-
vatica and M. minor, and also M. laurina, which may prove to be phylogeneti-
cally very closely related.
2.5 Interspecific Molecular Characterization
Molecular biology techniques now make it possible to assess genetic related-
ness in a more precise way. Published data support some of the groupings based
on anatomical characters (Kostermans and Bompard, 1993) but not entirely.
RAPD (random amplification of polymorphic DNA) markers were first
used in mango by Schnell and Knight (1993) and Schnell et al. (1995). Nine
Mangifera species were analysed and compared to the traditional taxonomic
groupings. The unweighted pair group method of arithmetic averages
(UPGMA) cluster analysis for the subgenus Limus was not supportive of the
separation between sections Perennes and Deciduae, which, admittedly, has a
weak taxonomic basis. It confirmed the relatedness between M. foetida and
M. pajang. The UPGMA cluster analysis of the subgenus Mangifera supported
the current taxonomy based on flower morphology. It showed the related-
ness between M. quadrifida and M. torquenda (both placed in the group of
species with tetramerous flowers), but also with M. casturi, although the lat-
ter species has tetra- and pentamerous flowers. One of the most significant
results was the evidence for the existence of interspecific hybridization within
the studied species of the section Mangifera (see also Yonemori et al., 2002).
Phylogenetic relationships among 14 Mangifera species of Thailand were
analysed by comparing amplified fragment length polymorphism (AFLP)
Taxonomy and Systematics 31
markers (Eiadthong et al., 2000b), and by comparing sequences of the inter-
nal transcribed spacer (ITS) region of nuclear ribosomal DNA (nrDNA)
(Yonemori et al., 2002). They demonstrated that the common mango was
closely related to M. laurina, M. sylvatica and M. oblongifolia of subgenus
Mangifera to which M. indica belongs. A close relationship between M. indica
and M. sylvatica has been corroborated by Nishiyama et al. (2006), who com-
pared signal intensity of genomic in situ hybridization (GISH) on somatic
metaphase chromosomes of M. indica, using labelled DNA of eight wild
Mangifera species.
Furthermore, Eiadthong et al. (2000b) and Yonemori et al. (2002) have
demonstrated that M. odorata, M. foetida and M. macrocarpa (of subgenus
Limus) were related to M. indica. It is not surprising in the case of M. odorata
whose hybrid origin (M. foetida × M. indica) has now been established, but
this calls into question the position of the section Perennes.
Results of molecular studies do not permit a comprehensive view of the
phylogenetic relationships among the genera. So far, they are rather support-
ive of the groupings based on phenotype within the subgenus Mangifera
(notably sections Rawa and Euantherae), but not for the subgenus Limus which
will need to be redescribed, and likely restricted to the group of species
related to M. caesia (M. kemanga, M. lagenifera, M. superba and possibly M.
decandra). More studies will be needed to infer phylogenetic relationships
within the section Mangifera. Keeping in mind the frequent misidentifications
in collections and botanic gardens, herbarium specimens of studied material
must be deposited in the national herbaria so that its taxonomic position can
be ascertained in case of doubt.
2.6 Region of Origin of the Genus
Based on morphological, phytogeographical and fossil evidence, Mukherjee
(1953) argued that:
although the highest number of species of both sections is concentrated in the
Malay Peninsula [19 were then recorded], the centre of origin of the genus
cannot be restricted to that area alone, as both the phylogenetically older
species, i.e. with pentacyclic flowers (M. duperreana, now reduced to M.
caloneura, and M. lagenifera), occur in Siam and Indochina, and the former is
absent from Malay Peninsula.
He concluded that the genus had its origin somewhere in the Myanmar–
Thailand–Indochina area or in the Malayan area. Careful identification of the
greatest part of herbarium materials available has allowed a more accurate
delimitation of the distribution ranges of the Mangifera species, notably of the
subgenus Limus, and has revealed, among other things, that M. lagenifera
does not occur north of Kra isthmus contrary to Mukherjee’s assertions. Fur-
thermore, the ten-stamen species, M. decandra, which was described by Ding
Hou in 1972 and hence was unknown to Mukherjee, is confined to Borneo
and Sumatra, and to date has not been recorded from peninsular Malaysia.
J.M. Bompard 32
Without overemphasizing the present great species diversity of subgen-
era in the Malay Peninsula, Borneo and Sumatra, the available evidence
points to a Sundaic origin for the genus. This, however, must not minimize
the particular importance of the region stretching from Myanmar to Indo-
china as another centre of diversification, as attested by the range of species
belonging to the section Euantherae. Unfortunately, many of the species of
this region remain poorly known, and it can be expected that plant collecting
in the region will yield interesting new findings. The speciation that occurred
in this region with a likely radiation centre today traced by the range of the
section Euantherae, is of special significance as it has given rise to the com-
mon mango.
2.7 Origin of the Common Mango
The common mango apparently originated in regions on the western border
of the secondary centre of diversification mentioned above. Truly wild mango
trees have been recorded in Bangladesh (Chittagong Hill tract, c.23°N), north-
eastern India (‘undoubtedly indigenous in the evergreen tracts of valley of
Assam’ according to Kanjilal et al., 1937), and in Myanmar where it was
reported as ‘not unfrequent in the tropical and lower mixed forests all over
Burma from Arracan and Pegu down to Tenasserim’ (Kurz, 1877). It would
be desirable to assess its affinity with the species of the section Euantherae, as
well as with species of other sections of the subgenus Mangifera that occur in
the same area and region. It is also believed to be wild ‘in the sub-Himalayan
tract, in deep gorges of the Baraitch and Gonda hills in Oudh, and the outer
hills in Kamaon and Garhwal’ (Brandis, 1874). The common mango has been
grown and disseminated for such a long time in India that semi-wild trees
can be found in the forests throughout the subcontinent. The fruits of wild
trees are said to be small and of poor quality. Watt (1891) mentioned two so-
called ‘almost unaltered wild varieties’ existed under cultivation in Tirhoot,
‘one originating from Kangra, a very variable one, and the other from Sikkim
which was evidently the progenitor of the varieties cultivated in Malda’.
The common mango in South-east Asia
The Linnean binomial (Mangifera indica) indicates in this instance the place
where the common mango was selected and improved, and not necessarily its
place of its origin. It has been traditionally accepted that mango was domesti-
cated several millennia ago in India (see Mukherjee and Litz, Chapter 1, this
volume); however, it cannot be excluded that domestication occurred inde-
pendently in several areas, possibly in the south-western and south-eastern
regions of its centre of origin, or later differentiated in those two regions. This
hypothesis would account for the differences that exist between the local
polyembryonic cultivars of Myanmar, Thailand, Indochina and Indonesia,
and the monoembryonic Indian cultivars. Note that polyembryony occurs
Taxonomy and Systematics 33
also in the cultivated M. casturi, M. laurina and M. odorata. Aron et al. (1998)
have demonstrated that polyembryony in mango is under the control of a
single dominant gene.
According to Juliano (1937), Bijhouwer suggested that there were two
main centres of domestication of mango, ‘one in India with monoembryonic
mangoes, the other in the Saigon area, Indonesia and the Philippines with
polyembryonic mangoes’. The ‘Saigon’ area must in fact be extended to
southern Vietnam, other parts of Indochina, Thailand and Myanmar, which
were recognized by Valmayor (1962) as homes of polyembryonic mangoes.
Notwithstanding, the origin of polyembryonic mangoes is probably better
placed in Myanmar, and possibly the eastern part of Assam. According to
Brandis (1874), ‘in Burma, the mango is not generally grafted, and seeds of a
good kind, as a rule, produce fruit of a similar description’. There are only a
few polyembryonic mango cultivars in India. They are restricted to the south-
western coastal region, and geographically isolated from the polyembryonic
mangoes of Myanmar and South-east Asia. Analysis of genetic relatedness
using RAPD markers among polyembryonic and monoembryonic cultivars
grown in the west coast of southern India suggest that the polyembryonic
types are unlikely to have originated from India and might have been intro-
duced from South-east Asia (Ravishankar et al., 2004).
Indian Buddhist monks might have introduced the common polyembry-
onic mango to South-east Asia, first along land trade routes through Myan-
mar, where they might have found better races, and from there into insular
South-east Asia. It is well established that some local names of the common
mango currently used in parts of Indonesia are of Sanskrit origin (‘ampelam’
and its cognates), and are sometimes used to designate M. laurina, which is a
truly native species. Vernacular names do not always travel with a plant, and
even if they did so in the case of the common mango, it is very unlikely that
these introductions were the first ones and that they came obligatorily from
India. In the absence of a comprehensive classification of the innumerable
South-east Asian cultivated forms of the common and wild mangoes, includ-
ing the countless primitive races, we have to rely on linguistics and the rich
history and prehistory of this region.
Vernacular names
The different local names of the common mango in Indonesia (‘pauh’, ‘ampe-
lam’ and its variants, and ‘mangga’) bear evidence of a long history of con-
tacts with mainland Asia and India, and point to possible introduction at
different times from different places. In some parts of Indonesia, the vernacu-
lar names ‘paoh’ or ‘pauh’ refer either to primitive races of the common
mango, or to native species, as a rule the ones most closely resembling the
common mango, for example: ‘pauh asal’ (= native mango) for M. pentandra
in peninsular Malaysia; ‘pahohutan‘ or ‘pahutan’ (= forest mango) for M.
altissima in the Philippines; and ‘pao pong’ (= forest mango) for M. minor in
Flores, Lesser Sunda Islands. ‘Pau’ is a word belonging to Austronesian lan-
guages, nowadays spoken over a very wide area from Madagascar to the
Easter Islands by people who originate from mainland Asia. These languages
J.M. Bompard 34
are still spoken by certain minority populations in Vietnam, Cambodia and
the Mergui Archipelago off the coast of Myanmar (Bellwood et al., 1995). In
Cambodia, which was occupied by the Chams from about the 3rd to the 15th
century ad, ‘pa:uh’ is a Chamic word. ‘Sva:y’, used by the Khmers (as in
‘sva:y srok’ meaning mango of the village (M. indica), and ‘sva:y prey’, wild
mango (i.e. M. caloneura) as attested in pre-Angkorian Khmer inscriptions
dating from the 6th to the 8th century ad (Pou and Martin, 1981)) is of Austro-
Asiatic origin. ‘Sva:y’ has cognates in south Vietnam (‘xoay’) and in Asian
languages spoken by aboriginal people in peninsular Malaysia. ‘Wai’, another
cognate, is a vernacular name of M. minor in several parts of New Guinea.
Pawley and Ross (1995) proposed ‘wai’ and ‘pau(q)’ as the reconstructed
Proto-Oceanic terms referring, respectively, to a generic name for mango,
and a species that is probably M. indica.
Nowadays, these two words are generic terms for mango fruits that
rather closely resemble M. indica. In the same way, ‘thayet’ which is the com-
mon vernacular name referring to M. indica in Myanmar (‘sinnin thayet’ and
‘taw-thayet’ for M. caloneura and M. sylvatica, respectively), or ‘mamuang’ in
Thai languages are probably generic names.
Obviously, linguistic evidence alone provided by these vernacular names
is not sufficient to prove the time and place of an introduction. None the less,
it is significant that in mainland South-east Asia none of the vernacular names
of the common mango exhibits signs of an Indian influence, moreover, cog-
nates of these names are also applied to primitive races in some parts of
insular South-east Asia.
Evidence of early trade in South-east Asia
The history of plant domestication in mainland South-east Asia has undoubt-
edly involved introduction of plants by people migrating from the mainland
into insular South-east Asia. In more recent times, there is evidence of con-
tacts and sea trade since at least the first centuries ad between mainland and
insular South-east Asia to indicate that there have been numerous opportuni-
ties for introduction of the common mango from different places at different
times prior to the 4th century (before the Indianization of early South-east
Asian states) into present-day Malaysia and Indonesia.
Recent studies based on archaeological evidence stress the long unrecog-
nized importance of South-east Asian trade (emanating from South-east
Asia) between ports established along the Java Sea, those of mainland Asia,
and India, back to the 1st century ad, and possibly earlier (Walker and San-
toso, 1984). Trade routes connected the developing population centres of the
mainland, such as the earliest known South-east Asian political entity, Funan,
an advanced agrarian society located on the southern Vietnam coast, which
became influenced by the Indians and reached the zenith of its commercial
prosperity in the middle of the 3rd century (Hall, 1985). Increasingly, king-
doms organized according to the Indian concept of royalty were established
in the Indonesian archipelago, for example Kutai in East Kalimantan (4th
century) and Central Java (8th to 9th century), the latter being famous for the
Buddhist temple at Borobudur, where sculptures depict the mango tree.
Taxonomy and Systematics 35
It is highly probable that the eventual introductions of superior cultivars
of polyembryonic mangoes from the south-west coast of India, ‘between the
6th and 14th century, the height of classical South-east Asian civilization and
also the golden age of early south Indian civilization’ (Hall, 1985), were not
the first ones.
During the 16th and 17th centuries, the Portuguese and Spaniards con-
tributed to the widest distribution of superior varieties in the archipelago, espe-
cially to the east. The name mango itself derives from the Tamil ‘man-kay’ or
‘man-ga’ (see Mukherjee and Litz, Chapter 1, this volume), which the Portu-
guese adapted as ‘manga’ and ‘mangueira’ when they colonized west India.
Superior Philippine cultivars originated through introduction of culti-
vars from Indonesia, for example ‘Dodol’ into Mindanao, and from Indo-
china, for example ‘Carabao’ and ‘Pico’ in Luzon, the Visayas and northern
Mindanao (Wester, 1920; Bondad et al., 1984). However, these introductions
dating from the first half of the 17th century were also preceded by the intro-
duction of primitive races of the common mango as well as other species into
the Sulu Archipelago and Mindanao through contacts with north Borneo, as
attested by their local names quoted by Wester (1920), that is mampalam
(M. indica, and possibly also M. laurina), baonoh (M. caesia) and wannih (M.
odorata).
The South-east Asian M. indica germplasm includes many races that defy
classification. Natural cross-pollination has undoubtedly occurred with
native species, such as M. laurina, which was also brought into cultivation in
several areas before the introduction of M. indica.
2.8 Conclusion
Potential contribution of wild species to mango cultivation
To date, the improvement and breeding of common mango has depended on
the use of genetic variability within a single species, M. indica. Mukherjee
(1957) observed that ‘similarity in chromosome number and pollen morphol-
ogy in different species suggests close compatibility during hybridization
and stock-scion relationship if other species are used as stock for the com-
mon mango’. Biotechnology opens new perspectives for mango improve-
ment (Litz, 2004). As noted by Litz et al. (Chapter 18, this volume), the
transformation of mango with genes from other species could address a
number of plant breeding objectives.
Source of rootstock
Grafting experiments between M. indica and other species are reported in the
literature, for example budding of M. indica on M. foetida and M. odorata in
Java (Ochse and Bakhuizen, 1931), M. odorata on M. indica in the Philippines
(Wester, 1920), and M. indica on M. zeylanica in Sri Lanka (Gunaratman, 1946).
J.M. Bompard 36
Mangifera indica ‘Madu’ in Java, and M. laurina in Sabah have been used as
rootstocks for M. casturi. Trials of grafted M. caesia on M. indica (Wester, 1920)
and M. indica on M. kemanga or M. caesia (Ochse and Bakhuizen, 1931) were
unsuccessful, as these two species have distinct bark features and only remote
affinity with the common mango. Better compatibility can be expected using
species more closely related to the common mango within the subgenus
Mangifera. In West Kalimantan, M. laurina is occasionally used as a rootstock
for the common mango on periodically inundated riverbanks. It has been
tried as a rootstock by the Department of Agriculture in Sabah (Lamb, 1987).
Campbell (2004) reported that M. casturi, M. griffithii, M. laurina, M. odorata,
M. pentandra and M. zeylanica grafted on M. indica had a high percentage of
success.
Several species that can grow in permanently inundated areas (i.e. M.
gedebe, M. quadrifida, M. griffithii and other species of the section Rawa) repre-
sent a potential source of rootstock for the development of mango cultivation
on poorly drained soils or in areas liable to prolonged flood. Other species
may be a source of dwarfing rootstocks.
Hybridization
From our observations in Borneo, natural interspecific hybridization involv-
ing various cultivated Mangifera species can occasionally occur. Suspected
hybrids were observed between wild M. gedebe and cultivated M. laurina in
the lakes area along the Mahakam River in East Kalimantan, where impor-
tant populations of M. gedebe occur; between cultivated M. foetida and M. pajang,
two species showing close affinity, in different areas of Kalimantan where
both species are grown together; and between closely related M. caesia and
M. kemanga in cultivation. A hybrid origin has been suggested for M. odorata
(M. indica × M. foetida), which is unknown in the wild (Ding Hou, 1978a).
Based on AFLP analysis, Teo et al. (2002) and Kiew et al. (2003) have con-
firmed that M. odorata is a hybrid between M. foetida and M. indica. The index
of similarity showed that M. odorata is closer to M. foetida (76% similarity)
than it is to M. indica (66%). Yamanaka et al. (2006) showed a high genetic
similarity among 11 landraces of M. odorata from the Malaysian Agricultural
Research and Development Institute (MARDI) gene bank. Higher variability
can be expected from Sumatra and Java samples.
Existing information about experimental interspecific hybridization is
scarce. According to Mukherjee et al. (1968), successful crosses between
M. odorata and M. zeylanica were made in India.
Potential of wild species
There is little doubt that wild mangoes are potentially valuable in breeding
programmes. Some species have important horticultural implications as they
demonstrate many desirable characteristics (Bompard, 1993). Fairchild (1948)
Taxonomy and Systematics 37
noted that crosses between the common mango and related five-stamen spe-
cies of the section Euantherae might produce hybrids with better pollinating
quality. Mangifera pentandra, which is grown in peninsular Malaysia and
Sabah, is a prolific bearer, due to its high proportion of hermaphrodite to
male flowers.
Stress resistance
In the Malesian rainforests, wild mangoes thrive well under an ever-humid
climate, without a prolonged dry season, i.e. is in areas with an annual rain-
fall > 4000 mm and no monthly mean < 100 mm and where the common
mango cannot be grown satisfactorily. Species, occurring in subtropical areas,
including primitive races of the common mango, or in high altitude tropical
forests, should be evaluated for cold tolerance, opening up the possibilities
for mango production in subtropical and Mediterranean areas. Mangifera lau-
rina and other species related to the common mango that grow in the rainfor-
est (e.g. M. minor in New Guinea) are apparently immune to anthracnose.
Sharma and Choudhury (1976) also observed that trees of an unknown wild
race found in the Tripura State (north-eastern India) were free from mango
malformation.
Potential new fruits
Extensive, yet largely unrecorded variability also exists among the non-indica
species under cultivation. Sadly, this gene pool is barely represented in exist-
ing collections, and is rapidly vanishing. An increasing number of horticul-
turists are demonstrating a keen interest in the wild relatives of the mango.
It is hoped that local peoples who have contributed to the recognition and
maintenance of these species can benefit from future innovative mango
breeding.
Since early times, local peoples have planted seeds collected from trees
that were observed to produce better quality fruits in the forests around their
settlements. In areas now completely devoid of lowland primary forest, espe-
cially in Sumatra and Borneo, the only wild relatives still found are those
which have been integrated into indigenous agroforests which represent
gene banks for an amazing diversity of fruit trees. A tenuous but constant
selection pressure over many centuries has resulted in improved selections
of several species. Today, some of these selections hold economic importance
for their intrinsic characteristics. In Malesia, forms of M. odorata and M. foetida
with sweeter and less fibrous flesh have been identified. The ‘wani’, a form of
M. caesia from Bali and Borneo, has green-skinned fruit with milky white soft
flesh and a sweet taste quite different from the fruit of common forms of M.
caesia. In addition, there are many interesting selections of M. casturi, M. grif-
fithii and M. torquenda.
Further improvement of these wild mangoes is especially desirable
owing to their local economic importance in the wet tropical regions. Use of
vegetative propagation methods must be encouraged. With proper selection,
there is every reason to believe that other Mangifera species can become valu-
able commercial fruits.
J.M. Bompard 38
Acknowledgement
Thanks are due to Dr Dawn Frame who assisted in correcting the text.
Note
1
Surveys were carried out in Kalimantan in cooperation with the Indonesian Institute of
Science (LIPI) and the Indonesian Commission on Germplasm, and in Malaysia with the
Forest Research Institute of Malaysia (FRIM).
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© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
42 (ed. R.E. Litz)
3 Important Mango Cultivars and
their Descriptors
R.J. Knight, Jr,¹ R.J. Campbell² and I. Maguire¹
1
University of Florida, Florida, USA
²Fairchild Tropical Botanic Garden, Florida, USA
3.1 Introduction 43
3.2 Criteria for Cultivar Description 44
3.3 Mango Cultivars 45
‘Alfa’ (Brazil) 45
‘Alphonso’ (India) 45
‘Amelie’ (West Africa) 46
‘Arumanis’ (Indonesia) 46
‘Ataulfo’ (Mexico) 46
‘B74’ (‘Calypso’™) (Australia) 47
‘Banganpalli’ (India) 47
‘Beta’ (Brazil) 47
‘Bombay Green’ (India) 47
‘Cambodiana’ (Vietnam) 48
‘Carabao’ (Philippines) 48
‘Chausa’ (India) 48
‘Cogshall’ (Florida, USA) 49
‘Coração de Boi’ (Brazil) 49
‘Dasheheri’ (India) 49
‘Espada’ (Brazil) 49
‘Ewais’ (Egypt) 50
‘Excellent Succari’ (Egypt) 50
‘Extrema’ (Brazil) 50
‘Fajri’ (India) 50
‘Fernandin’ (India) 51
‘Genovea’ (Egypt) 51
‘Glenn’ (Florida, USA) 51
‘Golek’ (Indonesia) 51
‘Haden’ (Florida, USA) 52
‘Himsagar’ (India) 52
‘Hindi Besennara’ (Egypt) 52
‘Hindi Khassa’ (Egypt) 52
Mango Cultivars and Descriptors 43
‘Irwin’ (Florida, USA) 53
‘Julie’ (West Indies) 53
‘Keitt’ (Florida, USA) 53
‘Kensington’ (Australia) 54
‘Kent’ (Florida, USA) 54
‘Khanefy’ (Egypt) 54
‘Kyo Savoy’ (Thailand) 55
‘Langra’ (India) 55
‘Mabrouka’ (Egypt) 55
‘Madame Francis’ (Haiti) 55
‘Mallika’ (India) 56
‘Manila’ (Mexico) 56
‘Manzanillo’ (Mexico) 56
‘Mesk’ (Egypt) 56
‘Mulgoa’ (India to Florida, USA) 57
‘Nabeel’ (Egypt) 57
‘Nam Doc Mai’ (Thailand) 57
‘Neelum’ (India) 58
‘Nuwun Chan’ (Thailand) 58
‘Okrung’ (Thailand) 58
‘Osteen’ (Florida, USA) 59
‘Pairi’ (India) 59
‘Palmer’ (Florida, USA) 59
‘Rosa’ (Brazil) 59
‘Sensation’ (Florida, USA) 60
‘Suvarnarekha’ (India) 60
‘Tahar’ (Israel) 60
‘Taimour’ (Egypt) 61
‘Tommy Atkins’ (Florida, USA) 61
‘Totapuri’ (India) 61
‘Turpentine’ (West Indies) 62
‘Vallenato’ (Colombia) 62
‘Van Dyke’ (Florida, USA) 62
‘White Succari’ (Egypt) 62
‘Zebda’ (Egypt) 63
3.4 Conclusion 63
3.1 Introduction
The mango (Mangifera indica L.) has traditionally been grown in an area that
extends southwards and eastwards from India through Myanmar and Vietnam
to Indonesia. It probably is not indigenous to the Philippines, where it has long
been cultivated (Valmayor, 1962), but Mangifera species are endemic there (Bon-
dad, 1982). This crop is best adapted to a warm tropical monsoon climate, with a
pronounced dry season followed by rains. Fruit of the best quality is usually
produced in such areas, but specific races are known to fruit in humid regions.
For example, some Mangifera species bear dependably on the island of Borneo,
where most standard cultivars do not mature normal crops of fruit. Numerous
Mangifera species closely related to the common mango are indigenous to
R.J. Knight et al. 44
Borneo and nearby parts of Malaysia and Indonesia, and this region is the prob-
able centre of diversity for this genus (see Bompard, Chapter 2 this volume).
Most crops long cultivated over an extended time and area show consider-
able diversity, reflecting different selection criteria in different regions of cul-
ture as well as genetic responses to varied environmental influences. Certainly
this is true of the mango. Indian cultivars differ markedly from those grown
in South-east Asian countries and in Egypt. An additional factor that has
promoted genetic diversity in the group recently has been the widespread
introduction of this crop into new areas of cultivation, many in the western
hemisphere, over the last 500 years. In this manner, genetically diverse germ-
plasm has been brought from widely dispersed areas of the original range of
the species and grown in mixed plantings where, through the cross-pollination
natural to the species, new genetic combinations have been made and selected
under many varying conditions of microclimate. In Florida, since the late 18th
century, enough such importations and genetic recombinations have occurred
to qualify the southern part of this state as a secondary centre of diversity for
the crop. A new group of mango clones designated the Florida cultivars has
been exported to Brazil, Israel, Australia and other places where the process of
increasing diversity under new and varying cultural and environmental con-
ditions continues (Knight and Schnell, 1994; Schnell et al., 2006).
Some Florida cultivars, most notably ‘Haden’, have been important in aid-
ing the establishment of a modern mango industry in other parts of the world
(Knight and Schnell, 1994), and the phenomenon first observed in Florida is now
occurring elsewhere; we are presented with the prospect of the importation of
cultivars of outstanding merit from their countries of origin to be grown, first
experimentally and then commercially, in new regions. For this reason it is
important to become familiar with the characteristics of a group of cultivars that
currently are known in the commerce and/or horticulture of one or more coun-
tries, and that may have potential for expanded culture or use in breeding.
3.2 Criteria for Cultivar Description
In the recent past, efforts to assemble lists of mango descriptors produced
two publications that cover the subject (Mukherjee, 1985; IBPGR (now Inter-
national Plant Genetic Resources Institute, IPGRI), 1989) and provide people
who manage collections with morphological criteria to identify cultivars. The
Descriptor List used by IPGRI documents passport data (identifying the acces-
sion and information recorded by collectors), characterization (recording char-
acters considered to be highly heritable which can easily be seen in the field
and are expressed in all environments) and preliminary evaluation, which records
a limited number of additional traits thought desirable by a consensus of users
of the crop. Plant data are important in preliminary evaluation, and include for
the tree, habit and height of the mature tree; for the leaf, shape, length and
width, and colour of the young leaf; and for the inflorescence, position, shape,
density of flowers, length, colour, hairiness, presence or absence of leafy bracts,
and percentage of flowers in an average inflorescence. (Some research indicates
Mango Cultivars and Descriptors 45
that both leafy bracts and number of perfect flowers are influenced by local
conditions and vary in their expression with differing environments.)
Additional plant data used in initial evaluation include those for the flower,
diameter in millimetres, type (pentamerous, tetramerous or both), nature of
disc (swollen, broader or narrower than ovary, reduced or absent) and number
of stamens; fruit, length, width and thickness, weight, shape, skin colour (which
may be compared with reference cultivars), skin thickness, skin texture, ratio
of pulp to skin and stone, texture of pulp, adherence of skin to pulp, fibre in
pulp and its quantity and length, and stem insertion; and stone, length, weight,
veins and pattern of venation, presence or absence of fibres and their length.
Additional plant data for leaves, inflorescence and fruit have been col-
lected and some of these, notably season (maturity period), productivity, eat-
ing quality and attractiveness are quite important. Unfortunately, from the
viewpoint of those who expect to apply these criteria outside the Indian sub-
continent, reference cultivars are for the most part Indian and many are not
readily available outside India. Other important characters that have been
evaluated or proposed for evaluation include susceptibility to stress (drought,
wind, flooding), susceptibility to specific diseases and pests, molecular
markers, cytological characters and identified genes. Because of the extreme
comprehensiveness of this list and the limited availability of many of the
proposed descriptor evaluations at this time, we have tried to utilize such
information as is available to make the comparison, identification and evalu-
ation of specific well-known cultivars a practical possibility.
3.3 Mango Cultivars
A list of mango cultivars that are of interest in areas other than their places of
origin, with descriptions intended to help differentiate them, follows (see
Plates 4–40). Spelling and name variants in some cases represent efforts to
transliterate from other orthographies to the Roman alphabet, and in others
reflect regional differences in usage.
‘Alfa’ (Brazil)
A monoembryonic cultivar developed by EMBRAPA Cerrado, Brazil, from
crossing ‘Mallika’ × ‘Van Dyke’. The tree is semi-dwarf in habit and high-
yielding, resistant to Oidium mangiferae and malformation, and moderately
resistant to anthracnose (Colletotrichum gloeosporioides); the fruit is large (435 g),
pink-red, firm, medium fibrous and of good quality (16% total soluble solids
(TSS), 0.23% acidity) (Pinto et al., 2004).
‘Alphonso’ (India)
Also known as ‘Appus’, ‘Badami’, ‘Gundu’, Haphus’, ‘Kagdi’, ‘Khader’ and
‘Khader Pasand’. The tree is moderately large, with broadly rounded, dense
R.J. Knight et al. 46
canopy; the fruit (Plate 4) is yellow, ovate-oblique, averaging 6 cm long by
5 cm broad, weighing 225–325 g (mean 226 g); the skin is thin; the flesh is
firm to soft, low in fibre, yellow, sweet with characteristic aroma and with a
very pleasant taste preferred by many who know this cultivar, bringing pre-
mium prices on Indian and international markets. Seed is monoembryonic in
a large, woody stone; the quality is excellent; ripening fruit in late to midsea-
son. Bearing is irregular, medium to heavy in India, but light and irregular in
Florida (Prasad, 1977; R.J. Knight, Jr, personal communication, 1995).
‘Amelie’ (West Africa)
Also known as ‘Gouverneur’ in the Caribbean. The tree is tall with a rounded,
dense canopy; the fruit is green to orange-yellow with the advance of the season,
rounded, 10–15 cm long by approximately 10 cm broad by approximately 7.8 cm
thick and weighing 300–600 g (average 360 g). The skin is thick and separated
with difficulty; the flesh is soft, juicy, melting, without fibre, a deep orange
colour, sweet and perfumed, free from turpentine, and provides the best of
mango tastes. Seed is monoembryonic in a medium-sized, elongate, narrow
stone that adheres to the flesh, having a few short, pliable fibres that are not
objectionable; the quality is excellent; the season early. The fruit closely resem-
bles that of ‘Julie’. ‘Amelie’ is exported to France, along with ‘Kent’, from
Burkina Faso, Ivory Coast and Mali. ‘Amelie’ is increasing in popularity on the
French market, chiefly in Paris and the surrounding area. It brings lower prices
than cultivars with blushed fruit because the consumer is not always aware
when it is ripe (Naville, 1985, 1986; R. LePrette, personal communication, 1996).
‘Arumanis’ (Indonesia)
Also referred to as ‘Harumanis’. The tree is vigorous and tall with a slightly
open canopy. The fruit (Plate 5) is greenish yellow with large, light-yellow
dots, elongate oblong with rounded base, 11–14 cm long by 6.6–7.5 cm broad
by 4.75–6.5 cm thick, weighing 200–350 g. The skin is thick, tough and easily
separated, the flesh firm and juicy with little fibre, lemon yellow, sweet,
slightly insipid with a strong aroma, of poor to fair quality. Seed is polyem-
bryonic in a thick, woody stone; this cultivar ripens midseason and bears
regularly. Relatively easy to propagate by graftage, scionwood survives well;
widely planted in humid parts of the world where many better-quality culti-
vars fail to fruit (R.J. Knight, Jr, personal communication, 1995).
‘Ataulfo’ (Mexico)
A polyembryonic cultivar sold in North American markets under the name
‘Ataulfo’ and as ‘Champagne’™. Originated in Tapachula, Chiapas, Mexico
reportedly from seed brought from Costa Rica in about 1930. The tree is
Mango Cultivars and Descriptors 47
vigorous and upright, a mid-range producer with production averages of
10–20 t/ha possible. The tree is not highly adaptive to different climatic/
edaphic conditions. It is moderately resistant to anthracnose disease. The
fruit (Plate 6) is small (200–300 g), elongate, of good quality, sweet with slight
acidity, yellow, firm, standing shipping stress well, and ripens from early to
midseason (Campbell et al., 2002; Magallanes-Cedeño, 2004).
‘B74’ (‘Calypso’™) (Australia)
A monoembryonic cultivar that originated from the controlled cross of ‘Sensa-
tion’ × ‘Kensington’. The tree is upright, with low to moderate vigour and is
highly productive, with good tolerance of flower and fruit diseases; the fruit
(Plate 7) is moderately large (457.4 ± 38.1 g), ovate (10.12 ± 0.27 cm long by
9.13 ± 0.28 cm wide), fibre-free and firm, bright yellow overlaid with red blush,
with extended shelf life and potential for shipment to overseas markets; ripens
late in the season; patented (Whiley, 2001; Whiley and Hofman, 2006).
‘Banganpalli’ (India)
Also called ‘Beneshan’ and ‘Chappatai’. The tree is medium sized with a
rounded canopy; the fruit is primrose-yellow, ovate-oblique, large and the
skin smooth, thin and shiny, flesh firm to meaty with juice moderately abun-
dant, without fibre, maize-yellow, with pleasant aroma and sweet taste. Seed
is monoembryonic, in an oblong stone covered with sparse fibres; quality
good; ripens midseason and bears heavily (Singh, 1960).
‘Beta’ (Brazil)
A cultivar developed by EMBRAPA Cerrado, Brazil, from crossing ‘Amra-
pali’ × ‘Winters’ (M20222 United States Department of Agriculture (USDA)).
The tree is moderately vigorous and free of malformation, high-yielding but
irregular, moderately resistant to anthracnose and Oidium; the fruit is small
(310 g), yellow, firm with low fibre, of excellent quality (24.8% TSS, 0.16%
acidity) (Pinto et al., 2004).
‘Bombay Green’ (India)
Also called ‘Bhojpuri’, ‘Bombai’, ‘Hiralal Bombai’, ‘Kali Bombai’, ‘Laile Alipur’,
‘Malda’, ‘Sarauli’ and ‘Sheeri-Dhan’. The tree is tall and erect; the fruit (Plate
8) is apple green with ochre blush at the base and on some exposed parts,
dots abundant, with brown specks in the middle, ovate with beak almost
missing, medium sized, with tough, thick, non-adhering smooth skin; the
flesh is cadmium-orange, firm and juicy with scanty fibre just under the skin,
R.J. Knight et al. 48
very sweet with pleasant aroma, of very good quality; seed is monoembryonic
in a full, thick, medium-sized stone. This cultivar ripens early in the season
and is a medium bearer. ‘Bombay Yellow’ is said to be practically identical to
this cultivar but for a slight difference in fruit colour. The present ‘Bombay
Green’ is said to be a degenerate form of the original one (Singh, 1960). In
Jamaica it is sometimes called ‘Peter’, which suggests a confusion with ‘Pairi’,
but the Jamaican ‘Peter’ is without the bright red blush normal to ‘Pairi’.
‘Cambodiana’ (Vietnam)
Also known as ‘Xoai Voi’. The tree is moderately vigorous, with a dense,
rounded canopy; the fruit (Plate 9) is greenish yellow, unblushed with a few
small white dots, oblong to ovate, 9–11.5 cm long by 6.5–7.5 cm broad by
5–6 cm thick, weighing 220–340 g; the skin is thin, tender and adherent; the
flesh contains little fibre, is tender and melting, lemon yellow, sweet and
mildly subacid with a pleasant aroma; the seed is polyembryonic in a thick,
woody stone. Ripens early in the season. Brought to Florida in 1902, where it
gave rise to the ‘Saigon’ landrace (Campbell, 1992).
‘Carabao’ (Philippines)
The tree is vigorous, forming a large and dense canopy; the fruit (Plate 10) is
greenish to bright yellow, brushed with a few small green dots, long and
slender, with base rounded to slightly flattened, 11–13 cm long by 6.5–7 cm
broad by 6–6.5 cm thick, weighing 270–440 g; the skin is thick, medium tough
and easily separated; the flesh is without fibre, tender and melting, lemon
yellow, spicy and sweet with a mild aroma, of good to excellent quality; seed
is polyembryonic in a thin, papery stone. Ripens early in the season (Camp-
bell, 1992). This is a heavy bearer that may alternate; however, it can be
induced to fruit by potassium nitrate treatment in the tropics (Bondad and
Linsangan, 1979). It is highly resistant to bacterial black spot (Xanthomonas
campestris pv. mangiferaeindicae) in Queensland (Mayers et al., 1988). It was
introduced to Florida in 1909. ‘Carabao’ is important in commerce between
the Philippines and Japan and is increasingly imported into the USA.
‘Chausa’ (India)
Also called ‘Samar Bahisht Chausa’ and ’Khajari’. The tree is tall and spread-
ing; the fruit is canary yellow to raw sienna when fully ripe, with numerous
obscure medium-sized dots with minute specks inside them, oblong with
prominent beak, obtuse to rounded, medium sized; the skin is thin and some-
what adhering, pulp raw sienna, soft and juicy with scanty fine, long fibres
near the skin; the fruit is very sweet with a luscious, delightful aroma, of
excellent quality; seed is monoembryonic in a thick, medium-sized oblong
Mango Cultivars and Descriptors 49
stone with fine, short fibres all over the surface and a tuft of long fibres on
the ventral edge. Ripens late in the season and is a light bearer (Singh, 1960).
‘Cogshall’ (Florida, USA)
A monoembryonic cultivar that originated on Pine Island in Lee County. The
tree is relatively small, forming a rounded canopy, moderately susceptible to
anthracnose and consistently productive; the fruit is medium to large, aver-
aging about 350 g, yellow with a bright crimson blush, oblong (11–14 cm long
by 7.5–8.5 cm broad by 6.2–8 cm thick) of excellent quality, rich and sweet in
taste, with tender skin and soft flesh. Ripens early to midseason over about 4
weeks, a season longer than some cultivars. It is recommended for the home
garden, not commercial planting, in Florida but is now grown commercially
on Mauritius and marketed in France. Seedling of ‘Haden’ (Campbell and
Campbell, 1995; Schnell et al., 2006).
‘Coração de Boi’ (Brazil)
The tree is vigorous, precocious and productive; the fruit is greenish yellow and
intense red on the side exposed to the sun, cordiform, medium sized, pulp yel-
low and fibrous. The seed is polyembryonic. There are two seasons in São Paulo,
January–February and September–December. This is one of the best-known
commercial cultivars in São Paulo state (Sampaio, 1980; A.C. Pinto, personal
communication, 1996; L.C. Donadio, personal communication, 1996).
‘Dasheheri’ (India)
Also known as ‘Dasheri’ and ‘Aman Dusehri’. The tree is of medium height
and moderate vigour, spreading, with a rounded, medium-dense canopy; the
fruit is primrose to canary yellow with abundant light-yellow dots, oblong to
oblong-oblique with base rounded to obliquely rounded, medium sized, skin
smooth, medium thick, tough and non-adhering; the flesh is yellow, firm,
with almost no fibre, scanty juice and a delightful aroma, very sweet taste, of
excellent quality; seed is monoembryonic in a thick, medium-sized stone. Rip-
ens midseason and is heavy bearing; fruit keeps well (Singh, 1960).
‘Espada’ (Brazil)
The tree is tall and develops rapidly, with a dense canopy, very productive;
the fruit is intense green or greenish yellow, oblong-elongate with a concave
base, medium sized, with smooth, thick skin; the flesh has much fibre, is egg-
yellow, with a strong aroma of turpentine. The quality is considered good for
fresh consumption. The polyembryonic seed is in an oblong stone, covered
R.J. Knight et al. 50
with soft fibres and many nerves. There are two seasons per year in São
Paulo, January–February and November–December (Sampaio, 1980; A.C.
Pinto, personal communication, 1996).
‘Ewais’ (Egypt)
A polyembryonic cultivar of major commercial importance. The tree is vigor-
ous, the fruit small (275 g), yellow with no blush, with small, light-brown
slightly corky dots, ovate-oblong in shape (11.7 cm long by 7.2 cm wide by
6.3 cm thick), with adherent skin of intermediate thickness, relatively free of
disease; flesh orange, juicy but susceptible to jelly seed, with no objectionable
fibre, sweet and agreeable in taste, of very good quality. The stone is large
(38.5 g). Fruit ripens midseason (Knight and Sanford, 1998). In warm subtrop-
ics this cultivar has shown a tendency for flowering in the warm season, with
fruit ripening during the cool winter. It has good anthracnose tolerance.
‘Excellent Succari’ (Egypt)
A polyembryonic mango of minor commercial importance. The tree is vigor-
ous, ripening fruit in late midseason. The fruit is small (280 g), green with a
yellow overlay and small, yellow smooth dots, ovate-oblong in shape (11 cm
long by 7 cm wide by 6.4 cm thick), with non-adherent skin of intermediate
thickness quite free of surface disease; the flesh is orange, melting (without
jelly seed) and juicy with no objectionable fibres, a delightfully sweet taste
and excellent quality; stone large (36.6 g) (Knight and Sanford, 1998). It has
moderate to good anthracnose tolerance in the warm subtropics.
‘Extrema’ (Brazil)
The tree is upright, vigorous and productive. The fruit is yellow with green-
ish areas, ovate-reniform, weighing 350–400 g, with smooth and thin skin,
and yellow, watery flesh with almost no fibres with a moderately resinous,
agreeable taste. The quality is considered good for fresh consumption and
processing. The polyembryonic seed is in a large, fibrous stone. Ripens early
in the season (Sampaio, 1980; A.C. Pinto, personal communication, 1996).
‘Fajri’ (India)
Also spelled ‘Fazli’. The tree is of medium size and moderately vigorous,
with rounded, open canopy. The fruit is light chrome yellow with small,
dark-coloured fairly sparse dots, obliquely oval with base slightly rounded
and beak distinct to slightly prominent, large (averaging 14.3 cm long by
9.8 cm broad, weighing 500 g on average) with a medium-thick skin that is
Mango Cultivars and Descriptors 51
smooth with some inclination to be warty, and firm to soft, fibreless flesh of
a light cadmium yellow with a pleasant aroma and a sweet taste, having juice
that may be scanty to moderately abundant, of good to very good quality.
The seed is monoembryonic in a large, oblong stone that is covered with a
sparse, short and soft fibre. Ripens midseason to late (Gangolly et al., 1957; N.
Balasundaram, India, personal communication, 1990).
‘Fernandin’ (India)
The tree is moderately vigorous with a dense, rounded canopy; the fruit is
bright yellow with an attractive bright-red blush, ovate-oblique, averaging
12.2 cm long by 8.5 cm broad, weighing 450 g; the skin is rough and warty,
thick and adherent, flesh bright yellow, moderately to abundantly juicy,
thick, with no objectionable fibre, with delightful to piquant aroma and sweet
to very sweet, delicious taste, of superior quality; seed is monoembryonic;
season late (Gangolly et al., 1957; Singh, 1960).
‘Genovea’ (Egypt)
A polyembryonic cultivar of minor commercial importance. The fruit is small
(234.5 g), green with a yellow overlay and medium-sized smooth yellow
dots, ovate-oblong in shape (11 cm long by 6 cm wide by 5.6 cm thick), a thin
adherent skin relatively free of surface disease; flesh orange, firm (no jelly
seed) and juicy with no objectionable fibres, a sweet agreeable taste of ac-
ceptable quality; stone large (53 g) (Knight and Sanford, 1998).
‘Glenn’ (Florida, USA)
The tree is moderately vigorous, small to medium with dense, rounded can-
opy of compact growth; the fruit (Plate 11) is bright yellow with orange-red
blush, with numerous small yellow and white dots, oval to oblong with a
rounded base, 9.5–12.5 cm long by 7.5–8.5 cm broad by 7–8 cm thick, weigh-
ing 400–620 g; the skin is thin, tough and easily separated, flesh soft and juicy,
with little fibre, deep yellow, rich and spicy with a strong, pleasant aroma, of
excellent quality; seed is monoembryonic in a thick, woody stone. Ripens
early in the season and is a regular bearer. This is a seedling of ‘Haden’
(Campbell, 1992; Schnell et al., 2006).
‘Golek’ (Indonesia)
The tree is moderately vigorous with an upright, open canopy; the fruit
(Plate 12) is greenish yellow with an orange overlay and prominent white
dots, oblong with rounded base, 9.5–12.5 cm long by 6–8 cm broad by 5.5–
6.5 cm thick, weighing 200–365 g; the skin is thin, tough and easily separated;
R.J. Knight et al. 52
the flesh is soft and juicy with abundant fibre (not objectionable), deep yel-
low, sweet, insipid with a mild aroma, of poor to fair quality; the seed is
polyembryonic in large, woody stone with abundant fine fibre. Ripens mid-
season (R.J. Knight, Jr, personal communication, 1995).
‘Haden’ (Florida, USA)
The tree is vigorous, with a large, spreading canopy; the fruit (Plate 13) is
bright yellow with a deep crimson or red blush and numerous large yellow
dots, oval with a rounded base, 10.5–14 cm long by 9–10.5 cm broad by 8.5–
9.5 cm thick, weighing 510–680 g; the skin is thick, tough and adherent; the
flesh is firm and juicy with abundant fibre, deep yellow, rich and sweet with
a pleasant aroma, of good to excellent quality; the seed is monoembyonic in
a medium-thick woody stone. Ripens early to midseason and bearing is
sometimes irregular. This is a seedling of ‘Mulgoba’ × ‘Turpentine’ and is the
first of the Florida mango cultivars, introduced in 1910 and since grown in
many other countries. It is the seed parent of numerous other cultivars
(Campbell, 1992; Knight and Schnell, 1994; Schnell et al., 2006).
‘Himsagar’ (India)
The tree is vigorous, tall, with a dense, spreading canopy; the fruit (Plate 14)
is greenish yellow to bright yellow with no blush, with light-yellow dots,
ovate with a flattened base, 12–15 cm long by 8.5–9.5 cm broad by 7.5–8.5 cm
thick, weighing 465–585 g; the skin is thin, tough and easily separated; the
flesh is firm and juicy with no fibre, orange, rich and sweet with a mild aroma,
of good to excellent quality; the seed is monoembryonic in a thick, woody
stone. This is a late midseason cultivar that bears well (R.J. Knight, Jr, per-
sonal communication, 1995).
‘Hindi Besennara’ (Egypt)
A polyembronic cultivar of major commercial importance. The tree is of me-
dium vigour, ripening fruit early to midseason. The fruit (Plate 15) is of small
to medium size (319 g), green with orange overlay, with small white corky dots,
oblong-cylindrical in shape (15.4 cm long by 6.7 cm wide by 6.5 cm thick) with
thick, non-adherent skin relatively free of surface disease; the flesh is orange,
yielding and juicy with no objectionable fibres, pleasantly sweet in taste, of
very good quality; the stone is large (47.2 g) (Knight and Sanford, 1998).
‘Hindi Khassa’ (Egypt)
A polyembryonic cultivar of major commercial importance. The tree is vig-
orous, ripening fruit in late midseason. The fruit is of medium size (461 g),
Mango Cultivars and Descriptors 53
yellow with no blush, with intermediate-sized smooth, light-yellow dots,
oblong-cylindrical in shape (16 cm long by 6.6 cm wide by 6.9 cm thick), with
thick, adherent skin relatively free of surface disease; flesh is orange, firm
and juicy with no objectionable fibres, of mediocre taste and a quality not
suitable for export; stone is large (55 g) (Knight and Sanford, 1998).
‘Irwin’ (Florida, USA)
The tree is small to medium, moderately vigorous, with open canopy. The fruit
(Plate 16) is bright yellow with a crimson or bright red blush, numerous large
white dots, ovate with rounded base, 11.5–13 cm long by 8–9 cm broad by 6.5–
7.5 cm thick, weighing 340–450 g; the skin is medium-thick, tender and adherent;
the flesh is soft, tender, melting and juicy without fibre, lemon yellow, sweet and
mild with a pleasant aroma, of good quality; the seed is monoembryonic in a
thin, papery stone. The stone may be seedless following cool weather at flower-
ing time. This is an early, regular and heavy bearer. The fruit is usually soft with
a short postharvest life, but it is often exported from tropical America to Europe.
It is a seedling of ‘Lippens’ × ‘Haden’ (Campbell, 1992; Schnell et al., 2006).
‘Julie’ (West Indies)
Also called ‘St Julienne’. The tree is compact (dwarf), with a dense canopy;
the fruit (Plate 17) is greenish yellow with a light pink to maroon blush and
numerous small white dots, rounded with flattened apex, pronouncedly
compressed laterally, 7–9.5 cm long by 4–7.5 cm broad by 2–5.5 cm thick,
weighing 200–325 g with a thin, tender skin and soft, melting, juicy, orange
flesh with scanty fibre, of a rich, spicy flavour with a strong, pleasant aroma,
of good quality; seed is monoembryonic in a thin, papery stone. This cultivar
ripens midseason and is a regular producer of small crops. The fruit is often
severely infected with anthracnose disease, but its unique taste is preferred
by many West Indians, and it is exported to the London market (C.W. Campbell,
personal communication, 1996).
‘Keitt’ (Florida, USA)
The tree is medium sized, moderately vigorous, upright with open canopy;
the fruit (Plate 18) is greenish yellow, with a pink or red blush, numerous
small white or yellow dots, oval, with rounded base, 13–15 cm long by
9–11 cm broad by 8.5–10 cm thick, weighing 510–2000 g; the skin is thick,
tough and adherent; the flesh is firm and juicy, with little fibre, lemon yellow,
sweet and mild with a pleasant aroma, of good to excellent quality; the seed is
monoembryonic in a thick and woody stone. This cultivar ripens late in the
season. It is a seedling of ‘Brooks’. After ‘Tommy Atkins’ it is the most com-
mercially important cultivar in the export mango industry of the western
R.J. Knight et al. 54
hemisphere. It is resistant to anthracnose disease, packing and shipping stress
and is heavily productive (Campbell, 1992; Schnell et al., 2006). It is highly sus-
ceptible to bacterial black spot in Queensland (Mayers et al., 1988).
‘Kensington’ (Australia)
Also known as ‘Kensington Pride’ and ‘Bowen’. ‘Kensington’ has a large, vig-
orous tree with spreading canopy; the fruit (Plate 19) is yellow with an orange-
red blush on the shoulder, round ovate with a flattened base and a slight beak,
10.5–13 cm long by 8.5–9.6 cm broad by 7.5–8.5 cm thick, weighing 350–750 g;
the skin is thick, tender and adherent; the flesh is soft and juicy, with moderate
to little fibre, sweet with a characteristic flavour that makes it the most popu-
lar cultivar in Australian markets, of excellent quality; seed is polyembryonic
in a moderately thick, woody stone. This cultivar ripens midseason and it
bears well. It is unusually susceptible locally, in Florida, to damage by red-
banded thrips (Selenothrips rubricinctus (Giard.)), and may be killed by this pest
without adequate countermeasures (R.J. Campbell, personal communication,
1994; R.J. Knight, Jr, personal communication, 1995). It is moderately suscep-
tible to anthracnose and bacterial spot (Mayers et al., 1988).
‘Kent’ (Florida, USA)
The tree is large and vigorous with a dense, upright canopy; the fruit (Plate
20) is greenish yellow with a red or crimson blush, numerous small yellow
dots, oval, with rounded base, 11–13 cm long by 9.5–11 cm broad by 9.9.5 cm
thick, weighing 600–750 g; the skin is thick, tough and adherent; the flesh is
firm, tender, melting and juicy with little fibre, deep yellow to orange-yellow,
sweet with a rich flavour and pleasant aroma, of excellent quality; the seed
is monoembryonic in a thick, woody stone. Fruit ripens late midseason to
late and bearing may be alternate. It is a seedling of ‘Haden’ × ‘Brooks’, which
is a seedling of ‘Totapuri’ (‘Sandersha’) (Schnell et al., 2006). ‘Kent’ is not
commonly commercial in Florida because it is prone to storage disease, but
is a successful commercial cultivar in drier parts of Mexico, Central and
South America and West Africa (Campbell, 1992). It is highly susceptible to
bacterial black spot in Queensland (Mayers et al., 1988).
‘Khanefy’ (Egypt)
A cultivar of minor commercial importance. The fruit is large (475 g), green
with a yellow overlay and large, brown, smooth dots, ovate in shape (10.7 cm
long by 8.3 cm wide by 8.6 cm thick), with an adherent skin quite free of sur-
face disease; the flesh is yellow, often with jelly seed, juicy, with no objection-
able fibres and a bland flavour unacceptable to many Western palates. The
stone is moderately large (53 g) (Knight and Sanford, 1998).
Mango Cultivars and Descriptors 55
‘Kyo Savoy’ (Thailand)
The tree is large, vigorous, with an open canopy made up of long branches;
the fruit is green when harvested (before the ripening process begins) turn-
ing to greenish yellow, oblong, 11.5–12.5 cm long by 5.5–6.5 cm broad by
5–6 cm thick, weighing 230–340 g; the skin is thin, tender and adherent; the
flesh is medium firm, tender and not very juicy with no fibre, pale yellow,
very sweet with an insipid taste and a mild, pleasant aroma, of fair to good
quality; the seed is highly polyembryonic in a medium-thin stone. This is a
regular producer (C.W. Campbell, personal communication, 1995). The fruit
is often consumed green.
‘Langra’ (India)
Also called ‘Darbhanga’, ‘David Ford’, ‘Hadialaziz’, ‘Hajipur Langra’, ‘Har-
doi Langra’, ‘Lan Garhi’, ‘Langra Faquirwala’, ‘Sylhet’ and ‘Tikari’. The tree
is moderately vigorous, forming a dense canopy; the fruit is greenish yellow
with medium to big dark-green dots, ovalish to oblong, 8–10.5 cm long by
6.5–7.5 cm broad by 6–7 cm thick, weighing 235–375 g; the skin is medium
smooth, thick; the flesh is firm to soft, fibreless, lemon yellow, very sweet
with a strong, pleasant aroma, juice moderately abundant; seed is monoem-
bryonic in a medium-sized, flattened stone covered with dense, short and
soft fibre; quality is very good. Fruit ripens early to midseason (Gangolly et al.,
1957; R.J. Knight, Jr, personal communication, 1995).
‘Mabrouka’ (Egypt)
A major commercial cultivar considered to have been originally introduced
from India. The tree is moderately vigorous; the fruit (Plate 21) is large (481 g),
yellow with an orange to red blush and small, light-yellow, smooth dots;
ovate-oblong in shape (13.7 cm long by 8.9 cm wide by 8.2 cm thick) with a
thick, non-adherent skin relatively free of surface disease; the flesh is yellow,
firm and juicy with no objectionable fibre, moderately agreeable in taste, of
acceptable quality. The monoembryonic seed is in a moderately large (51 g)
stone. Fruit ripens late midseason, ships well and has been marketed in
Poland (Singh, 1960; Knight and Sanford, 1998).
‘Madame Francis’ (Haiti)
The tree is moderately vigorous, medium sized, forming an open canopy; the
fruit (Plate 22) is greenish to bright yellow, with no blush and a few large rus-
set dots, oblong, sigmoid with rounded base, 15–17 cm long by 8.5–11 cm
broad by 5.5–7.5 cm thick, weighing 370–520 g; the skin is thin, tender and
adherent; the flesh is soft and juicy with medium fibre, orange, rich spicy and
sweet with a pleasant aroma, of fair to good quality; seed is polyembryonic
R.J. Knight et al. 56
in a thin, papery stone. This cultivar ripens early to midseason and bears
well. Shipped to North American markets from Haiti nearly 10 months of the
year (R.J. Campbell, personal communication, 2007).
‘Mallika’ (India)
The tree is a moderately vigorous dwarf with a dense canopy; the fruit (Plate
23) is bright yellow with no blush and numerous small, light-yellow dots,
oblong with rounded base, 10–12 cm long by 6.5–7.5 cm broad by 5–5.5 cm
thick, weighing 280–450 g; the skin is thick, tough and easily separating; the
flesh is soft, tender and juicy with little fibre, deep yellow to orange, rich,
strongly aromatic and sweet, of excellent quality; seed is monoembryonic in
a medium-thick and woody stone. This cultivar ripens midseason and is an
irregular producer. This cultivar came from crossing ‘Neelum’ and ‘Dashe-
hari’ (Singh et al., 1972; Campbell, 1992).
‘Manila’ (Mexico)
The tree is large, vigorous, with an upright, open canopy; the fruit (Plate 24)
is bright yellow, sometimes with a light-pink blush, a few small reddish dots,
long and slender with rounded base and bluntly pointed apex sometimes
with a small beak, 12.5–14 cm long by 5.5–6 cm broad by 5–5.5 cm thick,
weighing 180–260 g; the skin is thin, medium tough and easily separating;
the flesh is medium firm and juicy, with little to abundant fibre, deep yellow,
sweet, rich and spicy in taste with a pleasant aroma, of good to very good
quality; seed is polyembryonic in a medium-thick and woody stone. This
cultivar ripens early midseason and crops fairly dependably. For a long time
‘Manila’ has been the most popular mango in Mexico.
‘Manzanillo’ (Mexico)
The tree is large, of medium vigour with an upright canopy; the fruit is yel-
lowish orange with 75% of the surface blushed an intense dark red with
numerous dots, oval with moderately flattened base, averaging 12 cm long
by 10 cm broad by 7.5 cm thick, and 660 g in weight; the flesh is low in fibre,
slightly subacid and very palatable, quality high; seed is monoembryonic in
a relatively small stone. This cultivar ripens early in the season but spread
over a 60-day harvest period. It bears heavily without pronounced alterna-
tion and the fruit stores and ships well (Núñez-Elisea, 1984).
‘Mesk’ (Egypt)
A major commercial cultivar. The tree is vigorous, the fruit small to medium
sized (312.5 g), yellow with a red blush, with small, corky yellow dots;
Mango Cultivars and Descriptors 57
ovate-oblong (11.3 cm long by 7.4 cm wide by 6.5 cm thick) with adherent
skin intermediate in thickness and fairly free of surface disease; the flesh is
orange, frequently jelly-seeded, with no objectionable fibres and a sweet,
agreeable taste of very good quality. The polyembryonic seed is in a moder-
ately large (52.5 g) stone. Fruit ripens late in the season (Knight and Sanford,
1998).
‘Mulgoa’ (India to Florida, USA)
Also spelled ‘Mugoba’ and ‘Mulgova’. The tree is large, vigorous with open,
spreading canopy; the fruit (Plate 25) is bright yellow with a pink blush and
numerous large white dots, oval to ovate with flattened base, 8.5–10.5 cm
long by 6.5–7.5 cm broad by 5–6 cm thick, weighing 340–450 g; the skin
is thick, medium tough and adherent; the flesh is soft, tender, melting and
juicy, with little fibre, lemon yellow, rich spicy and sweet with strong, pleas-
ant aroma, of good to excellent quality; seed is monoembryonic in a thick,
woody stone. This cultivar ripens midseason to late and is a shy, irregular
bearer. Introduced to Florida in 1889 and called ‘Mulgoba’, this is the seed
parent of ‘Haden’, first of a series of cultivars known as the Florida group. A
question exists whether the cultivar known in Florida is identical with the
Indian cultivar or is a seedling rootstock that survived after the scion was
killed by cold. In either case its superior quality ensured its retention and
propagation (Campbell, 1992). Literature serves to compound the nomencla-
tural confusion, as illustrated by Gangolly et al. (1957) whose ‘Mulgoa’ fruit,
yellow overall and roundish oblique with a deeply depressed stem insertion,
does not resemble the cultivar introduced to Florida. Singh (1960), on the
other hand, portrays a rounded, lightly blushed greenish yellow fruit that
closely resembles the Florida mango. Furthermore, vegetative propagation
of selected chance seedlings has resulted in a variety of clonal types carried
under this name in India (Ratnam and Chellapa, no date, post-1954).
‘Nabeel’ (Egypt)
A minor commercial cultivar. The fruit is large (495 g), green with small yel-
low dots that are smooth; ovate-oblong (14 cm long by 9 cm wide by 7 cm
thick), with adherent skin relatively free of surface disease; the flesh is
orange, firm and juicy, without objectionable fibre, with a passable but not out-
standingly pleasing taste and acceptable quality. The seed is polyembryonic
in a large (56.6 g) stone (Knight and Sanford, 1998).
‘Nam Doc Mai’ (Thailand)
The tree is vigorous, medium sized with upright, dense canopy; the fruit
(Plate 26) is greenish to bright yellow with a slight pink blush and numerous
R.J. Knight et al. 58
small green dots, long and slender, sigmoid in shape with a rounded base,
17–19 cm long by 7.5–8.5 cm broad by 6.5–7.5 cm thick, weighing 340–580 g;
the skin is medium thick, tender and easily separated from the flesh which is
soft, tender and juicy with no fibre, lemon yellow, rich, spicy and very sweet
with pleasant aroma, of excellent quality; seed is polyembryonic in a thin,
papery stone. This cultivar ripens early midseason, fruits regularly and may
have multiple crops in one season (Campbell, 1992). It is highly resistant to
foliar infection, and resistant to fruit infection by bacterial black spot in
Queensland (Mayers et al., 1988).
‘Neelum’ (India)
The tree is moderately vigorous with a small, compact canopy; the fruit is
bright yellow with no blush and numerous small white dots, oval with flat-
tened or slightly rounded base, 9.5–11 cm long by 7.5–8.5 cm broad by
6–6.5 cm thick, weighing 230–300 g; the skin is thick, tender and easily sepa-
rating; the flesh is soft, melting and juicy with no fibre, deep yellow, mild and
sweet with a delightfully pleasant aroma, of good to excellent quality; seed is
monoembryonic in a medium-thick, woody stone. This cultivar is a late,
heavy bearer (Campbell, 1992).
‘Nuwun Chan’ (Thailand)
The tree is moderately vigorous, small, upright with a dense canopy; the fruit
(Plate 27) is greenish yellow with a pink to red blush, numerous small green
dots, long and slender with a flattened base, 16–18 cm long by 7–8 cm broad
by 6–6.5 cm thick, weighing 340–500 g; the skin is thick, tough and easily
separating; the flesh is soft, melting, juicy with little fibre, pale yellow, mild
and sweet with a faint, pleasant aroma, of good eating quality; seed is poly-
embryonic in a thick, woody stone. This cultivar is an early, regular bearer.
Fruit is often eaten green (Campbell, 1992).
‘Okrung’ (Thailand)
The tree is moderately vigorous, medium sized and upright, forming a dense
canopy; the fruit (Plate 28) is green to greenish yellow with no blush and
numerous small white dots, oblong and sigmoid with a rounded base,
11–13 cm long by 5–6 cm broad by 4.5–5.5 cm thick, weighing 160–240 g; the
skin is thick, tough and medium adherent; the flesh is soft and juicy with
abundant fibre, yellow or greenish, mild, somewhat insipid and very sweet
with a pleasant aroma, of good quality; seed is polyembryonic in a thick,
woody stone. This cultivar ripens midseason, is a heavy producer and some-
times bears more than one crop/year (Campbell, 1992).
Mango Cultivars and Descriptors 59
‘Osteen’ (Florida, USA)
The tree is vigorous, medium sized, forming a dense canopy; the fruit (Plate
29) is yellow-orange with a purple or lavender blush and numerous small
white dots, oblong with rounded base, 12–15.5 cm long by 9–10.5 cm broad by
8.6–9.5 cm thick, weighing 500–760 g; the skin is thick, tough and easily sepa-
rating; the flesh is firm and juicy, with little fibre, lemon yellow, mild and sweet
with a pleasant aroma, of good quality; seed is monoembryonic in a thick and
woody stone. This cultivar ripens late midseason to late and is a regular
producer. It is a ‘Haden’ seedling (Campbell, 1992; Schnell et al., 2006).
‘Pairi’ (India)
Also written ‘Pairie’, ‘Paheri’ and ‘Pirie’; synonyms are said to be ‘Peter’,
‘Peter Pasand’, ‘Grape’, ‘Gohabunder’, ‘Nadusalai’, ‘Rasjuri’ and ‘Yerra Goa’.
The tree is moderately vigorous, forming a dense, rounded canopy; the fruit
(Plate 30) is medium sized, green to yellow with a bright red blush, roundish,
skin smooth, thick, flesh golden yellow, slightly juicy, fibreless, with a deli-
cious subacid taste, of excellent quality; the thick stone covered with short,
bristly fibre encloses monoembryonic seed (Popenoe, 1927; Singh, 1960). This
cultivar has long been popular as a dooryard fruit tree in Hawaii.
‘Palmer’ (Florida, USA)
The tree is moderately vigorous, forming a large, upright, tight canopy; the
fruit (Plate 31) is yellow-orange with a dark-red to crimson blush and a few
small white dots, oblong with rounded base, 12–15 cm long by 8.5–10 cm
broad by 6.5–7.5 cm thick, weighing 510–850 g; the skin is medium thick,
tough and adherent; the flesh is firm and melting with little fibre, orange-
yellow, mild and aromatic, of good quality; seed is monoembryonic in a
medium-thick woody stone. This is a late midseason cultivar and is a regular
bearer. It is a seedling of ‘Haden’ (Schnell et al., 2006). In Florida it is of minor
commercial importance (Campbell, 1992). It is grown in Israel and is the seed
parent of ‘Naomi’. It is attracting increased attention in the western hemi-
sphere export market as a result of its superior eating quality.
‘Rosa’ (Brazil)
The tree is medium sized, of slow growth with a rounded canopy; the fruit
(Plate 32) is yellow to rose-red on the side exposed to sun, oblong-cordiform
and medium sized; the skin is thick and smooth; the flesh is firm and mod-
erately juicy, fibrous, golden yellow, moderately sweet with a turpentine
aroma, of ordinary quality, susceptible to anthracnose disease; the seed is
polyembryonic in a small, oblong stone. This cultivar ripens midseason to
R.J. Knight et al. 60
late. It is one of the most important commercial cultivars in the Federal Dis-
trict of Brazil, used for juice as well as fresh consumption, and is one of the
most well-known cultivars in Brazil (Sampaio, 1980; L.C. Donadio, personal
communication, 1996; A. Pinto, personal communication, 1996).
‘Sensation’ (Florida, USA)
The tree is vigorous, with a moderately open, symmetrical canopy; the fruit
(Plate 33) is dark yellow with a prominent dark-red to purple blush that cov-
ers most of its surface, oval with rounded base and rounded apex, 9–11.5 cm
long by 7–8 cm broad by 6.5–7 cm thick, weighing 280–340 g; the skin is
medium thick, tough and easily separating; the flesh is firm and medium
juicy, fibreless, deep yellow, mild and sweet with a weak, pleasant aroma,
of fair to good quality; seed is monoembryonic in a thick, woody stone.
This cultivar ripens midseason to late (Campbell, 1992). It is a seedling of
‘Haden’ × ‘Brooks’ (Schnell et al., 2006), and the seed parent of ‘B74’. It alter-
nates severely, and in ‘on’ years the fruit may be clustered so heavily that it
becomes diseased before maturity, thus ‘Sensation’ is not of commercial
importance. It is highly resistant to bacterial black spot in Queensland (May-
ers et al., 1988), but often has severe internal breakdown (browning, water
soaking) (A.W. Whiley, personal communication, 1996).
‘Suvarnarekha’ (India)
Also called ‘Swarnarekha’ and ‘Sundri’. The tree is moderately vigorous and
tall, with a rounded, dense, spreading canopy; the fruit is light cadmium yel-
low with a blush of jasper red and abundant small, light-coloured dots, ovate
oblong with a base slightly flattened, of medium size, 11 cm long by 8.2 cm
broad, weighing 400 g; the skin is medium thick, easily separated, flesh soft,
fibreless, primrose yellow with a pleasant aroma, sweet taste and abundant
juice, of medium to good quality; seed is monoembryonic in an oblong-oval
stone covered with soft, short fibre. Ripens early in the season early and is
heavy bearing (Gangolly et al., 1957).
‘Tahar’ (Israel)
The tree is vigorous, medium sized, with an upright, dense canopy; the fruit
is bright yellow with a dark-red blush and numerous small white dots, ovate
with flattened base, 11.5–13 cm long by 8.9–9.5 cm broad by 7.5–8 cm thick,
weighing 360–520 g; the skin is thick, tough and easily separating; the flesh is
soft and juicy with little fibre, deep yellow, mild, aromatic and slightly insipid
with a strong odour not appreciated by many, of fair to good quality; seed is
monoembryonic in a medium-thick woody stone. This cultivar ripens in late
midseason and bears well in Israel (Campbell, 1992).
Mango Cultivars and Descriptors 61
‘Taimour’ (Egypt)
A major commercial cultivar. The tree is vigorous; the fruit (Plate 34) is large
(500 g), ripening late in the season, dark green with large light-brown dots,
smooth in texture, ovate-oblong (12.8 cm long by 8.4 cm wide by 8 cm thick),
with non-adherent skin of intermediate thickness, quite free from surface
disease; flesh is orange, firm (free of jelly seed) and juicy with no objection-
able fibre, of a delightfully rich, sweet taste and excellent quality. The seed is
polyembryonic in a medium-sized (50.8 g) stone (Knight and Sanford,
1998).
‘Tommy Atkins’ (Florida, USA)
The tree is vigorous, with a dense, rounded canopy; the fruit (Plate 35) is
orange-yellow, with a crimson or dark-red blush and numerous small, white
dots, oval to oblong, with broadly rounded base, 12–14.5 cm long by 10–13 cm
broad by 8.5–10 cm thick, weighing 450–700 g; the skin is thick, tough and
adherent; the flesh is firm and medium juicy; with a medium amount of fibre,
lemon to deep yellow, mild and sweet with a strong pleasant aroma, of fair to
good quality; seed is monoembryonic in a thick, woody stone. This cultivar
ripens early to midseason. It is a ‘Haden’ seedling (Schnell et al., 2006).
‘Tommy Atkins’ is the most important commercial cultivar in the western
hemisphere export mango market; it is highly resistant to anthracnose dis-
ease and handling and shipping stress, and a consistent, heavy producer
(Campbell, 1992). ‘Jelly seed’ (internal breakdown) is a serious problem in
the moist subtropics and tropics outside Florida, where the mango is grown
on calcareous, well-drained soil (A.W. Whiley, personal communication,
1996).
‘Totapuri’ (India)
Also called ‘Bangalora’, ‘Collector’, ‘Kallamai’, ‘Killi’ (‘Gilli’), ‘Mukku’, ‘Sand-
ersha’ and ‘Thevadiyamuthi’. The tree is of medium size, vigorous, spread-
ing with an open canopy; the fruit (Plate 36) is greenish yellow with a pink
blush and a few small, white dots, oblong, base rounded, apex rounded to
bluntly pointed with a large beak, 17.5–20 cm long by 9–11.5 cm broad by
8.5–10.5 cm thick, weighing 800–1100 g; the skin is thick, tough and adherent;
the flesh is firm and medium juicy with a weak, somewhat repugnant aroma,
of poor to fair quality; seed is monoembryonic in a thin, papery stone. This
cultivar ripens late midseason, is productive and regular bearing. Fruit cracks
when exposed to heavy rains at ripening time. ‘Totapuri’ was imported to
Florida twice, as ‘Sandersha’ in 1901 and as ‘Totapuri’ in the early 1960s. It is
the seed parent of ‘Anderson’ and ‘Brooks’, which is itself the parent of
‘Kent’. It is called ‘Totapuri’ in Bangalore, and ‘Bangalora’ in much of the rest
of India (Gangolly et al., 1957; Campbell, 1992).
R.J. Knight et al. 62
‘Turpentine’ (West Indies)
The tree is vigorous, with a large, spreading, rounded canopy; the fruit (Plate
37) is bright yellow with a few large white dots, occasionally with a pink
blush, oval with a flattened base, 7.5–8.5 cm long by 6.5–7.5 cm broad by
6–6.5 cm thick, weighing 140–200 g; the skin is thick, tough and easily sepa-
rating; the flesh is firm and juicy, with abundant coarse fibre, lemon yellow,
rich, aromatic, spicy, resinous and sweet with a strong, pleasant aroma, of
poor to fair quality; seed is polyembryonic in a thick, woody stone. This cul-
tivar ripens early midseason to late midseason and is a heavy producer but
may alternate. It is commonly used as a grafting stock (Campbell, 1992).
‘Vallenato’ (Colombia)
The tree is vigorous, with an upright, dense canopy; the fruit (Plate 38) is
bright yellow, with a crimson blush, oblong with flattened base, 8–9 cm long
by 7–8 cm broad by 6–7 cm thick, weighing 195–340 g; the skin is thin, tough
and adherent; the flesh is firm, juicy with abundant fine fibre (not objection-
able), pale yellow, mild and sweet with a strong, pleasant aroma, of good to
excellent quality; seed is monoembryonic. This cultivar ripens in early mid-
season (R.J. Campbell, personal communication, 1995).
‘Van Dyke’ (Florida, USA)
The tree is moderately vigorous, with a large, open canopy; the fruit (Plate
39) is bright yellow with a bright red or crimson blush, oval with rounded
base, 9–11.5 cm long by 7.5–9.5 cm broad by 7–8 cm thick, weighing 250–
520 g; the skin is thick, tough and easily separating; the flesh is quite firm,
melting and juicy with little fibre, orange-yellow, rich, spicy and sweet with
a strong, pleasant aroma, of good to excellent quality, but susceptible to inter-
nal breakdown; seed is monoembryonic in a medium-thick, woody stone.
This cultivar ripens in late midseason and is a regular, heavy producer. It is a
seedling of ‘Haden’ (Campbell, 1992; Schnell et al., 2006).
‘White Succari’ (Egypt)
A cultivar of major importance. The tree is vigorous; the fruit is medium
large (410 g), greenish yellow with yellow overlay and small, brown dots of
smooth texture, ovate-oblong in shape (11.25 cm long by 8.3 cm wide by
8.0 cm thick), with thin adherent skin reasonably free of surface disease;
flesh is orange, yielding and juicy with no objectionable fibres, of an agree-
able sweet taste and very good quality. The seed is polyembryonic in a
moderate to large-sized stone (49 g), the season early (Knight and Sanford,
1998).
Mango Cultivars and Descriptors 63
‘Zebda’ (Egypt)
A cultivar of major importance. The tree is vigorous and regularly produc-
tive; the fruit (Plate 40) is large (660 g), green with no overlay and small,
brown dots of smooth texture, oblong-cylindrical in shape (14.6 cm long by
9.7 cm wide by 8.3 cm thick), with non-adherent skin quite free of surface
disease; flesh is deep orange, firm and juicy, with no objectionable fibre and
a mild, sweet taste, of acceptable quality. The seed is polyembryonic in a
moderately small (52 g) stone. This cultivar is of late-midseason maturity
(Knight and Sanford, 1998). It is highly tolerant of anthracnose and resistant
to malformation (R.C. Ploetz, personal communication, 2007).
3.4 Conclusion
The mango fruit’s nutritional value, aesthetic and gustatory appeal have
assured its growing importance in non-traditional markets since the late
1950s, as it has been introduced to consumers previously unacquainted with
it. Furthermore, the migration of large populations from South-east Asia and
other regions where this fruit is a traditional crop to metropolitan centres
where it has not been well known has created a permanent demand for it in
these new markets. An additional factor permitting market expansion has
been the growing mango production in areas previously unimportant in
world commerce such as Mexico, Brazil, Australia, West Africa, Israel, Flor-
ida and the Canary Islands. The fact that most new markets are remote from
areas of production has necessitated selection of cultivars for fresh market
sale that are dependably productive and resistant to harvest, handling and
shipping stress, with relatively long shelf life, for example ‘Tommy Atkins’,
‘Keitt’ and ‘Madame Francis’. The fruit quality of mango cultivars well suited
to packing and shipping has been a secondary consideration, and is gener-
ally not so high as that of cultivars acknowledged to be superior for eating.
Economic factors obviously must dictate what is grown for the fresh market.
The commercial market for processed mango products permits other
cultivars to be utilized, and these may vary with the product that is mar-
keted. Cultivars chosen for purée or juice preparation are likely to be quite
different from those used for manufacture of chutney or other products
requiring pulp that maintains its integrity after it is cooked. ‘Totapuri’ (‘Sand-
ersha’) or ‘Turpentine’, for example, considered mediocre for fresh consump-
tion, can be used to prepare excellent chutneys, as can many ‘criollo’ types in
the West Indies. ‘Tommy Atkins’ makes outstandingly good dried fruit sec-
tions, sweet and aromatic, even though its fresh-fruit quality is generally
conceded not to be high. Mango butter and mango leather are other products
that are appreciated by many who know them (see Raymundo et al., Chapter
17, this volume; Campbell and Campbell, 1983; Campbell and Smith, 1987).
As more fruit that is wholesome, but not of export quality, becomes available
in areas of increasing production, it is likely that processed mango products
will become more common.
R.J. Knight et al. 64
Despite the recognized high quality of many well-known mango cultivars,
considerable cultivar improvement is still needed in most regions of culture
before anything approaching perfection is likely to be achieved (Table 3.1).
For any given area, cultivars that combine adequate resistance to disease and
packing and shipping stress, regular heavy production, high quality, and
attractive appearance throughout a long bearing season are all requisites.
Production of seedlings from controlled crossing of different parents having
desired characters, followed by vigorous selection and evaluation of the
resultant selections, can produce such improved cultivars. Pursuit of com-
mon goals, including the cooperative exchange and testing of elite germ-
plasm in different regions of production, can accelerate progress towards this
objective (Lavi et al., 1989, 1993; Knight, 1993). Such interregional and inter-
national activities are to be encouraged because of their potential for advanc-
ing mango production and utilization in the world.
Acknowledgements
For their help in reviewing portions of this chapter and/or contributing vast
quantities of information on mango cultivar descriptors and attributes, the
authors are profoundly grateful to the following: Dr N. Balasundaram, Head,
Sugarcane Breeding Institute, Regional Centre, Karnal, India 132001; the late
Dr Carl W. Campbell, Tropical Research and Education Center, 18905 SW 280
Table 3.1. Ratings of selected mango cultivars grown in Florida (Source: Knight, 1993).
Cultivar Shape
a
Size
b
Firmness
c
Colour
d
Anthracnose
e
Fibre
f
Taste
g
Yield
h
Score
i
‘Alphonso’ 3 5 7 2 3 7 9 1
h
x
‘Boribo’ 3 8 8 4 7 9 5 6 x
‘Carabao’ 5 6 7 3 5 9 8 6
h
x
‘Haden’ 3 9 8 8 5 7 7 3
h
x
‘Keitt’ 4 10 9 6 8 9 8 8 ///
‘Kensington’ 3 8 7 7 7 8 7 6 /
‘Langra’ 2 6 8 3 5 8 8 3
h
x
‘Pope’ 3 9 5 7 2 8 8 1 x
‘Tommy
Atkins’
3 9 9 9 9 6 6 7 ///
‘Van Dyke’ 3 7 10 9 7 8 7 6 ///
a
Ratings of 1 (round) to 5 (long) indicate fruit shape, not its desirability.
b
Ratings below 6 justify discard; those of 7 and above show size only, not merit.
c
Ratings of 1–10 where 1 = least and 10 = most.
d
Ratings of 1–10 where 1 = least and 10 = most.
e
Ratings of 1–10 where 1 = most and 10 = least susceptible.
f
Ratings of 1–10 where 1 = most and 10 = least.
g
Ratings of 1–10 where 1 = worst and 10 = best.
h
Trends markedly towards alternate bearing.
i
One or more checks (/) show overall value; (x) indicates no commercial acceptability.
Mango Cultivars and Descriptors 65
Street, Homestead, Florida 33031 USA; Luis C. Donadio, Universidade Estad-
ual Paulista, Rodavia Carlos Tonanni KM5, Jaboticabal, 14870, São Paulo,
Brazil; Dr Shmuel Gazit, Department of Horticulture, Hebrew University of
Jerusalem, Rehovot 76100, Israel; Mr Remy LePrette, Directeur, Interfel, 155
Rue F.G. Poissonierre, Paris 75009, France; Dr Alberto C. Pinto, Lider de Pro-
jeto/CPAC, EMBRAPA, Caixa Postal 08 223, CEP 73301-970, Brasilia, DF,
Brazil; Dr Eli Tomer, Department of Fruit Trees, the Volcani Centre, Institute
of Horticulture, PO Box 6, Bet Dagan 50-250, Israel; Dr Anthony W. Whiley,
Maroochy Horticultural Research Centre, PO Box 5093 SCMC, Nambour,
Queensland 4560, Australia.
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nitrate. HortScience 14, 527–528.
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(ed. R.E. Litz) 67
4 Breeding and Genetics
C.P.A. Iyer
1
and R.J. Schnell
2
1
Indian Institute of Horticultural Research, Bangalore, India
2
USDA ARS, National Germplasm Repository, Miami, Florida, USA
4.1 Introduction 68
4.2 Origin of Cultivars and Distribution 68
Mangifera species and mango 68
History of cultivation 69
Impact of Florida mangoes 70
4.3 Reproductive Mechanisms 70
Polyembryony 70
Floral biology and pollination 71
Incompatibility 72
Cytology 72
4.4 Inheritance of Characters 73
Dwarfness, regular bearing and precocity 73
Flesh colour 74
Skin colour 74
Flowering time 74
Beak 74
Disease resistance 74
Other horticultural traits 75
4.5 Breeding Objectives 75
General objectives 75
Specific objectives 76
4.6 Methods of Breeding 78
Selection from open-pollinated seedlings 78
Controlled pollination 79
4.7 Handling of Hybrid Populations and Selection 80
Criteria for initial selection 80
Pre-selection 81
Potential for marker assisted selection (MAS) 81
Molecular markers 81
4.8 Minimizing Problems in Breeding 83
Heavy fruit drop 83
C.P.A. Iyer and R.J. Schnell 68
Long juvenile phase 83
Polyembryony 84
4.9 Achievements of Conventional Breeding 84
India 85
Other countries 86
4.10 Mutations 87
Somatic mutations 87
Induced mutations 88
4.11 Breeding Potential of Wild Species 88
4.12 Conclusions 89
4.1 Introduction
Mango has been considered to be a difficult plant species to improve in
breeding programmes because of certain inherent characteristics including:
(i) a long juvenile phase; (ii) a high level of heterozygosity resulting in unpre-
dictable outcomes in hybridization; (iii) only one seed per fruit; (iv) heavy
fruit drop leading to low retention of crossed fruits; (v) polyembryony in
many cultivars; and (vi) the large area required for a meaningful assessment
of hybrids. Despite these drawbacks, mango breeding can be successful
because of its wide range of genetic variation and the ease with which a
selected hybrid can be vegetatively propagated. Barring a few hybrid variet-
ies resulting from planned hybridization programmes, which are now gain-
ing increased attention, almost all known cultivars have resulted from the
selection of chance seedlings from natural cross-pollinations. However, in
Florida, a number of cultivars have resulted from the screening of seedlings
from known mother plants. Most of the present-day mango cultivars were
selected on the Indian subcontinent; these selections were made based
mainly on fruit quality, with very little emphasis on modern horticultural
and industrial requirements. These requirements include precocity, dwarf-
ness, heavy and regular bearing, absence of physiological disorders, resis-
tance to disease and pests and good shipping qualities. With decreasing land
availability and the rising cost of labour, tree architecture requirements have
also changed. The need for new cultivars to meet these requisites pinpoints
the importance of planned hybridization rather than merely depending on
chance seedlings. Current knowledge of hybridization techniques, inheri-
tance patterns, management of hybrid populations and the development of
genetic markers have greatly reduced the uncertainty in mango breeding.
4.2 Origin of Cultivars and Distribution
Mangifera species and mango
Almost all the commercial cultivars belong to Mangifera indica. However,
a few commercial varieties of South-east Asia belong to other species, i.e.
M. altissima, M. caesia, M. foetida, M. griffithi, M. odorata, M. pentadra, M. sylvatica,
Breeding and Genetics 69
M. zeylanica, M. laurina, M. lagenifera, M. cochinchinensis, etc. The monoem-
bryonic mango (M. indica) originated in north-eastern India (Assam), the
Indo-Myanmar border region and Bangladesh (Chittagong Hill tract), where
it is still found as a wild tree, with very small fruits. It may also occur in the
lower Himalayan tract, near Nepal, Bhutan and Sikkim. Polyembryonic
mangoes are considered to have originated in South-east Asia. Wild man-
goes, representing different Mangifera species, can be found in tropical Asia,
particularly north-eastern India, Sri Lanka, Myanmar, Thailand, Indo-China,
southern China, Malaysia, Indonesia, Papua New Guinea, the Philippines
and as far as the Solomon and Caroline Islands in the east. There are more
than 60 species worldwide. The highest specific diversity is found in the
heart of the distribution area of the genus Mangifera; the Malay Peninsula,
Borneo and Sumatra (Bompard, 1993).
History of cultivation
Mango has been cultivated in India for at least 4000 years and over 1000 vari-
eties are recognized there (Mukherjee, 1953). Almost all of them are selec-
tions made from naturally occurring, open-pollinated seedlings. However,
based on random amplification of polymorphic DNA (RAPD) analysis, Rav-
ishankar et al. (2004) felt that the mono- and polyembryonic types of Indian
mango cultivars have a different genetic base, and that the polyembryonic
types might have been introduced from South-east Asia and are unlikely to
have originated in India. Mango culture gradually spread to tropical and
subtropical countries throughout the world, where selections were made
that were adapted to particular growing conditions. Thus, selection by man
has played the most significant role in the development of new mango culti-
vars. The explorers who tasted the mango in the regions of its origin were
enchanted with its aromatic qualities, ambrosial flavour and creamy, smooth
and silky texture, and introduced the fruit to other tropical regions. The
spread of Hinduism and Buddhism assisted in the distribution of mangoes in
South-east Asia. The Chinese traveller Hwen T’sang who visited India in the
first half of the 7th century ad returned to China with the mango. The mango
was known in Baghdad in the 7th century.
The Persians or Omanis may have taken mangoes to East Africa around
the 10th century ad. The fruit was introduced to East and West Africa in the
early 16th century by the Portuguese and thence into Brazil. After being
established in Brazil, the mango was carried to the West Indies, being first
planted in Barbados by about 1742 and later in the Dominican Republic and
Jamaica (about 1782). The mango was introduced into Mexico from the Phil-
ippines by the Spanish and also from the West Indies (Morton, 1987). Duval
et al. (2006) developed microsatellite markers for studying the genetic vari-
ability of Caribbean mangoes and concluded that there were two routes of
mango to the French West Indies, namely, cultivars grown in Central Amer-
ica (Mexico) and South America (Colombia) introduced from South-east Asia
and also from former French colonies in the Indian Ocean. As the mango
C.P.A. Iyer and R.J. Schnell 70
adapted to new locales, new cultivars were selected based on local adapta-
tion and fruit preferences.
Impact of Florida mangoes
The first recorded successful introduction of mango into Florida (USA) was
made in 1861 (Knight, 1980). The earliest introductions were from the West
Indies and India, followed by the introduction of several hundred accessions
in the 20th century from South-east Asia, India and from other mango-growing
areas of the world (Florida Mango Forum, 1951). The introduction of mangoes
into Florida and subsequent development of a Florida group of mangoes has
been reviewed by Knight and Schnell (1994). The Florida mango cultivars are
unique in that they are hybrids between Indian cultivars (primarily mono-
embryonic) and the South-east Asian cultivars (primarily polyembryonic)
selected under south Florida conditions. The mango breeding system favours
out-crossing. Therefore, the proximity of numerous genotypes of disparate
geographical origins led to the production of many new seedlings by inter-
pollination in Florida (Knight and Schnell, 1993). Florida selections are
therefore not the result of a formal breeding programme. Early Florida selec-
tions were made by growers and enthusiasts and historical information is
often anecdotal. The Florida Mango Forum, established in 1938 for the
advancement of mango production, documented historical information on
the parentage of Florida cultivars in their proceedings. In addition to the
United States Department of Agriculture (USDA) Germplasm Resources In-
formation Network (GRIN) database, several sources compile information
on Florida mango selections and introduction of accessions to Florida ( Ruehle
and Ledin, 1956; Singh, 1960; Campbell and Campbell, 1993; Schnell et al.,
2006). With the exception of South-east Asia, Australia and some African
countries, which cultivate mostly locally selected varieties, the majority of
countries produce cultivars developed in Florida, i.e. ‘Haden’, ‘Tommy
Atkins’ and ‘Keitt’ (Galan Sauco, 1997). These Florida selections are now
widely grown commercial cultivars affording production stability across
many environments (see Mukherjee and Litz, Chapter 1, this volume).
4.3 Reproductive Mechanisms
Polyembryony
Nucellar embryos
Mangoes can be classified into two groups, monoembryonic and polyembry-
onic, based on their mode of reproduction from seeds. In general, monoem-
bryonic seeds are found in the sub-tropical group (Indian type) and the
polyembryonic seeds in the tropical group (South-east Asian). Monoembry-
onic mango seeds each contain a single zygotic embryo, and hence only one
seedling per seed that is of probable hybrid origin. Polyembryonic mango
Breeding and Genetics 71
seeds can contain one or more embryos, one of which is usually, but not
always zygotic. Adventitious embryos develop from the nucellus, a maternal
tissue surrounding the embryo sac, and consequently the seedlings of poly-
embryonic mangoes are genetically identical to the maternal parent. Adven-
titious embryos can also originate by direct budding from the cotyledons and
hypocotyls of other nucellar embryos (Juliano, 1934). According to Mahesh-
wari and Rangaswamy (1958), the nucellar cells destined to form adventi-
tious embryos are recognizable by their dense cytoplasm and starchy
contents. They gradually push into the embryo sac cavity where they divide
and differentiate into embryos.
Inheritance of polyembryony
Polyembryony is genetically determined. Leroy (1947) considered that adven-
tive embryony probably reflects the effect of one or more recessive genes.
This view was supported by Sturrock (1968), whose study of the progenies of
monoembryonic mango hybridized with polyembryonic cultivars indicated
that monoembryony was possibly a dominant trait. In contrast, Aron et al.
(1998) and Brettell et al. (2004) observed that polyembryony in mango is con-
trolled by a single dominant gene. Schnell et al. (2006) reported that 58 of the
Florida cultivars had been classified with 50 being monoembryonic and eight
polyembryonic. Information from the Florida cultivars parentage analysis
using 25 microsatellite markers supported the findings of Aron et al. (1998)
where polyembryony was found to be dominant. ‘Haden’ is a cross of the
monoembryonic ‘Mulgoba’ and the polyembryonic ‘Turpentine’. If we assume
that a single dominant gene controls this trait, all of the Indian cultivars in
Florida must be homozygous recessive and the ‘Turpentine’ parent of ‘Haden’
must have been heterozygous. The evidence suggests that ‘Haden’ inherited
the recessive allele from ‘Turpentine’, as all identified progeny of ‘Haden’ are
monoembryonic with the exception of ‘Winters’. The most probable pollen
parent of ‘Winters’ is ‘Ono’, a polyembryonic cultivar from Hawaii. The fre-
quency of this dominant allele is low in the Florida population and absent
from the Indian cultivars in Florida. In view of these interesting findings, and
since a thorough knowledge of inheritance of polyembryony is essential for
speculating the origin of M. indica, more work on these lines is warranted.
Floral biology and pollination
The mango inflorescence is primarily terminal, although axillary and multi-
ple panicles may also arise from axillary buds. Both perfect (hermaphrodite)
and staminate (male) flowers occur in the same inflorescence. The total num-
ber of flowers in a panicle may vary from 1000 to 6000, depending on the
cultivar (Mukherjee, 1953). Initial fruit set in mango is directly related to the
proportion of perfect flowers, although the final fruit set does not necessarily
depend on this ratio (Iyer et al., 1989). It appears that the proportion of per-
fect flowers in a cultivar becomes critical for optimum fruit set only when the
proportion drops to 1%.
C.P.A. Iyer and R.J. Schnell 72
Flowers begin to open early in the morning and anthesis has generally
been completed by noon. The greatest number of flowers opens between 9
and 10 a.m. Although the receptivity of the stigma continues for 72 h after
anthesis, it is most receptive during the first 6 h; however, there are reports
that the stigma can become receptive even before anthesis has occurred
(Singh, 1960). The minimum time required for pollen grains to germinate is
1.5 h (Sen et al., 1946; Singh, 1954; Spencer and Kennard, 1955). Singh and
Singh (1952) observed 98% pollen viability after 11 months in storage at 7°C
and 25% relative humidity (RH), and 65.7% viability after 24 months of stor-
age at 0°C and 25% RH.
Mango is cross-pollinated, which is carried out by insects such as the
common housefly, honeybees and thrips, and possibly by other insects
al-though to a lesser extent. Pollination by wind and gravity has been sug-
gested to occur in mango (Popenoe, 1917; Maheshwari, 1934; Malik, 1951). In
nature, > 50% of flowers do not receive any pollen. Some workers had sug-
gested that self-pollination in certain cultivars can also occur quite frequently
(Dijkman and Soule, 1951). Studies by Issarakraisila and Considine (1994)
have shown that for polyembryonic ‘Kensington’, a night temperature of
< 10°C results in pollen grains with low viability (< 50%). The optimum

temperature for normal meiosis is between 15 and 33°C with 70–85%
viability.
Incompatibility
Although the existence of self-sterility in mango was suspected several years
ago (Ruehle and Lynch, 1948, cited in Sharma and Singh, 1970; Dijkman and
Soule, 1951), the prevalence of self-incompatibility was clearly established in
monoembryonic ‘Dashehari’ by Singh et al. (1962). Subsequently, detailed
studies indicated that the four popular monoembryonic cultivars of northern
India (i.e. ‘Dashehari’, ‘Langra’, ‘Chausa’ and ‘Bombay Green’) were self-
incompatible (Mukherjee et al., 1968; Sharma and Singh, 1970). Embryo-
logical studies have shown that although fertilization takes place after
self-pollination, degeneration of endosperm occurs 15 days after pollination
involving self-incompatible parents (Mukherjee et al., 1968). The self-
incompatibility system operating in mango appears to be of the sporophytic
type. Instances of cross-incompatibility among certain mango cultivars
have also been reported (Ram et al., 1976), necessitating the identification of
suitable pollinizers for mango.
Using an approach involving isozyme analysis, Dag et al. (2006) have
initiated studies in many commercial mango cultivars in Israel and con-
cluded that self-pollination is not a yield-limiting factor in monoembryonic
‘Maya’ and the practice of planting ‘Maya’ in solid blocks is sound. They had
obtained similar results earlier with monoembryonic ‘Tommy Atkins’ (Dag
et al., 1997).
Breeding and Genetics 73
Cytology
Chromosome number
Information on the cytology of mango is quite limited. Only a few Mangifera
species (i.e. M. indica, M. caloneura, M. sylvatica, M. foetida, M. caesia, M. odorata
and M. zeylancia) have been studied, and were found to have chromosome
numbers of 2n = 2x = 40 and n = x = 20 (Mukherjee, 1950, 1957; Roy and
Visweswariya, 1951). Chromosome numbers and ploidy status of other species
are yet to be studied. The only exception to this chromosome number that has
been reported to date (Roy and Visweswariya, 1951) involves ‘Vallikolamban’,
which was reported to be tetraploid (2n = 4x = 80), although subsequent stud-
ies have indicated that it is only a diploid (Majumder and Sharma, 1990).
Polyploidy
Mango has been referred to as an allopolyploid. Due to the presence of
secondary associations at metaphase of meiosis, Mukherjee (1950) suggested
that the basic chromosome number of Mangifera is n = 8. In addition, the high
number of somatic chromosomes and the correspondingly high number of
nucleolar chromosomes led him to conclude that mango is an allopolyploid.
However, the evidence used to arrive at this conclusion is not unequivocal.
In fact, the molecular marker evidence is antithetical to this conclusion.
Results from Duval et al. (2005), Viruel et al. (2005) and Schnell et al. (2005,
2006) using microsatellite markers all indicate that M. indica is diploid.
Although many wild Mangifera species are potentially valuable for crop
improvement, they are yet to be exploited. Mukherjee (1963) felt that the
different Mangifera species could intercross easily, based on the success ob-
tained with interspecific crosses between M. zeylanica and M. odorata.
4.4 Inheritance of Characters
High heterozygosity in the cultivars that are used in hybridization and the
inadequate number of hybrid progenies realized has made accurate genetic
analysis in mango very difficult. However, based on limited data, some indi-
cations are available which would be useful in selecting parents in breeding
programmes designed with specific objectives. In studies of the distribution
of different traits in seedlings derived from open-pollination (where the pol-
len parent is unknown), Lavi et al. (1989) observed: (i) there is no maternal
effect on juvenile period and fertility; (ii) there is a slight effect of the female
parent on fruit taste and size; (iii) there is a maternal parent effect on harvest
season and fruit colour.
Dwarfness, regular bearing and precocity
An analysis based on observations of more than 1000 hybrids, involving sev-
eral combinations, has revealed that dwarfness, regular bearing and precocity
C.P.A. Iyer and R.J. Schnell 74
are controlled by recessive genes (Sharma and Majumder, 1988a). Regularity
of bearing appears to be linked with precocity. Characters contributing to
biennial bearing are dominant over those governing regular bearing habit.
Flesh colour
Sharma (1987) considered that additive gene action may be involved in the
inheritance of flesh colour; however, studies involving monoembryonic
‘Alphonso’ and ‘Neelum’ have indicated that light yellow is dominant over
orange-yellow (Iyer, 1991).
Skin colour
With regard to skin colour of fruit, Sharma (1987) observed that when red
cultivars were crossed with green cultivars, the F
1
seedlings exhibited vari-
ous gradations of red. Iyer and Subramanyam (1987) also found a wide array
of colours in the hybrids when the coloured monoembryonic ‘Janardhan
Pasand’ was crossed with green-fruited cultivars, indicating that colour is
mediated by a number of loci.
Flowering time
The flowering response of mango cultivars in subtropical and tropical envi-
ronments differs greatly (see Davenport, Chapter 5 this volume). Trees can be
stimulated to flower under certain conditions in tropical environments using
ethephon; however, this is ineffective in subtropical environments. Schnell and
Knight (1998) investigated the repeatability of flowering using eight cultivars
over six harvest cycles (years), collecting data weekly. Three characters were
evaluated: days to bloom (DTB) (from 1 November in each year), days in bloom
(DIB), and days in bloom and fruit (DIBF). Significant differences were detected
for all three characters for both years and cultivars. Significant differences were
not detected for replicate trees within cultivars. Repeatability (R) of the flower
phenology characters was high (R = 0.73, 0.88 and 0.77 for DTB, DIB and DIBF,
respectively). This indicates that much of the variation is heritable and useful
for extending flowering times in subtropical environments.
Beak
The presence of a beak on mango fruit appears to be a dominant character
since most of the hybrid plants had this feature when monoembryonic ‘Ban-
galora’ (‘Totapuri’) was used as one of the parents in controlled crosses (Iyer
and Subramanyam, 1979). Bunch bearing was found to be a dominant char-
acter, as indicated in many crosses (Sharma et al., 1972) involving bunch-
bearing types with single-fruited cultivars.
Breeding and Genetics 75
Disease resistance
Bacterial canker (Xanthomonas campestris pv. mangiferaeindicae) resistance
appears to have cytoplasmic inheritance. Whenever ‘Neelum’, a susceptible
cultivar, is used as the female parent, susceptibility is transmitted to all the
hybrids, irrespective of the male parent (Sharma and Majumder, 1988a). It
has been suggested that internal breakdown (spongy tissue) is mediated by
recessive genes (Iyer, 1991). Susceptibility to ‘mango malformation’ appears
to be dominant, since crosses involving resistant ‘Bhadauran’ did not yield
any resistant hybrids (Sharma and Majumder, 1988a).
Other horticultural traits
The genetics of inheritance of various horticultural traits appears to be
unclear. More knowledge will be forthcoming only when large-scale con-
trolled hybridization experiments are undertaken at different mango research
centres. However, the information now available for some of the characters
is very useful in deciding which parental genotypes ought to be used in
hybridization programmes.
Brettell et al. (2004) subjected the large number of mango hybrids obtained
from the Australian National Mango Breeding Programme to a biometrical
analysis. Their data indicated that many of the important fruit quality aspects,
including fruit weight, fruit shape, ground skin colour, fruit width and pulp
depth have high heritabilities and can therefore be readily selected in a breed-
ing programme. Of particular interest is the observation that a high frequency
of hybrids with a red or burgundy blush can be recovered from crosses where
one parent has an intense red blush. Similarly, while the unique flavour com-
pounds associated with ‘Kensington Pride’ are also found in nearly 50% of
the hybrids involving ‘Kensington Pride’, leaf fragrance was not found to be
a reliable predictor of fruit flavour.
4.5 Breeding Objectives
General objectives
Breeding objectives vary from region to region, depending on the specific
trait(s) for which improvement is sought. However, they can be broadly gen-
eralized to consist of the development of cultivars with: (i) regular bearing;
(ii) dwarf tree habit; (iii) precocity; (iv) attractive, good sized (300–500 g),
good quality fruits (appealing flavour and firm flesh without fibres); (v)
resistance to major diseases and pests; (vi) freedom from physiological dis-
orders; and (vii) good shipping qualities and shelf life. While it would be
hard to combine all these characteristics within a relatively short time, espe-
cially resistance to all major diseases and pests, all of these characteristics are
basic for commercial success.
C.P.A. Iyer and R.J. Schnell 76
With regard to the improvement of rootstocks by breeding, the main
desirable features are: (i) polyembryony; (ii) dwarfing; (iii) tolerance of
adverse soil conditions (high pH, calcareous soil, etc.); and (iv) good scion-
compatibility.
Specific objectives
In addition to improving general characters such as yield and quality, breed-
ing has also been undertaken for certain specific purposes.
Dwarfness
Because of the obvious benefits of comparatively dwarf trees for orchard man-
agement and fruit quality, attempts have been focused on obtaining hybrids
with a dwarf tree framework. Breeding for dwarfness is important in mango,
since a consistent dwarfing effect of any rootstock has not been established to
date. The Indian cultivars that could be useful as a source for dwarfness
include ‘Kerla Dwarf’, ‘Janardan Pasand’, ‘Manjeera’, ‘Amrapali’, ‘Creeping’
(Iyer and Subramanyam, 1986) and ‘Nileswar Dwarf’ (Singh, 1990).
Regular bearing
The causes of irregular bearing vary from region to region. In general, the
main reason for alternate bearing, particularly in subtropical India, is the
lack of initiation of vegetative growth soon after fruiting. However, two cul-
tivars, ‘Neelum’ and ‘Bangalora’ (‘Totapuri’), which are regular bearers, have
been extensively used as either of the parents in a hybridization programme
to transfer the regular bearing habit to hybrids. ‘Neelum’ has been observed
to be a good combiner and has contributed to the evolution of many regular-
bearing Indian hybrid cultivars. However, ‘Bangalora’ is not a suitable par-
ent since the hybrids possess very prominent beaks and their fruit quality is
invariably poor. The regular bearing Florida cultivars (i.e. ‘Tommy Atkins’,
‘Keitt’, etc.) also have potential as parents.
Fruit colour
Most of the commercial cultivars in South-east Asia possess green skin.
Efforts are underway to produce new hybrid cultivars that retain the good
qualities of these fruits together with attractive skin colour, so that they will
occupy a better position in international trade. Since good skin colour has
been shown to be transmissible to hybrids from suitably coloured parental
cultivars, a number of cultivars with coloured skin are being used for hybrid-
ization. In general, the attractively coloured Florida cultivars have been
found to be suitable parents. In addition, there are several Indian cultivars
(e.g. ‘Janardan Pasand’, ‘Suvarnarekha’, etc.) that would be suitable for use
as parents for this purpose.
In Florida, the skin colour of the mango is an important factor and red
skin is considered essential for mangoes shipped to northern markets. In the
past, the evaluation of mango colour has been subjective and based on visual
Breeding and Genetics 77
ratings. Large errors are associated with these types of ratings, which makes
evaluation based on fruit colour difficult. Ayala-Silva et al. (2005) used a colo-
rimeter to quantify fruit colour, quality and differentiation among cultivars.
Mango colour was measured with a Minolta Chroma Meter CR-400 (Osaka,
Japan) portable tristimulus colorimeter and fruit chromaticity was recorded
in Commission Internationale de l’Eclairage (CIE) L
*
, a
*
and b
*
colour space
coordinates. In this system of colour representation the values L
*
, a
*
and b
*

describe a uniform three-dimensional CIE colour space. If the L
*
, a
*
and b
*

coordinates are known, then the colour is not only described, but also located
in quantifiable space. Maternal half-sib families (MHS) of the mango culti-
vars, ‘Keitt’, ‘Tommy Atkins’, ‘Tyler Premier’, ‘Mamita’, ‘White Alfonso’ and
‘Sandersha’ were evaluated along with two parental check clones, ‘Tommy
Atkins’ and ‘Keitt’. Significant differences were found for each of the L
*
, a
*
and
b
*
colour space coordinates. Further work is underway to estimate the herita-
bility of these traits to estimate their usefulness for breeding and selection.
Disease resistance
MANGO MALFORMATION. Although no breeding work has been reported that spe-
cifically addresses disease or pest resistance/tolerance, cultivars are known to
show varying degrees of susceptibility to biotic stress (see Ploetz and Freeman,
Chapter 8, this volume ). Mango malformation, caused by Fusarium subglutin-
ans, is a very serious disease that has threatened the very survival of the
mango industry in many subtropical mango-growing regions. As there are
no reliable cultural and chemical control measures available, breeding for
resistance/tolerance has been attempted using ‘Bhadauran’ as the resistant
parent; however, all of the F
1
hybrids were susceptible to the disease (Sharma
and Majumder, 1988a). In this respect, the observations of Ram et al. (1987)
are very encouraging. Out of 102 cultivars screened, three of them, namely,
‘Bhydayam Dula’, ‘Samar Bahist Rampur’ and ‘Mian Sahib’, were free of
malformation and could be tried as one of the parents in hybridization.
BACTERIAL CANKER. Bacterial canker is a serious problem with many cultivars.
The only cultivar possessing true resistance to canker is ‘Bombay Green’
(Prakash and Srivastava, 1987) and hence could be a potential gene donor.
ANTHRACNOSE. Anthracnose, caused by Colletotrichum gloeosporioides Penz., is
the most widespread disease in all mango-growing countries, manifesting
itself in blossom blight, peduncle blight, leaf spot, twig blight, wither tip, fruit
russetting and fruit rot. ‘Tommy Atkins’ is moderately tolerant of anthracnose
and coupled with its other desirable qualities (i.e., regular bearing, fruit colour,
etc.) should be a good parent in breeding programmes. In addition, ‘Parish’
and ‘Fairchild’ have been reported to be relatively resistant (Yee, 1958).
POWDERY MILDEW. Powdery mildew caused by Oidium mangiferae Berthet, has
been reported to cause heavy loss of crops in years when RH is very high and
accompanied by cool nights during flowering. Cultivar differences with respect
to susceptibility are recognized, and ‘Pairi’ (‘Raspuri’) is highly susceptible.
C.P.A. Iyer and R.J. Schnell 78
Gupta (1976) has listed those cultivars that are most tolerant of this disease –
‘Neelum’, ‘Zardalu’, ‘Bangalora’, ‘Totapuri-Khurd’ and ‘Janardan Pasand’ –
and hence could be valuable in breeding programmes.
PEST RESISTANCE. Considerable variation is also known to occur among mango
cultivars with respect to their susceptibility to attack and injury by insect pests.
Although no resistant genotypes have been reported for the mango hopper
(Idiocerus spp.), the insect has been observed to avoid colonizing open plant
types where free movement of wind is possible, an observation that could be
useful in selection. Although complete resistance is not known to either fruit
fly (Bactrocera spp.) or seed weevil (Stenochetus mangiferae), variation in the
degree of susceptibility has been reported (Iyer, 1991). Rossetto et al. (2006)
observed that resistance to fruit fly is compatible with fruit quality and pro-
ductivity and advocated that resistance to fruit fly should be one of the objec-
tives of all mango breeding programmes. Their results also indicated that the
main factors for resistance of mangoes to fruit flies lie in the fruit peel and not
in the fruit pulp.
4.6 Methods of Breeding
Selection from open-pollinated seedlings
In India, almost all cultivars are selections that were made from naturally occur-
ring open-pollinated seedlings. All of the Florida cultivars were selected from
open-pollinated seedling progenies; none has come from a controlled breeding
programme. Among the 64 Florida cultivars evaluated in the parentage analysis
by Schnell et al. (2006), the genetic background was found to be based on as few
as four Indian cultivars and the polyembryonic cultivar ‘Turpentine’. Two Indian
cultivars, ‘Mulgoba’ and ‘Sandersha’, are in the background of most Florida
types with ‘Amini’, ‘Bombay’, ‘Cambodiana’, ‘Long’, ‘Julie’ and ‘Nam Doc Mai’
making lesser contributions. In the parentage analysis ‘Turpentine 10’ was iden-
tified as a most probable paternal parent for ‘Haden’. The polyembryonic seed-
ling races of Cuba and Florida were considered the same by Popenoe (1920) who
called them the West Indian race (commonly known as ‘Turpentine’ in Florida).
‘Haden’ was reported as the maternal parent for ten cultivars included in the
analysis, but based on the parentage analysis, 31 cultivars were found to have
‘Haden’ as one of the most likely parents. Likewise, the other important early
Florida selection ‘Brooks’ is the parent of seven cultivars. ‘Haden, ‘Brooks’ and
seedlings of ‘Haden’ and ‘Brooks’ have contributed disproportionately to the
Florida group. In Florida, modern selection and breeding programmes for
mango have focused on cultivars with exceptional production, red skin, disease
resistance and extended shelf life. Methodology for crop improvement consists
of collecting seeds from selected maternal parents with desired characteristics
and growing them in close proximity to desirable male parents. Seedlings are
screened by leaf aroma and horticultural traits, leading to a field population of
thousands of candidate seedlings (Campbell and Zill, 2006).
Breeding and Genetics 79
In the Canary Islands, Spain, breeding mainly involves selection of open-
pollinated seedlings of ‘Lippens’. Lavi et al. (2004) indicated that mother trees
should not be chosen entirely on the basis of their phenotypes and trees with
inferior performance could also be included since progeny performance is
quite unpredictable. They observed that most of the variance components of
the agriculturally interesting traits are non-additive (Lavi et al., 1998) and
most of these traits result from dominant and epistatic interactions.
The Israel mango-breeding programme therefore relies on open-pollination
involving many mango cultivars from various parts of the world and screen-
ing approximately 100 seedlings from each mother tree. The seedlings are
grown on their own roots in the nursery for about a year and then planted in
the field at spacings of 2 × 6 m. Fruiting occurs after 3–6 years and first selec-
tion is carried out based on field and laboratory data. Fruit characteristics at
this stage are good skin colour, fruit weight of 400–600 g and high fruit qual-
ity (good taste, absence of fibres and small seed). Where a long harvest sea-
son is desired both early and late harvest seedlings are selected and a general
idea about shelf life of these seedlings is obtained. The second selection is
carried out under commercial conditions by several experienced farmers
using grafted plants. Plants that successfully pass this stage are planted in
semi-commercial plots for a final assessment before recommendation to farm-
ers. The two selection stages are aimed at shortening the breeding programme
and minimizing both the false negatives (loss of interesting seedlings which
were not identified) and the false positives (wrong identification of interest-
ing seedlings which should actually be rejected). New mango cultivars have
also been selections made from open-pollinated seedlings. ‘Maya’ and ‘Nim-
rod’ are seedlings of the same mother tree (Oppenheimer, 1967); ‘Tahar’ is a
seedling of ‘Irwin’ (Slor and Gazit, 1982) and ‘Naomi’ is a seedling of ‘Palmer’
(Tomer et al., 1993). The other promising selections include ‘Shelly’ (late sea-
son), ‘Tango’ (early season), ‘Selection 20/1’ (large fruit with aborted seeds)
and ‘Selection 1/5’ (shiny red colour).
The South African mango-breeding programme also places a major
emphasis on open-pollinated seedling selection. The screening of seedlings,
besides undergoing tests for various plant and fruit characters, also includes
shipping quality tests in which fruits are packed in 4 kg crates and stored at
11°C for 28 days to simulate shipping conditions. After storage the fruits are
allowed to ripen at room temperature (25 ± 2°C) and screened thoroughly.
Promising seedlings are again field-tested with ‘Sabre’ as rootstock with a
row of 40 trees along with ten trees of commercial varieties. Two cultivars,
‘Joa’ (‘Palmer’ seedling) and ‘Chene’ (‘Kent’ seedling), were released from
the breeding programme in 1996. Another high yielding selection ‘A2-CD28’
(‘Fascell’ seedling) is a midseason clone with an attractive pink blush (Human
et al., 2006) and has been recommended for Plant Breeders’ Rights in 2005.
The other promising selections are ‘Osteen’ (‘Haden’ seedling) and ‘Neldica’
(‘Palmer’ seedling).
The 'R2E2' cultivar developed in Australia is a seedling progeny of the
Florida cultivar 'Kent', and is now the second most popular mango grown in
Australia.
C.P.A. Iyer and R.J. Schnell 80
Controlled pollination
Hand pollination
The traditional, cumbersome method involving the continued pollination of
flowers on a panicle over several days when the flowers are open has now
been replaced in India with more efficient methods. The current method
involves the pollination of a limited number of flowers per panicle (maxi-
mum of ten), utilizing a larger number of panicles since it is very rare that a
panicle bears more than one fruit to maturity. Using this method, fruit set as
high as 3.85% can be achieved compared to the 0.23–1.57% efficiency involv-
ing other methods (Mukherjee et al., 1968; Singh et al., 1980).
Caging
The enclosure of two desirable parents of synchronous flowering in a screen
house with pollinating insects provides a more practical method of hybridiza-
tion. This method has been used in Israel (Degani et al., 1993), Brazil (Pinto and
Byrne, 1993) and South Africa (Cilliers et al., 1996). A standardized caging
technique for mango breeding was previously used in India following the
discovery of self-incompatibility in some of the most popular commercial culti-
vars (Sharma and Singh, 1970). This procedure involves the planting of self-in-
compatible (female) and male parents in specially prepared breeding plots, and
are enclosed in an insect-proof cage into which freshly reared houseflies, or any
other suitable pollinator, are introduced to effect cross-pollination (Sharma
et al., 1972).
Polycross mating
A polycross is simply the use of a number of advanced selections or current
commercial clones planted in a design that maximizes the chance for
cross-pollination. The polycross design has been extensively used in
sugarcane breeding where small flower size and low numbers of seedlings
per cross make controlled pollinations difficult. At USDA Subtropical Horti-
culture Research Station (SHRS) in Miami the clones ‘Haden’, ‘Tommy
Atkins’, ‘Kent’, ‘Keitt’ and ‘Nam Doc Mai’ have been used to produce new
seedlings for selection. Five clones of each genotype were planted, with at
least one plant of each genotype next to all other genotypes. Over 1000 seed-
lings from known mother trees are planted as maternal half-sib families. The
pollen parent of superior selections is determined using microsatellite
markers.
4.7 Handling of Hybrid Populations and Selection
Criteria for initial selection
Primary selection from the hybrid progeny is based on: (i) precocity; (ii) fruit
size and shape; (iii) skin colour; (iv) fruit characteristics (high pulp to stone
ratio and freedom from fibre and physiological disorders); and (v) fruit qual-
ity. Following this preliminary evaluation, selected hybrids are retained for
Breeding and Genetics 81
further screening. It is important to graft the hybrids onto proper rootstocks
as early as possible, as grafted plants are precocious. At least ten grafted
plants of each selected hybrid are used in the final selection, which is based
on yield, regularity in bearing and response to diseases and pests, in addition
to other desirable fruit characters. At least 3 consecutive years’ performance
data should be collected before deciding on their suitability for release as
new cultivars.
Pre-selection
Trees have a long juvenile phase, and the development of pre-selection meth-
ods is important for discarding inferior seedlings at a very early stage, obvi-
ating the need for maintaining a large number of seedlings for long periods.
This can save time, land and labour. Leaf flavour has been reported to be
directly correlated with fruit flavour (Majumder et al., 1972; Whiley et al.,
1993). Emergence of new growth flushes, simultaneously with fruiting or
immediately after harvest, is indicative of regular bearing (Sharma et al.,
1972). A higher phloem to xylem ratio, associated with dwarfing, has been
used effectively as a pre-selection criterion. Genotypes in which the ratio
exceeds 1.0 are least vigorous, those with a ratio between 0.6 and 1.0 are of
medium vigour and those with a ratio of less than 0.6 are most vigorous
(Kurian and Iyer, 1992). In addition, higher levels of phenolics in the apical
bud is associated with reduced vigour and dwarfing (Iyer, 1991). Although
Majumder et al. (1981) indicated that low stomatal density is an indicator of
dwarfness this has not been confirmed by other workers (Iyer, 1991). Regular
bearing mango cultivars have low polyphenol oxidase (PPO) activity (cate-
cholase and cresolase) compared to alternate bearers (Sharma, 2003). Sharma
et al. (2000) observed that a strong positive correlation existed between the
incidence of floral malformation and both enzyme activity (catecholase and
cresolase) and phenolic content and speculated that PPO activity can be used
as a biochemical index for screening mango germplasm against malforma-
tion disease.
Potential for marker assisted selection (MAS)
More than 65 microsatellite markers have been developed for mango and
these are easily used to verify parentage using a software package such as
cervus (Marshall et al., 1998). When caging trees or using the polycross mat-
ing design it is possible to identify the male parent from a set of potential
male parents. This has been useful in cacao breeding where mistakes in pol-
lination have lead to the estimation of unreliable breeding values for parental
clones. The development of linkage maps and identification of quantitative
trait loci (QTL) for productivity and quality traits has led to a very successful
MAS in cacao (Schnell et al., 2007). This could serve as a model for future
mango breeding and selection efforts.
C.P.A. Iyer and R.J. Schnell 82
Molecular markers
Molecular markers can be used for estimating genetic relationships among
clones, for parentage analysis and for the development of a saturated linkage
map. Isozymes were the first markers to be used for fingerprinting mango
cultivars, to determine self- versus cross-pollination and to estimate genetic
relationships (Degani et al., 1990; Knight and Schnell, 1994). RAPD markers
were also used to fingerprint cultivars and estimate genetic relationships in
mango (Schnell et al., 1995). A group of ‘Haden’ seedlings and a random
group of seedlings were evaluated using 11 RAPD primers. This study sup-
ported the ‘Haden’ parentage of ‘Eldon’, ‘Lippens’, ‘Tommy Atkins’ and
‘Zill’; however, the parentage of ‘Glenn’ and ‘Osteen’ was questioned. Adato
et al. (1995) used DNA fingerprinting (DFP) to evaluate genetic relationships
between 26 mango cultivars and 14 rootstocks. They provided a pedigree
that further confirmed the relationship between many of the ‘Haden’ seed-
lings. Lopez-Valenzuela et al. (1997) used RAPD markers to estimate genetic
diversity among 15 rootstock cultivars using 13 markers, and identified a
specific RAPD band associated only with the polyembryonic types. Eiad-
thong et al. (1999) utilized anchored simple sequence repeat markers to anal-
yse 22 mango cultivars; they were able to distinguish genotypes, but were
unable to find markers unique to either monoembryonic or polyembryonic
types, or for the Thai cultivars selected for green harvest (crispy mango) from
the cultivars selected for ripe fruit production. Kashkush et al. (2001) utilized
amplified fragment length polymorphisms (AFLP) to estimate genetic rela-
tionships between 16 cultivars and seven rootstock cultivars. They also anal-
ysed 29 progeny from a cross of ‘Tommy-Atkins’ and ‘Keitt’ and produced a
crude linkage map that identified 13 of the 20 linkage groups.
Viruel et al. (2005) developed the first reported set of 16 microsatellite
markers for mango, of which 14 produced the expected one or two amplifi-
cation products per genotype. These 14 microsatellites were used to evaluate
28 mango genotypes that included 14 Florida cultivars. Discrimination of all
28 genotypes was possible and the average number of alleles per locus was
5.3. Previously known pedigree information for the ‘Haden’ family of man-
goes was confirmed and was in agreement with previously published RAPD
and DFP analyses (Adato et al., 1995; Schnell et al., 1995) with one exception.
Viruel’s clone of ‘Zill’ was not resolved as a seedling of ‘Haden’. Schnell et al.
(2005) developed a second set of 15 microsatellite markers and analysed 59
Florida cultivars and four related species. Two of the microsatellites were
monomorphic among the Florida cultivars; the other 13 had an average num-
ber of alleles per locus of 4.2 with polymorphism information content (PIC)
values varying from 0.21 to 0.63.
Schnell et al. (2006) used 25 microsatellite loci to estimate genetic
diversity among 203 unique mangoes (M. indica L.), two M. griffithii Hook. f.
and three M. odorata Griff. accessions maintained at the National Germplasm
Repository (NGR) and by Fairchild Tropical Botanic Garden (FTBG) in Miami,
Florida. The 25 microsatellite loci had an average of 6.96 alleles per locus
and an average PIC value of 0.552 for the M. indica population. The total
Breeding and Genetics 83
propagation error in the collection (i.e. plants that had been incorrectly
labelled or grafted) was estimated to be 6.13%. When compared by origin,
the Florida cultivars were more closely related to Indian than to South-east
Asian cultivars. Unbiased gene diversity (H
nb
) of 0.600 and 0.582 was found
for Indian and South-east Asian cultivars, respectively, and both were higher
than H
nb
among Florida cultivars (0.538). When compared by horticultural
type, H
nb
was higher among the polyembryonic types (0.596) than in the
monoembryonic types (0.571).
To date 63 microsatellite markers have been developed for mango
(Duval et al., 2005; Honsho et al., 2005; Schnell et al., 2005; Viruel et al., 2005).
This number is more than adequate for genetic diversity studies and for par-
entage analysis as has been demonstrated by Schnell et al. (2006); however, it
is not enough to develop a saturated linkage map for the 20 linkage groups
of mango. Developing an additional 200 microsatellite or single nucleotide
polymorphic markers is a major objective of the USDA Agriculture Research
Service (ARS) programme in Miami over the next 2 years. Three experimen-
tal populations have been developed and planted in the field as mapping
populations. The first population is an F
2
population derived from self-polli-
nation of ‘Tommy Atkins’ consisting of 168 seedlings that was planted in the
field in 1995. The second population is an F
2
population derived from self-
pollination of ‘Haden’. A total of 224 seedlings from a single isolated ‘Haden’
tree have been in the field for 3 years. Phenotypic data collection is in prog-
ress for both of these populations. The development of a saturated linkage
map and the identification of QTL for important traits are objectives for the
USDA-ARS programme in Miami for the next 5 years.
4.8 Minimizing Problems in Breeding
Heavy fruit drop
Heavy fruit drop ultimately results in few hybrid fruits, despite the large
number of flowers used for cross-pollination. While many recommendations
are available to minimize mango fruit drop with growth regulators, these
have not been very useful in breeding programmes where the number of
flowers remaining in a panicle is very low. Iyer and Subramanyam (1972)
suggested that embryo culture could be used to rescue hybrid embryos, and
Sahijram et al. (2005) developed in-vitro techniques to rescue immature mango
embryos from controlled crosses and recovered hybrid plants.
Long juvenile phase
Normally, mango seedlings require 3–10 years to flower, thereby prolonging
the breeding programme. Grafting individual hybrids on the proper
rootstocks at the earliest possible stage and growing them in a location
where climatic stress (particularly cold weather) prevails, induces precocious
C.P.A. Iyer and R.J. Schnell 84
flowering. Iyer (1991) has reported significant differences between seedlings
on their own roots and grafted plants of the same genotype with respect to
fruit size, quality and even colour in the early years. However, different
results have been reported in a recent study of the effect of rootstocks on the
performance of seedling scions with respect to ten horticultural traits (Lahav
et al., 1995). No difference of practical importance was found between the
original seedlings and their grafted duplicates.
Singh (1969) has suggested that young mango seedlings can be induced
to flower and fruit if they are grafted onto comparable shoots of a bearing
tree (a few days before flowering). The scions are defoliated and girdled.
Using this technique, it has been reported that the fruit characteristics of F
1

hybrids can be determined within 2 years and F
2
hybrids within 4 years, thus
eliminating at least 10 years from the period required to raise and evaluate F
2

populations (Singh, 1963).
Ethephon (Chacko et al., 1974) and paclobutrazol (Anonymous, 1984)
have flower-inducing properties in mango and could also be utilized for
shortening the juvenile phase. However, they must be used with caution
since chemical induction of flowering can alter fruit size and this could lead
to errors in judgement when making selections within the hybrid progeny.
Polyembryony
Seeds of polyembryonic mango cultivars characteristically contain several
nucellar embryos, and may also contain a zygotic embryo. While nucellar
seedlings are preferred as rootstocks for mango because of their uniformity,
the breeder, on the other hand, is generally interested in sexual seedlings for
the selection of improved rootstocks. Until recently, crosses involving poly-
embryonic cultivars as the maternal parents were generally not performed,
since reliable methods for identifying zygotic seedlings were not available.
The use of polymorphic enzyme systems (isozymes) (Degani et al., 1990,
1992) to identify zygotic seedlings (Schnell and Knight, 1992; Truscott, 1992;
Degani et al., 1993) is based on the fact that nucellar seedlings should have
the same isozyme alleles as the maternal parent. A variation at a locus coding
for an enzyme indicates that the plant has originated by sexual reproduction.
Zygotic seedlings arising from self-pollination are distinguished from nucel-
lar seedlings by being homozygous at one or more loci at which the female
parent is heterozygous. Statistically, when three or four heterozygous loci are
examined, up to 88 or 94%, respectively, of the selfed zygotic seedlings are
identifiable (Moore and Castle, 1988). Cross-pollination by another cultivar
of the same genotype is equivalent to self-pollination. Zygotic seedlings from
cross-pollination are distinguishable from those resulting from self-pollination
if they express an allele not carried by the female parent.
The frequency of occurrence of zygotic seedlings varies among the poly-
embryonic mango cultivars, i.e. 22% with ‘13-1’ (Degani et al., 1993), 20 and
24% with ‘Turpentine’ (Degani et al., 1993 and Schnell and Knight, 1992,
respectively), 2 and 4% with ‘Sabre’ (Truscott, 1992 and Schnell and Knight,
Breeding and Genetics 85
1992, respectively) and 36 and 64% with ‘Madu’ and ‘Golek’, respectively
(Schnell and Knight, 1992).
4.9 Achievements of Conventional Breeding
Despite the many problems associated with mango breeding for cultivar devel-
opment, many useful hybrids have been released. The earliest attempts were
probably made in the West Indies to combine the good qualities of the Indian
mango with the indigenous types by controlled pollination (Brooks, 1912).
India
Intervarietal hybridization in India has resulted in the release of many culti-
vars. The work at Sabour initially yielded two promising hybrids: ‘Mahmood
Bahar’ and ‘Probashanker’, both combinations of ‘Bombay’ and ‘Kalapady’
(Roy et al., 1956). Subsequently, four more hybrids have been developed.
These are: ‘Sundar Langra’ (‘Sardar Pasand’ × ‘Langra’) having ‘Langra’
quality and regular bearing habit; ‘Alfazli’ (‘Alphonso’ × ‘Fazli’) with ‘Fazli’
quality and early ripening; ‘Sabri’ (‘Gulabkhas’ × ‘Bombai’) having ‘Bombai’
fruit shape and colour of ‘Gulabkhas’ with regular bearing habit; and ‘Jawa-
har’ (‘Gulabkhas’ × ‘Mahmood Bahar’) having high pulp and early bearing
habit (Hoda and Ramkumar, 1993). Developed in Kodur, the hybrid ‘Swarna-
jehangir’, combining the high quality of ‘Jehangir’ and the attractive colour of
‘Chinnaswarnarekha’, is a prolific bearer and is the best of all hybrids devel-
oped at this centre. The other hybrids released from Kodur are ‘Neeludin’
(‘Neelum’ × ‘Himayuddin’), ‘Neelgoa’ (‘Neelum’ × ‘Yerra Mulgoa’) and
‘Neeleshan’ (‘Neelum’ × ‘Baneshan’).
Two excellent, regular-bearing hybrids, ‘Mallika’ and ‘Amrapali’, were
developed and released by the Indian Agricultural Research Institute (IARI),
New Delhi (Singh et al., 1972). ‘Mallika’ is a hybrid between ‘Neelum’ and
‘Dashehari’ with a high total soluble solids (TSS) content, a higher percentage of
pulp, fibreless flesh and a fruit size of about 300 g. ‘Amrapali’ (‘Dashehari’ × ‘Nee-
lum’) is precocious, distinctly dwarf and hence amenable to high-density
planting, a regular bearer with excellent quality and is also very rich in vitamin
A. Recently, two more cultivars, ‘Arunima’ (‘Amrapali’ × ‘Sensation’) and ‘Pusa
Surya’ (a selection from ‘Eldon’) have been released from the IARI. A promising
mango hybrid ‘Ambika’, a cross between ‘Amrapali’ and ‘Janardhan Pasand’,
having a yellow colour with red blush, firm flesh and scanty fibre was
released from the Central Institute of Sub-Tropical Horticulture, Lucknow.
Four hybrid cultivars were released from the Indian Institute of Horticul-
tural Research in Bangalore: ‘Arka Aruna’ (‘Banganapalli’ × ‘Alphonso’),
‘Arka Puneet’ (‘Alphonso’ × ‘Banganapalli’), ‘Arka Anmol’ (‘Alphonso’ ×
‘Janardhan Pasand’) and ‘Arka Neelkiran’ (‘Alphonso’ × ‘Neelum’). ‘Arka
Aruna’ is dwarf, and large fruited with a high percentage of pulp and high
TSS content. It is ideal for homesteads. ‘Arka Puneet’ is very similar to
C.P.A. Iyer and R.J. Schnell 86
‘Alphonso’ but free of ‘spongy tissue’, has a good shelf life and is not suscep-
tible to fruit fly attack. ‘Arka Anmol’ is a heavy bearer with good keeping
quality (Iyer and Subramanyam, 1993). ‘Arka Neelkiran’ is free of spongy
tissue and has excellent skin colour.
‘Ratna’ is a cross between ‘Alphonso’ and ‘Neelum’ that was carried out
at the Fruit Research Station, Vengurla, Maharashtra; it has a larger fruit size,
fruit quality similar to ‘Alphonso’ and is free of ‘spongy tissue’ (Salvi and
Gunjate, 1988). A parthenocarpic mango cultivar, ‘Sindhu’, has been devel-
oped at this station as a result of back-crossing ‘Ratna’ with ‘Alphonso’ (Gun-
jate and Burondkar, 1993).
Two hybrid cultivars were released from the Fruit Research Station in
Sangareddy, Andhra Pradesh. ‘Au-Rumani’ (‘Rumani’ × ‘Mulgoa’) is a regular
and prolific bearer with fibreless flesh. ‘Manjira’ (‘Rumani’ × ‘Neelum’) is a
dwarf, regular and prolific bearer with good quality fruits.
The Paria Research Station in Gujarat developed three mango hybrids,
‘Neelphonso’ (‘Neelum’ × ‘Alphonso’), ‘Neeleshan Gujarat’ (‘Neelum’ × ‘Bane-
shan’) and ‘Neeleshwar’ (‘Neelum’ × ‘Dashehari’). These hybrids are supe-
rior in TSS, total sugars and vitamin C, in addition to their dwarfing habit,
with respect to their parents (Sachan et al., 1988).
Other countries
USA
Mango hybridization was reported from Hawaii in the 1920s, but no out-
standing problem appears to have been addressed or solved (Pope, 1929). A
number of crosses have been reported in Florida (Young and Ledin, 1954;
Sturrock, 1969), but all of the Florida cultivars are chance seedlings and none
came from controlled pollinations.
Israel
There is an extensive breeding programme in Israel aimed at producing
higher yielding cultivars with good quality, attractive fruit and with longer
harvest periods. Several hundred seedlings from open and controlled polli-
nations have been evaluated, and 14 of them have been identified as being of
interest (Lavi et al., 1993). The rootstock breeding programme is aimed at
developing rootstocks resistant to or tolerant of soil stresses, i.e. calcareous
soils, saline irrigation water and heavy non-aerated soils that predominate in
the mango-growing regions of Israel. Several interesting mono embryonic and
polyembryonic rootstocks have been selected (Lavi et al., 1993), but none has
performed better than ‘13-1’, the currently preferred rootstock in Israel (Gazit
and Kadman, 1980).
Australia
A breeding programme to develop a new cultivar which retains the charac-
teristic flavour of ‘Kensington’, but with improved productivity, greater dis-
ease resistance, enhanced skin colour and better postharvest performance,
Breeding and Genetics 87
was initiated in Queensland, Australia. These features are found in many
Florida cultivars (i.e. ‘Irwin’, ‘Sensation’ and ‘Tommy Atkins’) which are
being used as maternal parents in crosses with ‘Kensington’ (Whiley et al.,
1993). Promising hybrids have been identified in crosses involving ‘Sensa-
tion’, for example ‘Calypso’™ (see Knight et al., Chapter 3, this volume).
‘Calypso’™ has increased shelf life, firmer fruit, extra blush for cosmetic
appeal, a higher flesh-to-seed ratio and consistent yields of high-quality fruit.
The Australian mango breeding programme was strengthened since 1994 by
launching a major effort involving various organizations located in different
agro-climatic zones in hybrid production, as well as regional testing.
Brazil
Breeding has been initiated in the tropical savannah of Brazil to develop cul-
tivars that are dwarf and with good quality fruit. Hybridizations have
involved local, Indian and Florida cultivars. ‘Amrapali’ and ‘Imperial’ were
good male parents to confer dwarfing in the progeny (Pinto and Byrne, 1993).
Out of 2088 seedlings in the field, 209 seedlings were selected in the first
year and 42 of these were later identified as promising, from which four have
been released as new cultivars (Pinto et al., 2004). These four are: ‘Alfa’ (‘Mal-
lika’ × ‘Van Dyke’), which is semi-dwarf, high yielding and regular bearing;
‘Beta’ (‘Amrapali’ × ‘Winter’), high yielding and moderately resistant to
anthracnose and Oidium; ‘Roxa’ (‘Amrapali’ × ‘Tommy Atkins’), with excel-
lent fruit quality; and ‘Lita’ (‘Amrapali’ × ‘Tommy Atkins’), high yielding
with excellent fruit quality.
South Africa
The South African breeding programme at the Citrus and Subtropical Fruit
Research Institute (CSFRI) is based on introductions, open-pollination and
mass selection. Four new cultivars have been released: ‘Heidi’, ‘Neldawn’,
‘Neldica’ and ‘Ceriese’. In addition, 12 promising selections have been iden-
tified for further evaluation (Marais, 1992).
4.10 Mutations
Somatic mutations
Asexual propagation enables the preservation of accumulated mutations
(macro and micro), which would normally be eliminated during sexual
propogation. In many fruit crops, bud mutations and chimeras occur rather
frequently and can provide an additional source of variability for selection.
However, such reported instances are relatively few in mango. Roy and
Visweswariya (1951) observed mutants of ‘Puthi’ in which the number of
palisade cell layers differed from the original cultivar. Naik (1948) observed
significant variation among trees of the same clone with respect to fruit shape,
size, colour and quality, which was ascribed to bud mutations. ‘Davis Haden’,
a sport of ‘Haden’, is larger than ‘Haden’ and its season of maturity is about
C.P.A. Iyer and R.J. Schnell 88
a month earlier (Young and Ledin, 1954). ‘Rosica’ from Peru, is a bud mutant
of ‘Rosado de lca’. Unlike its parent, ‘Rosica’ is high yielding and regular
bearing, and does not produce seedless fruits (Medina, 1977).
Oppenheimer (1956), after a survey of many orchards in India, reported
wide variability in the performance of trees of the same clone within a single
orchard. Mukherjee et al. (1983) conducted a survey of mangoes in eastern
India and identified some superior clones. Singh and Chadha (1981), in a
study of orchards of ‘Dashehari’, located four clones which were superior in
performance. Singh et al. (1985) isolated two high-yielding clones from
orchards of ‘Langra’. Within ‘Kensington’, strains have also been identified
that show improved resistance to bacterial black spot (Whiley et al., 1993).
Roy (1950) observed a mutant of ‘Alphonso’ with respect to fruit shape,
and suspected it to be a mericlinal chimera. Pandey (1998) has described
seven clones of ‘Alphonso’: ‘Alphonso Behat’ and ‘Alphonso Bihar’ from Bihar,
‘Alphonso Batli’, ‘Alphonso Black’ and ‘Alphonso Bombay’ from Maharashtra,
‘Alphonso Punjab’ from Punjab and ‘Alphonso White’ or ‘Bili Ishada’ from
the North Canara district of Karnataka. Rajput et al. (1996) assembled several
‘Dashehari’ variants and after 14 years of observation, reported that the clone
‘Dashehari 51’ was superior with respect to yield and regular bearing. Other
somatic mutants include: ‘Cardozo Mankurad’ with large fruits of attractive
colour and high yields from ‘Mankurad’ of Goa; dwarf selections from the
‘Rumani’ and ‘Bangalora’ (Ramaswamy, 1989); development of ‘Paiyur’, a
dwarf selection from ‘Neelum’ (Vijaya Kumar et al., 1991); ‘Rati Bangana-
palli’ and ‘Nuzuvid’ from ‘Banganapalli’ (Anonymous, 1999); and ‘MA-1’,
regular bearing and high yielding with resistance to ‘spongy tissue’ from
‘Alphonso’ (Mukunda, 2003).
In Thailand, Chaikiattiyos et al. (2000) selected clone ‘SKoo7’ (now known
as ‘Kaew Sisaket’) from 320 ‘Kaew’ plants; ‘SKoo7’ has higher yield and
superior quality. Jintanawongse et al. (1999) also made superior selections for
yield and fruit quality from ‘Nam Dok Mai’, ‘Khiew Sawoey’, ‘Rad’ and
‘Nang Klang Wau’ and DNA fingerprints of all these clones were made for
comparison with the parental clone.
For these studies, it is important to conduct a replicated cultivar evalua-
tion trial against standard commercial cultivars to establish that these varia-
tions are stable and not due to environmental responses. The use of genetic
markers should be explored to confirm that the new clones are genetically
distinct from the original cultivar.
Induced mutations
Mutation induction using ionizing radiation was attempted by Siddiqui et al.
(1966). Siddiqui (1985) irradiated dormant buds of ‘Langra’ with high doses
of rays, and grafted them onto 1-year-old seedlings. A bud graft exposed to
3.0 kR bore fruits which were heavier, larger and had a more cream-yellow
pulp than the control. This variability was stable over three seasons. Sharma
and Majumder (1988b) irradiated bud sticks, topworked them onto 10-year-old
Breeding and Genetics 89
seedlings, and found that dosages above 5 kR are lethal for mango and that
the lethal dose required for 50% mortality (LD
50
) lies between 2 and 4 kR.
Effective dosages of the chemical mutagens, ethane methyl sulfonate (EMS)
and N-nitroso methyl urea (NMU), were 1.5 and 0.05%, respectively. The
spectrum of mutations induced by physical and chemical mutagens was
observed to be more or less the same, indicating the high sensitivity of certain
loci. The mutants included dwarfness, changes in shape and serration of
leaves and in TSS content in ‘Dashehari’. As in other perennial crops, muta-
genesis techniques that can allow useful traits to be targeted, as well as
isolating mutated sectors from a chimera, are essential.
4.11 Breeding Potential of Wild Species
Bompard (1993; see Bompard, Chapter 2, this volume) has made a compre-
hensive study of the wild Mangifera species and enumerated their potential
use in breeding. Mangifera laurina, which has subglabrous and laxly flowered
panicles and is well adapted to areas with perpetual wet climates, is resistant
to anthracnose. Mangifera orophila from Malaysia and M. dongnaiensis from
Vietnam are both restricted to mountain forests 1000–1700 m above sea level
and their hybrids with mango could extend cultivation into temperate zones.
Mangifera magnifica is completely free of fibres; M. rufocostata and M.
swintonioides have an off-season bearing habit; M. pajang (endemic to Borneo)
and some strains of M. foetida have good quality fruits. Similarly ‘Wani’ from
Bali and Borneo, the best variety of M. caesia, has a distinctive taste. Mangifera
casturi from South Kalimantan is a prolific bearer with small, black, sweet
fruits having good potential. Mangifera altissima is reportedly resistant to
mango pests, such as hoppers, tip borers and seed borers (Angeles, 1991).
Sharma and Choudhury (1976) observed that wild Mangifera trees identified
in Tripura State (north-eastern India) were free of mango malformation. The
wild species could also contribute to higher productivity. Fairchild (1948)
observed that crosses between five-stamened mango and the Indian mango
(only one fertile stamen) could produce hybrids having better pollinating
quality. The interspecific compatibility of these species with M. indica must
be verified before they can be utilized in hybridization programmes (as sug-
gested by Bompard, 1993; Kostermans and Bompard, 1993; see Bompard,
Chapter 2, this volume).
4.12 Conclusions
Until recently, all mango cultivars arose as chance seedlings or as seedling
selections from known mother trees. Enthusiasm for controlled hybridiza-
tion by means of hand pollination waned because of the tedious nature of the
task and heavy fruit drop, resulting in only very few hybrids. This low hybrid
population was inadequate for selection and hence not many outstanding
hybrids were obtained. However, improvements in pollinating techniques
C.P.A. Iyer and R.J. Schnell 90
and more rapid screening of hybrid populations have enabled the release of
many hybrid mango cultivars of commercial value. Because of the world
market’s demand for mangoes with specific qualities, the synthesis of new
cultivars has become imperative. Rapid strides in molecular biology and in
other aspects of biotechnology have opened up new approaches in plant
breeding. The development of polymerase chain reaction (PCR)-based
genetic markers, specifically microsatellites, and their application to classical
breeding offer tremendous potential for mango improvement. The develop-
ment of a saturated linkage map and the identification of QTL for important
traits will allow the implementation of a MAS programme. The introduction
of specific genes for disease resistance from cultivars and wild species into
popular cultivars should soon be a reality. Without resorting to these new
technologies, mango breeding will continue to be a slow process.
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© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 97
5 Reproductive Physiology
T.L. Davenport
University of Florida, Florida, USA
5.1 Introduction 98
5.2 Phenology 99
5.3 Shoot Development 100
Vegetative shoots 102
Reproductive shoots 104
5.4 Flowering Mechanisms 105
Shoot initiation 105
Induction 106
Florigenic promoter (FP) or stimulus 108
Vegetative promoter (VP) 110
5.5 Environmental Influence on Vegetative and Reproductive Development 111
Temperature 111
Water relations 113
Effect of N on flowering 114
Photoperiod 116
5.6 Hormonal Influence on Flowering 116
Ethylene 116
Auxin 117
Cytokinins 118
Gibberellins 119
Plant growth retardants 121
5.7 Photoassimilate Influence on Flowering 123
5.8 Horticultural Manipulation of Flowering 123
5.9 Conceptual Flowering Models 124
Carbohydrate-regulated flowering models 124
Hormone-regulated flowering models 127
5.10 Floral Management 133
5.11 Floral Biology 134
Sex ratio 134
Environmental determinants of sex ratio 134
Physiological determinants of sex ratio 135
T.L. Davenport 98
Anthesis and dehiscence 136
Pollen 136
Pollination 137
5.12 Fruit Development 139
5.13 Stenospermocarpy 139
5.14 Fruit Set and Retention 140
Sex ratio 140
Mineral nutrients 141
Hormonal control 141
5.15 Alternate Bearing 145
5.16 Conclusions 145
5.1 Introduction
Flowering and fruit set are the most critical of all events occurring after estab-
lishment of a tree crop. Given favourable growth conditions, the timing and
intensity of flowering greatly determine when and how much fruit are pro-
duced. Many important details about flowering are becoming clearer, espe-
cially in herbaceous plants, at the physiological, biochemical and molecular
levels (see reviews by Searle, 1965; Zeevaart, 1976, 2006; Bernier et al., 1981,
1993; Halevy, 1985–1986; Bernier, 1988; Kinet, 1993; Boss et al., 2004; Komeda,
2004; Putterill et al., 2004; Corbesier and Coupland, 2005).
Cool temperatures in the subtropics stimulate mango flowering and age
of the last vegetative flush has an important bearing on its ability to flower in
marginally cool or warm temperatures of the tropics (van der Meulen et al.,
1971; Davenport, 2000, 2003). Consequently, mango flowering can be en-
hanced during its normal season or manipulated to occur at other times of
the year in the tropics. For example, potassium nitrate (KNO
3
) can stimulate
out-of-season flowering in mangoes in tropical latitudes (Barba, 1974; Núñez-
Elisea, 1985; Davenport, 1993; Protacio, 2000), although this treatment has
not always been dependable. Various aspects of mango flowering and/or
fruit set have been reviewed (Singh, 1958a, 1979; L.B. Singh, 1960, 1977;
Chacko, 1986, 1991; Chadha and Pal, 1986; Davenport, 1993, 2000, 2003; Dav-
enport and Núñez-Elisea, 1997; Singh et al., 2005), and M.J. Soule (1950) pub-
lished an extensive annotated bibliography of the older literature related to
mango reproduction.
Understanding mango flowering is essential to efficiently utilize man-
agement systems that extend the flowering and crop production seasons.
Recent studies of mango flowering have resulted in conceptual models that
help explain the physiological basis of flowering (Chacko, 1991; Cull, 1991;
Kulkarni, 1991, 2004; Whiley et al., 1991; Davenport and Núñez-Elisea, 1997;
Davenport, 2000, 2003). Control of flowering allows growers to harvest their
crops at the most profitable times. Increasing the season of availability
improves competitiveness in the international marketplace, and promotes
the most efficient use of resources as costs of inputs continue to rise.
This chapter addresses the physiology of mango flowering, early fruit set
and retention. Cultivar names and type of embryony (i.e. monoembryonic or
Reproductive Physiology 99
polyembryonic) are purposefully left out to focus on the physiological
aspects of reproduction regardless of whether they are tropically or subtrop-
ically adapted or from Indian or South-east Asian origin. Cultivars are
selected for their productivity in specific environments. Transfer of a cultivar
to a different environment often results in some alteration in performance. It
is reasonable to assume that the underlying mechanisms by which all culti-
vars respond to their environment within the framework of their genetic limi-
tations are similar. The concepts described herein, therefore, apply to all cultivars,
regardless of origin.
5.2 Phenology
Growth of mango and other tropical trees is not continuous (Nakasone et al.,
1955; Halle et al., 1978; Verheij, 1986; Davenport, 1993, 2000, 2003). Apical
buds spend most of the time in rest. Growth occurs as intermittent, ephem-
eral flushes of shoots from apical or lateral buds (Naik and Mohan Rao, 1942;
Singh, 1958a, b). Stems are quiescent or resting terminal vegetative structures
on branches from which shoot growth occurs. Shoots are elongating vegeta-
tive or reproductive structures that emerge from apical or lateral buds of
stems. Vegetative shoots develop a prescribed number of nodes during
growth before entering a resting state as a stem. Depending on environment,
periods of stem rest are generally short in young plants but usually last sev-
eral months between episodes of growth in mature trees. Vegetative growth
generally occurs up to three or four times a year on individual branches,
depending upon cultivar and growth conditions.
Development of the vegetative shoot from initiation of growth to full
elongation requires 3–6 weeks, depending on the cultivar and climatic condi-
tions (Whiley et al., 1991). During this period, 10–20 new leaves are generally
produced before returning to a resting state. These rhythmic episodes of
extension growth are recorded on each branch as segments consisting of
compressed internodes interspersed with long internodes, that is articulate
growth (Tomlinson and Gill, 1973). Davenport (1992, 2003, 2006) referred to
regions of compressed internodes as intercalations and the entire segment of
long internodes terminating in an intercalation as an intercalary unit. The
number of intercalations between each branching point indicates the number
of vegetative growth episodes or flushes that have occurred between each
flowering flush.
Flushes of vegetative growth occur on groups of stems borne on scaffold-
ing branches in isolated sections of tree canopy. Flushing stems are usually
connected at some common branch point within the tree limbs. Asynchro-
nous flushes of growth at various times in random portions of a tree canopy
may appear to be continuous growth but are simply flushes occurring in
various parts of the total canopy over time. Flowering flushes generally occur
after extended periods of stem rest in the low-latitude tropics or during cool
winter months in the high-latitude tropics and subtropics. Like vegetative
flushes, reproductive flushes are usually asynchronous in tropical climates
T.L. Davenport 100
(Verheij, 1986). In the subtropics, however, trees exposed to cold tempera-
tures (3–10°C) display synchronized flowering flushes throughout the tree
canopy approximately 1 month later. Subsequent vegetative flushes also tend
to be synchronous for one or two growth cycles depending upon the number
of retained fruit. Less intense, cool weather (10–18°C), however, results in
asynchronous reproductive flushes in responsive stems as is typical of trees
growing in the tropics. The timing of flowering flushes of cultivars in various
locations has been reviewed by L.B. Singh (1960), Chadha and Pal (1986) and
Pandey (1989). Variations in flowering patterns occur in all cultivars depend-
ing on their age and whether they are growing in dry or humid tropics or
subtropics (L.B. Singh, 1960).
5.3 Shoot Development
Flushes of vegetative extension growth of mango stems terminate with for-
mation of determinate panicles. Several weeks to a few months after separa-
tion of the last flower or fruit from these panicles are required for the central
axis of the panicle or rachis to dry and mechanically separate from the sup-
porting stem, depending on the longevity of attached fruit. Five to ten lateral
vegetative shoots typically develop from axillary buds located at the termi-
nal intercalation positioned in a compact whorl surrounding the panicle scar
of each stem (see Fig. 1 of Reece et al., 1949). These lateral shoots become the
branch points of stems. These branching shoots form 10–15 leaves before the
apical buds return to a resting state to establish them as individual stems. Ini-
tiation of these lateral vegetative shoots may occur 2–3 months after desicca-
tion of panicles which fail to set fruit. Fruit-bearing stems do not initiate new
lateral shoots until several months after separation of fruit and rachis from the
stem (Kulkarni and Rameshwar, 1989). Such delayed vegetative growth can
reduce the potential for new shoots to flower during the next flowering sea-
son (Singh and Khan, 1939; L.B. Singh, 1960, 1972; Monselise and Gold-
schmidt, 1982). The apical bud of stems is at rest for most of the year in mature
trees. Stems on centennial trees typically produce only one vegetative flush
during the year (N. Golez, personal communication, the Philippines, 1989).
The apical resting bud of each newly established lateral stem (intercalary
unit) is surrounded by a compact whorl of 10–12 leaves with short inter-
nodes (intercalation) (Fig. 5.1). Protective bud scales are green but may be
brown at the tips due to desiccation (Sen and Mallik, 1941; Mustard and
Lynch, 1946; Singh, 1958b; Ravishankar et al., 1979). Resting buds possess a
number of pre-formed nodes, each of which contains a leaf bract or leaf pri-
mordium and a lateral meristem (Fig. 5.2; see Figs 7–12 in Chaikiattiyos et al.,
1994). The outermost, proximally located dried leaf bracts (bud scales) pro-
tect the more distal interior leaf bracts, leaf primordia and lateral meristems
from mechanical damage and desiccation. Leaf bracts are vestigial non-
developed leaves. Scales abort upon evocation of new shoots. Proximally
located bracts in apical buds fail to further develop beyond some enlarge-
ment and also abort with elongation of shoots.
Reproductive Physiology 101
If apical buds are initiated during vegetatively inductive conditions,
bracts develop as small leaves and the leaf primordia develop as the full-
sized leaves of vegetative shoots. Additional leaves result from nodes formed
by renewed activity of the apical meristem. The number of leaves (nodes) is
dependent upon the mean temperature during initiation, and increases as
temperatures rise (Whiley et al., 1989). The lateral meristems of the apical
bud develop as axillary buds at the base of petioles in the elongating vegeta-
tive shoot, each bearing protective bracts, leaf primordia and lateral mer-
istems (Fig. 5.3; see Fig. 1 of Reece et al., 1949).
In contrast, if shoot growth is initiated under floral inductive conditions,
the leaf bracts and primordia fail to fully develop, but the lateral meristems
Fig. 5.1. Apical bud of resting mango stem.
Leaf primordia (bracts)
Apical dome
(includes meristem)
Lateral meristematic
primordia
Fig. 5.2. Stylized cross-section of apical bud showing positions of apical meristem,
lateral meristems and leaf primordia.
T.L. Davenport 102
begin to elongate and branch at each node forming secondary, tertiary and
quaternary lateral meristems. Each branch point in the lateral inflorescence
from the panicle axis to the floral pedicels bears a floral bract (i.e. partially
developed vestigial leaf primordium) (Fig. 5.4). The distal half of the panicle
structure is derived from newly formed nodes laid down by cell divisions in
the apical meristem prior to returning to a resting state. Mixed shoots, bear-
ing both leaves and inflorescences at each node, result from development of
both the primary leaf primordia and the lateral meristems, which form the
inflorescences in the same nodes as leaves.
Vegetative shoot induction, thus, involves stimulating development of
leaf primordia from resting buds while repressing development of lateral
meristems. Leaf primordia then follow a predetermined cascade of genetic
signals resulting in leaf development at each node. Because all shoots emerge
from resting buds, a vegetatively induced event does not involve simply
inhibition of flowering. The putative inductive signal directing differentia-
tion of leaf primordia onto leaves upon initiation is termed a vegetative pro-
moter (VP) rather than a floral inhibitor.
Shoots bearing only inflorescences (generative shoots) result from induc-
tive development of lateral meristems and suppression of leaf primordial
development. A predetermined cascade of flowering gene signals is activated
in lateral meristems resulting in lateral cymose inflorescences terminating
with flowers. A distinct florigenic promoter (FP) may be responsible for spe-
cific activation of the lateral meristems of mango. Mixed shoot induction
results in combined development of leaf primordia and lateral meristems.
Vegetative shoots
Vegetative shoots bear only leaves (Fig. 5.5). The anatomy of mango vege-
tative shoot development has been described (Singh, 1958b; Chaikiattiyos
Stem
Fig. 5.3. Axillary bud of resting mango stem. Leaf petioles (arrows).
Reproductive Physiology 103
et al., 1994). Vegetative shoots may arise either from axillary buds, if no apical
bud exists due to flowering in the previous flush, or from the apical bud
when present. The latter is considered extension growth or addition of an
intercalary unit on the existing stem, but the developmental events during
shoot formation from either apical or lateral buds are basically the same.
Cells in the leaf primordia of initiating buds begin to form individual leaves
in the proximal portion of the vegetative shoot. Soon thereafter, the apical
meristem activates to form more nodes bearing leaf primordia and lateral
meristems. These newly formed leaf primordia develop as the distal portion
of the vegetative shoot if environmental conditions remain vegetatively
inductive (Núñez-Elisea et al., 1996). Newly elongating vegetative shoots are
green in most cultivars but may be bronze or red in others. Fully expanded

Pedicel

Pedicel
1° Bract
2° Bract
3° Bract
2° Bract
1° Bract

Pedicel

Pedicel
1

Pedicel
1° Bract




º

Axis
Axis
Fig. 5.4. Diagram and photos of mango inflorescence depicting the panicle axis and
primary (1°), secondary (2°) and succeeding levels of pedicel and cymose floral archi-
tecture. Vestigial leaf promorida (floral bracts) are depicted at the base of each level of
pedicel architecture.
T.L. Davenport 104
leaves are a shade of red, depending upon cultivar and cultural conditions
and are thin and limp from lack of lignification. The apical buds of vegetative
shoots generally become quiescent before completion of the limp, red-leaf
stage (Núñez-Elisea and Davenport, 1995). Internodes are compressed at the
apex, and leaf development is arrested thereby forming a bud with protec-
tive outer scales, inner leaf primordia, lateral meristems and the apical mer-
istem. Fully expanded leaves become light green and stiff as they become
lignified and suberized. Vegetative shoots are mature when leaves become
dark green, which occurs when they are c.2 or 3 months old.
Reproductive shoots
Two types of reproductive shoots typically occur in mango. Generative
shoots display only flowers and have floral bracts or non-developed leaves
at the base of each lateral inflorescence (Fig. 5.5). Terminal inflorescences,
i.e. panicles or thyrsoids (Weberling, 1989), develop from dormant apical
buds. The anatomy of panicle development has been described (Juliano
and Cuevas, 1932; Musahib-ud-din, 1946; Mustard and Lynch, 1946; Singh,
VEGETATIVE GENERATIVE MIXED CHIMERIC V/F TRANSITION F/V TRANSITION
GENERATIVE CHIMERIC V/F TRANSITION VEGETATIVE MIXED F/V TRANSITION
Fig. 5.5. Stylized diagrams and photomontage of shoot types found in mango. Transition
shoots shift from vegetative to floral (V/F) or floral to vegetative (F/V). Arrow ( )
represents individual leaves; floral diagram ( ) represents lateral inflorescences.
Reproductive Physiology 105
1958b; L.B. Singh, 1960; Sturrock, 1966; Ravishankar et al., 1979; Scholefield,
1982; Scholefield et al., 1986). The complexes of primary to quaternary branch-
ing lateral structures of the inflorescence each terminate with three cymose
flowers. The terminal flower opens first, followed by two subtending lateral
flowers. These complexes form the lateral inflorescence structures emerging
from the central axis of the panicle. The central axis extension also terminates
in a similar fashion. Morphological stages of floral buds and panicle develop-
ment were described by Shu (1981) and Oosthuyse (1991a). Reece et al. (1949)
described the development of inflorescences initiated in lateral buds when
the terminal bud is missing. There are more nodes in dormant apical buds
and their bracts are more developed than in axillary buds; however, floral
evocation is indistinguishable.
Generative shoot development in apical buds initially involves swelling
of the lateral meristems and their bud scales. Each axillary meristem devel-
ops as an inflorescence on a primary peduncle. The apical meristem then
forms new lateral meristems and leaf primordia for the distal portion of pan-
icle development if floral inductive conditions persist (Núñez-Elisea et al.,
1996). Panicles may be open or compact, depending upon internode elonga-
tion, which is cultivar dependent (L.B. Singh, 1960), but the architecture gen-
erally conforms to that in Fig. 5.5. Mixed shoots develop under weak floral
inductive conditions (i.e. in the low-latitude tropics). Both leaves and pri-
mary pedunculate inflorescences develop from the same nodes (Fig. 5.5).
Leaf primordia and lateral meristems develop as leaf and floral structures,
respectively.
5.4 Flowering Mechanisms
Mango stems undergo varying periods of rest between episodes of growth,
depending on tree age and environmental influences. Resting mango buds
must, therefore, respond to two distinctly different signals for shoots to occur.
The first signal initiates growth of the shoot and the second determines if it
will be vegetative or reproductive. The signals that regulate initiation of
shoot growth in resting buds differ from the inductive signals that regulate
shoot type.
Shoot initiation
Initiation is the onset of shoot development, regardless of the type of shoot
evoked. It involves cell division and elongation of cells in leaf primordia
(vegetative shoots), lateral meristems (generative shoots) or both (mixed
shoots) in the nodes of the resting buds, and is followed by cell divisions in
the apical meristem to form more nodes. Shoot initiation is stimulated by
pruning, defoliation and irrigation during dry conditions, or transition from
the dry to rainy season in the tropics. Application of nitrogen (N)-containing
fertilizers, exposure to ethylene, or a shift from cool to warm temperatures
T.L. Davenport 106
also stimulates shoot initiation. Reece et al. (1946, 1949), Mustard and Lynch
(1946), Núñez-Elisea and Davenport (1992b), Núñez-Elisea et al. (1996) and
Davenport et al. (2006a) observed that the vegetative or reproductive fate of
mango buds remains undetermined until after shoot growth is initiated.
Reece et al. (1949) proposed that a putative signal that triggers initiation of
shoot development is separate and different from the inductive signal, which
determines the fate of the shoot. Removal of apical buds by pruning stimu-
lates initiation of axillary shoots (Singh and Singh, 1956; Núñez-Elisea and
Davenport, 1992b; Núñez-Elisea et al., 1996; Davenport et al., 2006a). Defolia-
tion of the apical whorl of five to ten leaves also stimulates shoot initiation
in dormant apical buds (Núñez-Elisea et al., 1991; Núñez-Elisea and Daven-
port, 1995). The fate of shoots that emerge in response to these initiation stim-
uli, however, is determined by other factors that are prevalent at the time of
initiation. Tip pruning, for example, during warm summer months results in
initiation of vegetative shoots from axillary buds, whereas pruning during
cool winter months usually results in initiation of axillary inflorescences.
Induction
Induction in mango is the temporary commitment of buds to evoke a par-
ticular developmental pathway (i.e. vegetative shoot, generative shoot or
mixed shoot) when growth is initiated. Initiation of herbaceous plant flower-
ing refers to the onset of floral bud growth in actively growing vegetative
shoots after the floral inductive event (Bernier et al., 1981, 1993; Halevy, 1985–
1986; Bernier, 1988; Huala and Sussex, 1993; Kinet, 1993). The inductive sig-
nal is formed in leaves, but the responsive buds are in continuous vegetative
growth at the time of floral induction in herbaceous plants and floral initia-
tion follows; whereas mango buds are in rest. Although the mango bud
must be initiated to grow, that growth is induced according to forces already
present.
Whereas the floral inductive signal in mango may be present prior to
bud initiation, it must be present at the time of initiation for flowering to
occur (Kulkarni, 1988a; Núñez-Elisea and Davenport, 1995; Núñez-Elisea et al.,
1996; Davenport and Núñez-Elisea, 1997; Davenport et al., 2006a). The induc-
tive signal can be shifted from floral (F) to vegetative (V) or vegetative to
floral, forming F/V or V/F transition shoots, by altering temperatures dur-
ing early shoot development (Batten and McConchie, 1995; Núñez-Elisea
et al., 1996) (Fig. 5.5). This shift in morphogenic responses during shoot devel-
opment demonstrates the plasticity and temporal nature of induction, indi-
cating that cells of the apical meristem do not become irreversibly determined
under inductive conditions. These results demonstrate that, rather than being
irreversibly committed to a vegetative or reproductive fate at the onset of
shoot initiation, the mango apical meristem provides progenitor cells, some
of which differentiate into specific target cells at each node in the apex. The
apical meristem, therefore, may not be directly involved in the flowering
process.
Reproductive Physiology 107
Target cells within leaf primordia and lateral meristems are competent to
respond to inductive signals; for example when initiated to grow under veg-
etatively inductive conditions, individual leaf primordia develop as leaves
and subtending lateral meristems associated with each developing leaf
develop as dormant axillary buds with protective bracts. These axillary buds
may develop in subsequent flushes as vegetative shoots when initiated in
vegetatively inductive conditions or as axillary inflorescences under floral
inductive conditions. Under strongly floral-inductive conditions, leaf pri-
mordia fail to develop beyond the bract stage, become dormant, and lateral
meristems develop. Each lateral meristem forms nodes consisting of leaf pri-
mordia and meristems that are influenced by the putative floral-inductive
stimulus, which suppresses development of newly formed leaf primordia.
Subsequently formed meristems form pedunculate structures that terminate
in cymose inflorescences borne on each tertiary peduncle (Fig. 5.4). Forma-
tion of the primary, secondary, tertiary and quaternary peduncles, as well as
pedicels of inflorescences are always accompanied by a subtending, aborted
bract or vestigial leaf at each node (Fig. 5.4). Such development is attributed
to a sequence of gene expression (Coen et al., 1990; Coen and Meyerowitz,
1991; Weigel et al., 1992; Coen and Carpenter, 1993; Lumsden, 1993; Yanofsky,
1995). Shoot initiation during weakly floral-inductive conditions activates
growth of leaf primordia to develop leaves and the lateral meristems to pro-
duce peduncles bearing lateral inflorescences in each node of mixed shoots.
The bases of each pedicel branch within each lateral inflorescence also bear a
vestigial leaf.
Upon termination of cell divisions in the apical meristem at the end of a
flushing period, no more nodes are formed. The apical bud of vegetative
shoots becomes quiescent, and the resting leaf primordia, bracts and lateral
meristems are poised to resume growth at a later date. When reproductive or
mixed shoots become quiescent, the lateral meristems ultimately develop
determinant cymose inflorescences. The most distally located meristem is
possibly the determinant extension of the central axis forming the terminal
cymose floral group.
Chimeric shoots (Fig. 5.5) can occur in mango trees when shoot initiation
occurs during floral inductive conditions. They display inflorescences on one
side of the longitudinally bisected shoot and leaves on the other. The shoot
axis is red on the floral side of red fruiting cultivars (typical of panicles) and
green on the vegetative side (typical of vegetative shoots). This difference
in the two sides extends to the apical bud, which bears an undeveloped
inflorescence on the floral side and leaf bracts on the vegetative side. The
explanation for this spatial differentiation is that target nodes on each side of
the apical bud respond to the different inductive signals at the same time.
The apical meristem is not implicated except to form more nodes for the lat-
eral inductive responses on each side in the second portion of growth. Differ-
ences in inductive signals on each side of an existing shoot probably cause
the differential response. This phenomenon indicates that the fate of nodes
on each side of the shoot cannot be attributed to a single mother cell in the
apical meristem. The inductive response must involve cells formed in later
T.L. Davenport 108
cell divisions and would be determined by their location within nodes of
the bud.
Florigenic promoter (FP) or stimulus
Early flowering work provided evidence for the presence of a graft transmis-
sible floral stimulus (i.e. florigen) that was induced in leaves and was trans-
located to buds to stimulate floral development (Chailakhyan, 1936; Zeevaart
and Boyer, 1987). Florigen was functionally conserved across plant species
(Lang, 1965, 1984; Zeevaart, 1976; Lang et al., 1977). Floral induction in most
plants involves sensing of some environmental cue (i.e. daylength, water
stress or vernalizing temperature) in some organ (e.g. leaves). A putative flo-
ral stimulus or alteration in the ratio of florigenic to anti-florigenic compo-
nents may be translocated to target cells in meristems (Bernier et al., 1981).
Photoassimilate movement from leaves in phloem facilitates its transport to
buds where it can interact to initiate flowering (King and Zeevaart, 1973).
Until recently, a floral stimulus could not be identified. Alternative hypoth-
eses were proposed that nutrient diversion to the meristems could be
involved (Sachs and Hackett, 1983) or that floral induction might be con-
trolled by multiple factors, including the putative floral stimulus, photoas-
similates and phytohormones (Bernier et al., 1993).
Molecular biology of flowering in the facultative, long-day, model plant,
Arabidopsis thaliana (reviewed in Zeevaart, 2006 and Aksenova et al., 2006),
has provided insight into the nature of the floral stimulus (FP). A network of
four interacting genetic signalling pathways may result in flowering in
response to photoperiodic, vernalization, gibberellin and autonomous envi-
ronmental cues (Perilleux et al., 1994; Mouradov et al., 2002; Perilleux and
Bernier, 2002; Boss et al., 2004; Komeda, 2004; Putterill et al., 2004; Corbesier
and Coupland, 2005). The photoperiodic pathway involves activation of
the CONSTANS (CO) gene that encodes a zinc-finger protein, which in
turn induces expression of the FLOWERING LOCUS T (FT) gene in the
phloem tissue of leaves. FT is the terminal, integrating gene of the four path-
ways regulating flowering in Arabidopsis. Its transcribed mRNA was initially
thought to be the FP that is transported in phloem to buds (Huang et al.,
2005); however, evidence indicates that the translated protein product of FT
is translocated to Arabidopsis buds (Corbesier et al., 2007). Analogous proteins
encoded by Hd3a, an ortholog of FT in rice (Tamaki et al., 2007), and the aspen
ortholog, PtFT1, which along with CO regulates the timing of flowering and
growth cessation of Populus trichocarpa (Bohlenius et al., 2006), appear to be
the FP. In the buds, the protein product of FT is thought to combine with the
bZIP transcription factor (FD) protein to activate transcription of floral iden-
tity genes (i.e. APETALA1) to begin floral expression (Abe et al., 2005; Wigge
et al., 2005). Similar mechanisms are likely to exist in mango.
Zhang et al. (2005) and Davenport et al. (2006b) isolated a CONSTANS-
like gene (MiCOL) from mango leaf DNA. CO is a circadian expression gene
interacting with the photoperiodic pathway in Arabidopsis (Putterill et al.,
Reproductive Physiology 109
2004), and is central to activation of the FT gene in Arabidopsis during long
days. Its role in mango flowering is unclear. The mango ortholog has 79%,
76% and 62% homology with two apple CO genes, MdCOL2 and MdCOL1,
and the Arabidopsis CO gene (AtCO), respectively. Isolation of the FT or
homologous gene responsible for synthesis of the FP has been unsuccessful.
Studies with mango indicate that a FP is synthesized in leaves during
exposure to cool, floral-inductive temperatures and moves to buds to induce
flowering (Reece et al., 1946, 1949; Singh and Singh, 1956; L.B. Singh, 1959,
1962, 1977; R.N. Singh, 1961; Sen et al., 1972; Núñez-Elisea and Davenport,
1989, 1992b; Davenport and Núñez-Elisea, 1990; Davenport et al., 1995,
2006a). Unlike receptor sites in buds of Thlaspi arvense (Metzger, 1988) and
other plants requiring vernalization for floral induction (Zeevaart, 1976;
Bernier et al., 1981), mango leaves appear to be where the putative floral stim-
ulus is produced. Complete defoliation of girdled branches during inductive
conditions results in vegetative shoots instead of generative shoots (Reece
et al., 1949; Sen et al., 1972; Núñez-Elisea and Davenport, 1989, 1992b; Núñez-
Elisea et al., 1996; Davenport et al., 2006a). It appears to be transported over
long distances from leafy branches to defoliated branches (Sen et al., 1972;
Núñez-Elisea et al., 1996).
The putative, temperature-regulated FP is short-lived in situ (Núñez-
Elisea and Davenport, 1989, 1992b; Davenport et al., 1995; Núñez-Elisea et al.,
1996). Leafless cuttings from trees during cool, floral inductive conditions
produce inflorescences when stimulated to grow within 7 days of transfer to
warm, non-inductive conditions; the influence of the removed leaves lasts
for 13 days when cuttings are stored at cool temperatures (Davenport et al.,
2001a). The same cuttings produce only vegetative shoots in both storage
conditions after the initial loss of reproductive shoot production. There are
more leaves on mango stems than are necessary for floral induction in cool
temperatures. Stems bearing as little as one-quarter of a cross-sectioned leaf
induce 95% generative shoots (Davenport et al., 2006a); the remaining shoots
are vegetative. Half of a leaf or more resulted in 100% generative shoots.
Thus, the limiting amount of leaf necessary for floral induction is less than a
quarter of a leaf per stem. Davenport et al. (2006a) demonstrated the quanti-
tative movement of mango FP from half to five leaves on a donor stem to five
leafless receiver stems located as far as 100 cm from the donor stem in isolated
branches during exposure to cool, floral inductive temperatures. The FP moves
with photoassimilates in phloem from donor leaves to buds in the receiver
stems.
The mango floral stimulus is graft transmissible (L.B. Singh, 1959, 1962;
Kulkarni, 1986, 1988b, 1991). Flowering of seedling stems is stimulated by
grafting onto mature trees or by grafting mature stems onto juvenile plants
(L.B. Singh, 1959, 1962). Some mango cultivars selected in the tropics can
flower at higher temperatures than others and are not restricted to winter
flowering (Kulkarni, 1991). Transfer of the FP from tropical to subtropical selec-
tions was accomplished using reciprocal grafts between the two cultivar types
(Kulkarni, 1986, 1988b, 1991). Subtropical cultivars that seldom flower in warm
temperatures flower in the ‘off’ season using these techniques. Three conditions
T.L. Davenport 110
were essential for summer flowering to occur in the low-temperature-requiring
cultivars (receptors) when grafted to the summer flowering type (donors): (i)
the summer-flowering donor cultivar stocks or scions were in a flowering
cycle; (ii) buds on the receptor scions or stocks of grafted plants had initi-
ated shoot growth during this cycle; and (iii) receptor stocks or scions had
been completely defoliated for transfer and/or expression of the floral stimu-
lus. The presence of any leaves on the receptor plants resulted in vegetative
shoots.
Girdling experiments to isolate treated mango branches from the rest of
the tree suggest that the FP is translocated via phloem to apical buds (King
and Zeevaart, 1973; Bernier et al., 1981; Núñez-Elisea and Davenport, 1989,
1992b; Núñez-Elisea et al., 1996; Davenport et al., 2006a). Shading experiments
to reduce photosynthate loading into the phloem also support this (Kulkarni,
1991). Reduced flowering responses were observed in isolated leafy branches
that were provided with 90% and complete shading, which stopped photosyn-
thate production entirely, mimicked defoliation during cool, floral inductive
conditions, resulting in a vegetative growth response (R. Núñez-Elisea, T.L.
Davenport and B. Schaffer, Florida, 1991, unpublished results).
Vegetative promoter (VP)
An independently regulated VP probably contributes to induction of vegeta-
tive shoots as opposed to a floral inhibitor or expression of a default vegeta-
tive status in the absence of sufficient FP at the time of shoot initiation.
Grafting studies (L.B. Singh, 1959, 1962; Kulkarni, 1986, 1988b, 1991, 2004)
demonstrated that complete removal of leaves from receptor stems is required
to express flowering of those receptors when they are grafted to flowering
donor stems. Kulkarni (1986, 1988b, 1991, 2004) considered that a putative
floral inhibitor in leaves of the non-induced receptor stems might antagonize
the influence of the floral stimulus from donor leaves. Others have noted a
relationship between leaf age and the ability of shoots to be reproductive
(Singh et al., 1962a; Scholefield et al., 1986). KNO
3
-stimulated early flowering
in the tropics is successful only on stems that are at least 4 (Davenport, 2003)
to 7 months old (Astudillo and Bondad, 1978; Bondad and Apostol, 1979;
Núñez-Elisea, 1985). Young stems often produce vegetative shoots when ini-
tiated under conditions that are floral inductive for more mature stems
(Núñez-Elisea and Davenport, 1995; Davenport, 2003). The putative VP
appears to be most active in leaves of young stems and slowly dissipates
over time to allow expression of the FP when shoots are initiated to grow in
warm conditions.
The VP may be a gibberellin or closely associated with the gibberellin
synthesis pathway as indicated by enhanced flowering responses of trees to
plant growth retardants. Mangoes growing in wet and humid, low-latitude
tropics tend to produce frequent vegetative flushes and flower sporadically,
perhaps due to higher levels of the VP in the young stems combined with
low levels of the putative FP when shoot initiation occurs. Paclobutrazol
Reproductive Physiology 111
(PBZ) reduces the time in rest necessary to allow floral induction during
warm temperature conditions by c.1 month (Davenport, 2003), thus increas-
ing the potential to produce reproductive shoots in younger stems when ini-
tiated to grow. PBZ and uniconazole, triazole compounds that inhibit kaurene
oxidase in the gibberellin-synthesis pathway (Dalziel and Lawrence, 1984;
Rademacher, 1991), stimulate production of flowering shoots during weakly
inductive conditions (Burondkar and Gunjate, 1991, 1993; Tongumpai et al.,
1991a; Voon et al., 1991; Nartvaranant et al., 2000; Yeshitela et al., 2004a).
Application of PBZ to mango trees bearing 1-month-old stems produced
inflorescences when bud break was initiated 3 months later by foliar applica-
tion of KNO
3
(Davenport, 2003).
Vegetative or reproductive induction at the time of shoot initiation is
governed by the ratio of the putative floral promotive to inhibitory compo-
nents (Lang et al., 1977; Lang, 1984; Kulkarni, 1988a; see Bernier et al., 1981 for
additional references). The mango floral inhibitor should be viewed as an
age-dependent VP. The presence of an age-regulated VP in mango leaves,
which moves with the temperature-regulated FP and photoassimilates in
phloem, may explain the induction of specific receptors by this promoter in
targeted leaf primordia to cause development of leaves in vegetative or
mixed shoots. A gradual decrease in the level or influence of the VP may
cause vegetative shoots to develop when initiation occurs on 2-month-old
stems, and generative or mixed shoots when initiation occurs in stems from
4- to 7-month-old stems, given the constantly warm daily temperatures
maintaining a low level of FP in both situations.
5.5 Environmental Influence on Vegetative and Reproductive
Development
The effects of temperature and water relations on determinating vegetative
and reproductive growth of mango have been addressed (Davenport and
Núñez-Elisea, 1997; Davenport, 2000; Kulkarni, 2004; Bangerth, 2006). This
section focuses on the impacts of temperature, plant water relations, mineral
nutrition and photoperiod on shoot initiation and induction.
Temperature
The developmental fate of mango buds is strongly influenced by tempera-
ture (Davenport and Núñez-Elisea, 1997). Cool night temperatures < 15°C in
combination with day temperatures < 20°C typically induce flowering if
shoot initiation occurs when plants are exposed to these conditions (Ou,
1980, 1982; Wolstenholme and Mullins, 1982a, b; Shu and Sheen, 1987; Whi-
ley et al., 1988, 1989, 1991; Núñez-Elisea et al., 1993; Núñez-Elisea, 1994;
Núñez-Elisea and Davenport, 1994a, b). The physiological and molecular
basis for temperature perception in leaves with respect to floral induction is
not understood (Samach and Wigge, 2005). Whiley et al. (1988, 1989, 1991)
T.L. Davenport 112
described the vegetative growth and flowering responses of several mo-
noembryonic and polyembryonic cultivars to four temperature regimes rang-
ing from vegetatively inductive (30°C day/25°C night) to floral inductive (15°C
day/10°C night). The effect of temperature on marcotted, container-grown
plants that were tip pruned or defoliated in order to stimulate shoot initia-
tion was also studied (Davenport, 1987; Núñez-Elisea et al., 1991, 1993, 1996;
Núñez-Elisea and Davenport, 1994b). Mango trees develop vegetative shoots
when shoot initiation occurs in warm temperatures (30°C day/25°C night),
whereas inflorescences develop when shoots initiate growth in cool tempera-
ture conditions (18°C day/10°C night; or 15°C day/10°C night) (Whiley et al.
1989; Núñez-Elisea and Davenport, 1991b, 1995; Núñez-Elisea et al., 1993,
1996; Batten and McConchie, 1995). Bangerth et al. (2004) reported changes in
the major phytohormones in stems of containerized mango trees during
exposure to cool, floral inductive temperatures. The minimum leaf age and
time of exposure to a low temperature regime (18°C day/10°C night) required
by stems for floral induction was examined (Núñez-Elisea and Davenport,
1995). Leaves are competent to respond to cool temperatures at 7 weeks,
forming a small percentage of generative shoots. As they age, higher propor-
tions of generative shoots are induced and warmer temperatures can stimu-
late floral induction. The response to temperature is moderated by age of the
previous flush. Stems that are 4–5 months beyond the limp, red-leaf stage of
development will be induced to form generative shoots if initiated to grow
at 25–30°C (Davenport, 2003).
Whiley et al. (1988, 1989, 1991) observed that at least 17 weeks are required
for initiation of reproductive shoots on non-clipped stems of trees maintained
at 15°C day/10°C night. In similar experiments with different cultivars with-
out previous clipping of distal leaves to stimulate initiation, inflorescences
were observed after 5 weeks at 15°C day/10°C night (Chaikiattiyos et al.,
1994). Although inductive conditions were present in each of these studies,
shoot initiation was delayed by the presence of distal leaves. The earlier ini-
tiation of inflorescence development in tip-pruned or tip-defoliated stems
compared to intact ones demonstrates that the floral stimulus may be pres-
ent, but the buds are not induced until initiation occurs. It demonstrates the
importance of stimulating initiation of stems by tip defoliation or pruning
at the onset of incubation in controlled environment conditions so that the
inductive response can be observed within a reasonable length of time.
The variable delays in shoot initiation in these studies occurred because the
experimental protocols depended on the plants’ internal initiation cycle to
initiate shoots. This cycle slows down when plants are exposed to lower tem-
peratures (Whiley et al., 1988, 1989, 1991).
Floral or vegetative induction occurs when shoots are initiated. Resting
buds of plants that are exposed to cool temperatures (18°C day/10°C night)
for > 3 weeks and then transferred to a warm temperature (30°C day/25°C
night) before initiation, produce only vegetative shoots (Núñez-Elisea et al.,
1996). Thus, the stems do not ‘remember’ that they had been exposed to floral
inductive conditions while still in rest. They responded to warm conditions
present when shoot initiation occurred.
Reproductive Physiology 113
This response to temperature conditions at the time of shoot initiation
extends to the formation of transition shoots if conditions change during
early shoot development. First reported by Naik and Mohan Rao (1943),
transition shoots are an unusual transition in expression of shoot type during
a single growth flush (Kulkarni, 1988b; Núñez-Elisea and Davenport, 1989,
1992b; Batten and McConchie, 1995). The transition typically occurs near the
middle of the extending shoot. Resting buds possess preformed nodes, each
of which contains a primordial leaf or bract and a lateral meristem. The api-
cal meristem initiates cell division at the same time or soon after the nodal
target tissues begin development (Mustard and Lynch, 1946; L.B. Singh, 1960;
Núñez-Elisea et al., 1996). Vegetative or inflorescence development in the
pre-formed primordia is underway before the apical meristem begins to pro-
duce differentiating cells. Transfer from a warm, vegetatively inductive con-
dition to a cool, floral inductive environment at early bud break results in
formation of V/F transition shoots (Fig. 5.5). Transfer from cool to warm con-
ditions at the same stage of bud break results in formation of F/V transition
shoots (Batten and McConchie, 1995; Núñez-Elisea et al., 1996).
The flowering response to temperature occurs in mangoes growing in
subtropical latitudes where cool temperature is the dominant induction fac-
tor. Many cultivars flower erratically in the low-latitude tropics, providing
continuously warm temperatures with high soil and atmospheric moisture.
Under such conditions, the age of stems is the dominant inductive factor
(Buell, 1954; Nakasone et al., 1955; Ravishankar et al., 1979; Ou and Yen, 1985;
Issarakraisila et al., 1992), and occasional cool night temperatures in the upper
latitude tropics have a positive moderating effect (Davenport, 2003).
Water relations
In the absence of cool temperatures, mango trees in the tropics may flower in
response to irrigation or rain following periods of water stress lasting 6–12
weeks or more (Pongsomboon, 1991). Plant water stress has been presumed
to provide the stimulus for flowering (reviewed in Whiley, 1993; Chaikiatti-
yos et al., 1994; Schaffer et al., 1994; Davenport and Núñez-Elisea, 1997); how-
ever, most of these studies have failed to substantiate prolonged tree water
deficit as a successful agent for floral induction.
Experiments with container-grown trees fail to produce inflorescences
after 8 weeks of water deficit (Wolstenholme and Hofmeyr, 1985). Under
glasshouse conditions (27°C day/22°C night; relative humidity (RH) ≥ 90%),
container-grown, monoembryonic cultivars were water stressed through
deficit irrigation for 14 days, resulting in an average leaf xylem water poten-
tial of −3.9 MPa (Davenport, 1992; Núñez-Elisea and Davenport, 1992a,
1994b). Following resumption of irrigation, all trees grew vegetatively. Sim-
ilarly, only vegetative growth was obtained when container-grown trees
were deprived of irrigation for 36 days during summer, although leaf xylem
water potentials of −3.78 MPa were attained (Núñez-Elisea and Davenport,
1994b). Water stress imposed on plants during the cool autumn months
T.L. Davenport 114
(night temperatures < 15°C) do not increase the proportion of apical buds
forming inflorescences, but expedited shoot initiation after rewatering
(Núñez-Elisea and Davenport, 1994b). These results demonstrated that cool
temperatures provide inductive conditions, whereas relief of water stress
accelerated shoot initiation under cool, inductive temperatures. Flowering
was delayed when container-grown monoembryonic mangoes were water-
stressed at 18°C day/15°C night (Chaikiattiyos et al., 1994). Water-stressed
trees held at 29°C day/25°C night did not flower.
Mango trees growing in the low-latitude tropics may flower after an
extended period of mild water stress (Harris, 1901; Collins, 1903; Kinman,
1918; Gangolly et al., 1957; Gangolly, 1960; L.B. Singh, 1960). Pongsomboon
et al. (1991) observed flowering in field-grown trees in the tropics following
6 weeks of withholding water. The primary impact of water stress appears to
be prevention of shoot initiation during stress. The accumulating age of stems
is greater in water-stressed trees than in trees maintained under well-watered
conditions that promote frequent vegetative flushes (Davenport, 1992, 1993;
Schaffer et al., 1994). This delay in flushing may provide more time for accu-
mulation of a putative FP (Schaffer et al., 1994) or reduction in the level of a
putative VP (Davenport and Núñez-Elisea, 1997; Davenport, 2000). Some
cultivars appear to be better adapted to such delays in growth and perform
better in dry environments in the tropics.
Effect of N on flowering
Subsequent to the discovery of ethephon to stimulate mango flowering (Gon-
zalez, 1923; Alcala and San Pedro, 1935), Barba (1974), Bueno and Valmayor
(1974), Astudillo and Bondad (1978), Bondad et al. (1978), Bondad and Apos-
tol (1979), Pantastico and Manuel (1978) and Bondad and Linsangan (1979)
reported that KNO
3
could be used for the same purpose. This has been
exploited in the low- and mid-latitude tropics (Mosqueda-Vázquez and de
los Santos de la Rosa, 1981; Mosqueda-Vázquez and Avila-Resendiz, 1985;
Núñez-Elisea, 1985, 1986; Ou and Yen, 1985; Winston and Wright, 1986;
Tongumpai et al., 1989; Goguey, 1993; Ravishankar et al., 1993; Sergent et al.,
1996; Yeshitela et al., 2004b, 2005). The nitrate (NO
3

) anion is the active
component of KNO
3
(Bueno and Valmayor, 1974), and ammonium nitrate
(NH
4
NO
3
) is twice as effective as KNO
3
(Núñez-Elisea, 1988; Núñez-Elisea
and Caldeira, 1988). In the low- and mid-latitude tropics, receptive trees
respond by developing visible reproductive buds within 2 weeks after
application. The effective spray concentration is 1–10% KNO
3
, depending on
the age of the trees and climate. Two to four per cent KNO
3
or calcium nitrate
(Ca(NO
3
)
2
) and 1–2% NH
4
NO
3
are effective for stimulating flowering in
most conditions. The physiological and temporal timing of application is
important. Old trees, non-vigorous trees, and trees in which vegetative
flushes have been discouraged by low water potentials produce the best
response to NO
3

induction (N. Golez, personal communication, the Philip-
pines, 1989).
Reproductive Physiology 115
Chemical bud forcing is most effective in the tropics where distinct wet
and dry seasons prevail. The response to chemical bud forcing by NO
3

and
ethephon diminishes at latitudes > 22° N or S (Mosqueda-Vázquez and de
los Santos de la Rosa, 1981). Their effect may involve the decline of night
temperatures from ≥ 20°C around the equator to ≤ 10°C between 22° and 27°
N or S latitude during winter months or by late summer vegetative flushes.
Trees in the wet or dry subtropics at 25° N or S have not responded to treat-
ments (Davenport, 1993).
Stems must be sufficiently mature, dark green with a minimum age of 4
months since the previous limp, red-leaf stage in easily induced cultivars
and 5 months for more recalcitrant cultivars to obtain a reproductive shoot
response in the low-latitude tropics (Davenport, 2003). Bueno and Valmayor
(1974) indicated that leaves must be brittle when hand-crushed. Núñez-
Elisea (1986, 1988) reported that stems must be at least 6 months old. Trees
that experience autumn dry periods become responsive to treatments as
early as October (northern hemisphere). Groups of stems within tree cano-
pies are produced through asynchronous flushes of growth, and vary in age;
only a few are responsive to the first inductive spray. Subsequent biweekly
applications cause flowering in canopy sectors as they reach the age-depen-
dent requirement for initiation. Early and out-of-season flowering and fruit-
ing can thereby be achieved.
KNO
3
may be floral inductive in mango (Barba, 1974); however, trees in
the upper latitude tropics typically flush vegetatively rather than produce
bloom when either KNO
3
or NH
4
NO
3
is sprayed between June and Septem-
ber (N. Golez, personal communication, the Philippines, 1989). The warm,
rainy season producing frequent flushes of growth during this period is con-
ducive to a vegetative response to the sprays. These results indicate that
KNO
3
and NH
4
NO
3
stimulate shoot initiation but do not determine bud
morphogenesis. In buds released after KNO
3
or NH
4
NO
3
treatments, the
ratio of leaf-generated FP to VP and not NO
3

causes initiating buds to become
reproductive. Kulkarni (1988b, 2004) suggested that the floral stimulus is
present in stems when buds are forced in response to KNO
3
and suggested
that KNO
3
may also sensitize buds to the floral stimulus. Davenport (2003),
T.L. Davenport and J. Oleo (2006, unpublished data) and F. Ramirez and T.L.
Davenport (submitted for publication) observed 100% vegetative shoots
when 4% KNO
3
was foliar applied to 2-month-old stems; whereas, applica-
tion of the same spray treatment to 4.5-month-old stems on trees in the same
orchards resulted in 100% reproductive shoots.
Trees with high leaf N levels rarely flower in the tropics. Lack of flower-
ing is always due to frequent vegetative flushes of growth, especially during
the rainy season. Mango trees must have leaf N levels of 1.4% or less in order
to suppress frequent flushes of vegetative growth (Davenport, 2003). Leaf N
levels of < 1.1% suppress frequent flushes but also provide insufficient nutri-
tion to support good cropping. Thus, 1.1–1.4% N levels in leaves appear to be
optimum for good commercial production and control of flowering time in a
managed orchard. The application of KNO
3
to the foliage of the resting stems
4–5 months after the limp, red-leaf stage will cause a flowering response.
T.L. Davenport 116
Photoperiod
Flowering in most trees does not appear to be under photoperiodic control
(Kozlowski et al., 1991). Mango cultivation is concentrated between 27° N
and 27° S where the shortest annual photoperiod is c.10.5 h and the longest
photoperiod is c.13.5 h. Cultivars in the upper-latitude tropics and subtropics
flower during the winter when photoperiods are short; however, trees in the
low-latitude tropics, where a 12-h photoperiod is nearly constant, can flower
at any time of the year. Furthermore, flowering on spring-initiated shoots in
the subtropics occurs during summer (Schaffer et al., 1994). Studies have
failed to demonstrate a correlation between 8-h photoperiods and flowering
(Maiti, 1971; Maiti and Sen, 1978; Maiti et al., 1978). Núñez-Elisea and Daven-
port (1995) studied the effects of 11-, 12-, 13- and 24-h photoperiods at 18°C
day/10°C night, or 11- and 13-h photoperiods at 30°C day/25°C night on
flowering of container-grown trees. Photoperiod had no effect on the fate of
buds, and the promotive effect of cool temperatures on flowering was inde-
pendent of photoperiod. Photoperiods of 11-, 12- or 13-h with 18°C day/10°C
night, caused flowering in trees within 40 days. The 24-h photoperiod with
12-h thermoperiods of 18°C and 10°C caused flowering of trees within 35
days. Photoperiods of 11- or 13-h at 30°C day/25°C night resulted in vegeta-
tive growth only. With warm temperatures, vegetative shoots were produced
in 17 days. These results confirm that floral induction is caused by cool tem-
peratures and not by short photoperiods and that warm temperature, not a
long photoperiod, caused vegetative induction.
5.6 Hormonal Influence on Flowering
FP is a protein product of the FT gene in Arabidopsis (Corbesier et al., 2007)
and the Hd3a gene in rice (Tamaki et al., 2007) and moves in phloem from
leaves to buds; there is little evidence that phytohormones are directly
involved as the FP. Phytohormones appear to be responsible for shoot initia-
tion in conditions that are floral inductive.
Ethylene
Smudging has been utilized to stimulate mango flowering in the Philippines.
Only branches that attain sufficient age respond to smudging by forming
reproductive shoots (Acala and San Pedro, 1935; Bueno and Valmayor, 1974).
Rodriguez (1932), investigating smoke-induced flowering of pineapple, pro-
posed that ethylene, generated by burning material, may stimulate flower-
ing. Dutcher (1972) confirmed that smoke from smudge fires contained
ethylene. Smudging and the use of ethephon in 1968 by F. Manuel (Barba,
1974) and others (Bondad, 1972, 1976) to promote mango flowering sug-
gested that endogenous ethylene is integral for floral induction (Barba, 1974;
Bondad, 1976; Chadha and Pal, 1986). Ethephon effectively promotes flowering
Reproductive Physiology 117
of mangoes under specific conditions in the low-latitude tropics (Davenport
and Núñez-Elisea, 1997).
The involvement of endogenous ethylene in flowering is supported by
observations that indirectly link it to symptoms of ethylene production.
Extrusion of latex from terminal buds occurs at the time of inflorescence ini-
tiation, and epinasty of mature leaves near the apex during expansion of the
panicle has been observed (Davenport and Núñez-Elisea, 1990, 1991). Both
are symptoms of plants exposed to high ethylene levels (Abeles, 1973). Indi-
rect support also comes from reports that KNO
3
-stimulated flowering of
mango is mediated by increased levels of endogenous ethylene (Thuck-Thye,
1978; Lopez et al., 1984). Mosqueda-Vázquez and Avila-Resendiz (1985)
reported that the efficacy of KNO
3
was negated by cobalt chloride (CoCl
2
)
and silver nitrate (AgNO
3
), which inhibit the synthesis and action of ethyl-
ene, respectively, when sprayed 1–4 h after KNO
3
. Saidha et al. (1983) reported
a gradual increase in endogenous leaf ethylene production as the season of
floral initiation approached. Ethylene production by stems producing repro-
ductive shoots was up to fivefold that of resting stems.
Inconsistent (Pandey et al., 1973; Sen et al., 1973; Winston and Wright,
1986) or non-responsive results with ethephon (Pandey and Narwadkar,
1984; Ou and Yen, 1985; Pandey, 1989) or smudging (Sen and Roy, 1935),
especially during warm, non-inductive conditions, have been reported. Dav-
enport and Núñez-Elisea (1990, 1991) reported elevated ethylene production
in mango stems in response to ethephon sprays without an accompanying
floral response. Experiments were conducted during floral-inductive and
non-inductive periods. Unlike Saidha et al. (1983), they observed no increase
in ethylene production rates prior to or during panicle development.
The effect of ethylene on flowering is unresolved. It is likely that ethylene
stimulates shoot initiation by inhibiting auxin transport from leaves to buds
and stems (Morgan and Gausman, 1966; Beyer and Morgan, 1971; Riov and
Goren, 1979, 1980; Ramina et al., 1986). This may increase the ratio of cytoki-
nin to auxin in buds and stimulate shoot initiation (Davenport, 2000). Other
factors (i.e. cool temperatures or aged leaves) may be responsible for floral
induction (Ona and de Guzman, 1982; Davenport, 1993).
Auxin
Although auxin may have a critical role in floral induction of mango (Chadha
and Pal, 1986; Hegele et al., 2006), there is little supporting evidence. The
application (L.B. Singh, 1961; Singh and Singh, 1963; Bakr et al., 1981; Pandey
and Narwadkar, 1984) and analysis of auxin in leaves (Paulas and Shanmu-
gavelu, 1989; Sivagami et al., 1989), stems (Chen, 1987) and shoots (Chacko et al.,
1972b) have been reported in relation to mango flowering. These studies are
inconclusive due to inconsistencies in purification and analytical methodolo-
gies (Davenport and Núñez-Elisea, 1997).
Auxin may indirectly stimulate root-produced cytokinins through initia-
tion of new root growth. Auxin is transported basipetally from growing
T.L. Davenport 118
shoots and leaves to roots (Goldsmith, 1968; Cane and Wilkins, 1970; Wilkins
and Cane, 1970; Goldsmith and Ray, 1973; Lomax et al., 1995) and stimulates
root initiation (Hassig, 1974; Wightman et al., 1980). The efficacy of various
auxins for stimulating adventitious rooting of mango marcots and cuttings
was reviewed by Davenport and Núñez-Elisea (1997).
Auxin inhibits shoot initiation (Davies, 1995) and confers apical domi-
nance by preventing axillary bud break. Leaf-produced auxin and petiolar
auxin transport capacity declines as leaves age (Veen, 1969; Veen and Jacobs,
1969; Davenport et al., 1980). The interaction of decreasing auxin and accu-
mulating cytokinins in resting buds may explain the cyclic nature of shoot
initiation. The ratio of cytokinin to auxin levels in buds regulates shoot
initiation (Skoog and Miller, 1957; Bangerth, 1994; Cline et al., 1997; Beveridge
et al., 2003).
Cytokinins
Relationships between mango flowering and the endogenous levels of cyto-
kinins in leaves (Paulas and Shanmugavelu, 1989; Kurian et al., 1992), stem
tips (Agrawal et al., 1980) and xylem sap (Chen, 1987) and the effect of cyto-
kinin applications on bud break and shoot development have been reported.
Chen (1985) described precocious flowering of mango shoots in response to
early October application of 6-benzylaminopurine (BA). Flowering was
observed 1 month following application and 3 months later on non-treated
trees. Núñez-Elisea et al. (1990) reported numerous reproductive shoots per
stem in response to the synthetic cytokinin, thidiazuron, during cool, floral
inductive conditions; however, numerous vegetative shoots per stem were
initiated when thidiazuron was applied during warm, vegetatively induc-
tive conditions. Early bud break was not achieved following foliar applica-
tion of Promalin (commercial formulation of BA and gibberellins A
4
+A
7
)
(Oosthuyse, 1991b), BA (A.K. Singh and Rajput, 1990) or kinetin (Singh and
Singh, 1974).
Chen (1987) reported the lowest levels of putative trans zeatin and its
riboside were translocated from roots during the vegetative shoot growth
and resting stages, whereas the highest levels occurred during early flower-
ing and full bloom. Paulas and Shanmugavelu (1989) observed no significant
difference in cytokinin levels of the fourth and fifth leaves during resting bud
and flowering. Cytokinin levels in mango stem buds increased during expo-
sure to cool, floral inductive temperatures (Bangerth et al., 2004). Agrawal
et al. (1980) described 11 cytokinin-like substances isolated from stem tips of
an alternate-bearing cultivar in ‘on’ and ‘off’ years. Kurian et al. (1992) reported
a link between PBZ applications and reduction in cytokinins in mango leaves
with treatments, perhaps caused by reduction in feeder root development
and formation of thick, blunt roots (Bausher and Yelenosky, 1987; Peng et al.,
1991; Burrows et al., 1992; Yelenosky et al., 1993). Concurrent with this response
was suppression of bud initiation and reduced internode lengths for c.2
years.
Reproductive Physiology 119
The role of cytokinins in flowering is unresolved due to sampling of dif-
ferent organs at non-comparable times or conditions. The elevated cytokinin
levels found prior to and during flowering and the flowering response to
applied BA led to the conclusion that cytokinins are involved in flowering of
mango (Chen, 1985, 1987; Bangerth, 2006); however, such responses can be
explained if cytokinins are involved in stimulation of bud break (i.e. shoot
initiation) during floral inductive conditions.
A well-documented role for cytokinins in higher plants, especially evi-
dent in vitro, is bud organogenesis (Skoog and Miller, 1957; Miller, 1963;
Takahashi, 1986; Salisbury and Ross, 1992; Davies, 1995; Haberer and Kieber,
2002). The primary cytokinins in higher plants are trans zeatin, dihydrozeatin,
isopentenyl adenine and their ribosides. They are translocated from roots and
accumulate in resting buds (Hendry et al., 1982a, b) or can possibly be synthe-
sized in nearby tissues as regulated by auxin (Nordstrom et al., 2004; Tanaka
et al., 2006). Their rate of accumulation may relate to periodic root flushes that
alternate with shoot flushes (Krishnamurthi et al., 1960; Bevington and Castle,
1986; Cull, 1987, 1991; Parisot, 1988; Williamson and Coston, 1989).
Gibberellins
Gibberellins are tetracyclic diterpenoid compounds that vary in biological
activity according to the type and location of substituted side groups on a
basic ent-gibberellane skeleton. The number of known gibberellins is > 100
(Pearce et al., 1994). Reproductive shoot initiation is suppressed in many
woody angiosperms by gibberellic acid (GA
3
) (Pharis and King, 1985). GA
3

inhibits mango flowering (older literature reviewed in Davenport and
Núñez-Elisea, 1997; Núñez-Elisea and Davenport, 1998).
GA
3
inhibition of mango flowering is correlated with the applied con-
centration (Kachru et al., 1971, 1972) and may cause buds to develop vegeta-
tively under floral-inductive conditions. Núñez-Elisea and Davenport (1991a,
1998) reported a delay in initiation of axillary shoots when GA
3
was foliar
applied to deblossomed stems during cool, floral inductive temperatures.
Higher concentrations caused longer delays in shoot initiation. GA
3
did not
inhibit floral induction, so long as cool, inductive temperatures were present
during axillary shoot initiation. Late initiating buds, which grew during
warm, spring temperatures, however, formed vegetative shoots. Similar
delays in reproductive shoot initiation in response to GA
3
application was
reported by Shawky et al. (1978) and Turnbull et al. (1996). Multiple applica-
tions, even at lower rates, are more effective than a single application (Tomer,
1984; Turnbull et al., 1996; Davenport and Smith, 1997). GA
3
treatment has
been recommended in the Canary Islands to delay flowering until the danger
of frost has passed (Galán-Saúco, 1990). In the subtropics of Australia, it is
used to prevent flowering in newly planted trees during the spring so that the
full growing period can be utilized for vegetative growth, thereby hastening
orchard establishment (A.W. Whiley, personal communication, Queensland,
1996).
T.L. Davenport 120
Response to GA
3
varies among cultivars, growing conditions and timing
of application (Tomer, 1984; Oosthuyse, 1995a; Turnbull et al., 1996; Sánchez-
Sánchez et al., 2004). GA
3
can delay shoot initiation beyond the floral induc-
tive window, resulting in a vegetative flush when shoots develop in warm
weather (Kachru et al., 1971, 1972; Núñez-Elisea and Davenport, 1991a, 1998;
S. Gazit, personal communication, Israel, 1993). The variable response to GA
3

may be related to levels of active gibberellin in buds at the time of applica-
tion, inconsistent uptake or differential sensitivity of buds, depending on
their position (apical versus axillary) or age (Núñez-Elisea and Davenport,
1991a, 1998). Efficacy is related to the timing of application; immediately
prior to normal shoot initiation appears to be most effective (Davenport and
Smith, 1997).
Reports of endogenous gibberellins in mango tissues, especially in buds,
are difficult to interpret with respect to a regulatory role in bud break or
flowering. Problems include sampling of tissues other than apical buds, i.e.
whole stems (Tongumpai et al., 1991b), leaves (Paulus and Shanmugavelu,
1989; Sivagami et al., 1989) and xylem sap (Chen, 1987), or at times when
developing shoots may contribute to the overall result (Chen, 1987). Pal and
Ram (1978) tentatively identified the presence of gibberellins A
1
, A
3
, A
4
, A
5
,
A
6
, A
7
and A
9
. Chen (1987) identified gibberellins A
1/3
, A
4/7
, A
5
, A
17
, A
20
and
A
29
. The estimated levels of gibberellins in apical buds for 6 months prior to
the flowering season were reported to be higher in the ‘off’ year than in the
‘on’ year of an alternate-bearing cultivar (Pal and Ram, 1978). Chen (1987)
reported the highest levels of gibberellins in xylem sap during leaf differen-
tiation and lower concentrations during rest, panicle emergence and full
flowering. Tongumpai et al. (1991b) observed increasing levels of gibberellins
in whole stems over the 16 weeks prior to vegetative shoot emergence and
decreasing levels over the same period prior to panicle development. Gib-
berellins A
1
, epi-A
1
, A
3
, A
19
, A
20
and an unidentified gibberellin in buds and
leaves from shoot and stem tips of different ages have been quantified (Dav-
enport et al., 2001b). The detected gibberellins are members of the early
13-hydroxylation pathway of gibberellin synthesis (Takahashi, 1986; Pearce
et al., 1994). Gibberellins A
3
and A
19
were the most abundant gibberellins in
apical stem buds. The concentration of GA
3
increased within buds with
increasing age of stems, although concentrations of other GAs were variable.
The concentration of GA
3
did not change significantly with age in leaves,
whereas that of most of the other GAs declined. Davenport et al. (2001b)
concluded that elevated GA
3
levels in buds may enhance or maintain the
synthesis or activity of endogenous auxin to maintain low cytokinin/auxin
ratios and enhance inhibition of shoot initiation (Jacobs and Case 1965; Scott
et al., 1967; Pharis et al., 1972; Ross et al., 1983; Law 1987; Law and Hamilton,
1989).
The roles of gibberellins and other phytohormones in shoot initiation
and induction is unclear. Endogenous levels in buds and leaves must be cor-
related with physiological events in individual stems. Experimental approaches
should include examination of resting buds up to both vegetative and repro-
ductive shoot initiation to avoid misinterpretation of results. Experiments
Reproductive Physiology 121
should utilize plants grown under defined conditions with specific environ-
mental controls for evaluation of cause and effect. Finally, extraction and
purification protocols should include quantifiable internal standards and
use of sensitive unambiguous analytical techniques.
Plant growth retardants
Plant growth retardants have been evaluated to stimulate early or more
intense flowering, especially in the ‘off’ year of alternate-bearing cultivars
(Davenport and Núñez-Elisea, 1997). They are in three main classes: (i) the
gibberellin transport inhibitor, daminozide (N-dimethylamino-succinamic
acid), known as alar or B-Nine; (ii) the onium type, chloremquat chloride
(2-chloroethyl trimethylammonium chloride), known as cycocel and CCC;
and (iii) the steroid-synthesis-inhibiting triazoles, for example PBZ (PP-333),
known as Cultar
®
, and uniconazole, known as XE-1019 or Sumagic (Rademacher,
1991, 2000a). The latter two classes of compounds inhibit ent-kaurene syn-
thetase, an enzyme in the gibberellin synthesis pathway (Nickell, 1983;
Dalziel and Lawrence, 1984; Rademacher, 1991, 2000a). Applying daminoz-
ide results in increased gibberellin levels, perhaps due to the inability to dis-
tribute it properly (Rademacher, 1991). Plant responses may depend upon
whether target tissues are near the site of gibberellin synthesis or sufficiently
removed from it to be affected by the inhibited translocation.
Daminozide and cycocel
The efficacy of daminozide and cycocel for increasing flowering in the ‘off’ sea-
son of alternate-bearing cultivars has been studied (Maiti et al., 1972; Mukhopad-
hyaya, 1978; Rath and Das, 1979; Suryanarayana, 1980; Rath et al., 1982; Ou and
Yen, 1985), together with their ability to stimulate early flowering (Suryanarayana
and Rao, 1977; Chen, 1985; Kurian and Iyer, 1993a, b). Enhanced, inconsistent
flowering occurs in response to these compounds, especially cycocel.
Triazoles
PBZ is being used (except in the USA where it has not been cleared for use)
to stimulate enhanced or early flowering. It is best applied to the soil due to
its low solubility, long residual activity and lack of efficient foliar uptake
(Rademacher, 2000b). PBZ applied as a soil drench (1–20 g active ingredient
(ai)/tree) reduces internode lengths and causes earlier and enhanced flower-
ing in mango trees (Hasdiseve and Tongumpai, 1986; Haw, 1986; Hongsb-
hanich, 1986). Depending on climate, residual activity lasts for c.2 years
(Kulkarni, 1988a). These results have been confirmed in different locations in
the tropics (Davenport and Núñez-Elisea, 1997; Yeshitela et al., 2004a, b).
Nartvaranant et al. (2000) recommended soil application of PBZ at 1–1.5 g
ai/m of canopy diameter to achieve flowering in 90–120 days if the trees are
stimulated to flush. Davenport (2003) observed that such treatments allowed
a reduction of c.1 month in the time required for stem rest before stimulating
them to initiate reproductive shoots using KNO
3
. PBZ also reduces alternate
T.L. Davenport 122
bearing of some cultivars (Hillier and Rudge, 1991; Burondkar and Gunjate,
1993; Rao, 1997; Rao et al., 1997; Rao and Srihari, 1998; Vijayalakshmi and
Srinivasan, 1999). Cultivars that tend to flower with minimal inductive impe-
tus are more responsive and can be induced to flower out-of-season using
PBZ (Tongumpai et al., 1989). Núñez-Elisea et al. (1993) demonstrated that
application of PBZ and uniconazole advanced bud break of containerized
trees in controlled environment chambers, but cool temperatures were neces-
sary to induce flowering. Initiated shoots were induced to be vegetative in
warm temperatures. The greater proportion of purely reproductive panicles
in treated plants (compared with controls) suggests that triazoles impact the
level of a putative VP, probably a gibberellin. Whiley (1993) suggested a sec-
ondary mechanism for the floral promotive action of PBZ on mangoes, not-
ing inconsistent responses in the literature between cultivars, environments
and application times.
Application of PBZ reduces the number of panicles, despite increased
fruit set (Goguey, 1990). Davenport (1987, 1994) observed neither growth
inhibition nor enhanced or early flowering in response to root drenches or
bark banding with uniconazole (1–5 g ai/tree) in trees growing in alkaline,
calcareous soil. He reported that new shoot growth was stunted with
extremely short internodes when trees were severely pruned soon after or as
long as 3 years after treatment. Yield was severely reduced due to the lack of
normal growth flushes. The growth stunting effect continued for 7 years after
pruning. Davenport (1994) warned that use of triazole plant growth retar-
dants for control of tree growth, flowering or yield must be done with con-
siderable caution, especially if severe pruning of the trees is anticipated.
Residual uniconazole or PBZ applied as a soil drench or bark band is appar-
ently retained in high concentrations in main scaffolding branches. In Cen-
tral and South America, growers utilize PBZ annually to stimulate early
flowering. A test tree should be severely pruned to determine if the trees are
affected by PBZ to anticipate the orchard response to later severe pruning.
Certain gibberellins (i.e. GA
1
) are necessary for shoot elongation. Inhibi-
tion of bud break and shoot elongation in response to application of the
growth retardants cycocel (Maiti et al., 1972) and triazoles (Kulkarni, 1988a;
Burondkar and Gunjate, 1991, 1993; Tongumpai et al., 1991a; Kurian et al.,
1992; Winston, 1992; Kurian and Iyer, 1993a, b; Núñez-Elisea et al., 1993; Wer-
ner, 1993) have been reported. Elongation of panicles is inhibited, especially
by high levels of triazoles (Kulkarni, 1988b; Winston, 1992; Davenport, 1994;
Salomon and Reuveni, 1994). Inflorescences in treated trees may become
compact, improving opportunities for disease and insect attack (Winston,
1992). Kurian et al. (1992) associated reduced cytokinin levels in leaves with
inhibition of shoot initiation in plants treated with soil drenches of PBZ. Ele-
vated, concentration-dependent levels of phenolic compounds were also
found in resting apical buds of PBZ-treated trees (Kurian et al., 1994). They
suggested that low cytokinin activity and high phenolic levels in buds con-
tributed to inhibition of shoot initiation.
The combined impact of the gibberellin synthesis-inhibiting triazoles on
shoot initiation, induction, and elongation implies that several different
Reproductive Physiology 123
gibberellins regulate specific activities in mango plants. This is supported by
the inhibitory effect of GA
3
on shoot initiation in contrast with early initia-
tion of flowering in triazole-treated trees. Compression of reproductive and
vegetative shoot internodes may involve inhibition of GA
1
synthesis. Stimu-
lation of flowering instead of vegetative growth during early initiation in
triazole-treated plants in marginal or non-floral inductive conditions, sug-
gests that the putative VP, a gibberellin other than GA
3
or GA
1
, is reduced
when gibberellin synthesis is inhibited.
5.7 Photoassimilate Influence on Flowering
Flowering may be regulated by C:N ratios with high levels being conducive
to flowering (Kraus and Kraybill, 1918). Photoassimilates reaching the apical
bud from leaves was central to several theories of floral induction (Sachs,
1977; Bernier and Sachs, 1979; Bernier et al., 1981, 1993; Bernier, 1988) includ-
ing mango (Mallik, 1951; L.B. Singh, 1960; Chacko and Ananthanarayanan,
1982; Rameshwar, 1989) and other species (Allsopp, 1965; Sachs, 1977; Mishra
and Dhillon, 1978; Ramina et al., 1979; Bernier et al., 1981; Sachs and Hackett,
1983). The theory of photoassimilate diversion to the apical bud (Sachs et al.,
1979) is the basis for the carbohydrate-regulated flowering models (see
below). Sugars are utilized during panicle development (Ravishankar and
Mohan Rao, 1982). Starch reserves and C:N ratios have been correlated with
flowering (Mishra and Dhillon, 1978; Suryanarayana, 1978a, b, c; Chacko and
Ananthanarayanan, 1982; Whiley et al., 1988, 1989, 1991; Robert and Wolsten-
holme, 1992; Shivashankara and Mathai, 1995), and the subject has been
reviewed (L.B. Singh, 1960, 1972; Singh, 1979; Chacko, 1986, 1991; Chadha
and Pal, 1986; Pandey, 1989). Starch accumulation during extended periods
of canopy rest prior to flowering provides supportive evidence, but there is
little consensus regarding the role of carbohydrates and N in flowering.
Photoassimilates may be necessary for floral induction. If a florigenic
promoting gene product is synthesized in leaves in small amounts, it must be
able to move to those buds via phloem. Due to the requirement for high sol-
ute concentrations to motivate phloem flow, the low concentration of the FP
could not cause fluid movement through sieve tubes of the phloem on its
own. The much higher concentrations of photoassimilated sugars carried by
water loading into the phloem in leaves passively transports the FP towards
the various sinks, including respiring buds, where they are utilized for floral
induction.
5.8 Horticultural Manipulation of Flowering
Mango flowering can be stimulated by trunk or branch girdling, defoliation
and deblossoming (Pandey, 1989). Responses vary with cultivar and envi-
ronment. Trunk girdling of mango trees to promote flowering is inconsis-
tently effective (Kinman, 1918; Gaskins, 1963; Winston and Wright, 1986) and
T.L. Davenport 124
can be detrimental to trees, especially if done in subsequent years. It has been
shown to increase flowering in the ‘off’ year of alternate-bearing cultivars;
however, it either has no effect or is only marginally beneficial in the ‘on’ year
(Mallik, 1951; Rath and Das, 1977, 1979; Rath et al., 1982; Rameshwar, 1989).
Girdling in late summer or early autumn usually results in less vegetative
flushing prior to flowering, which is enhanced in trees exposed to marginally
inductive conditions. Tree response is dependent on the width of the girdle.
Narrow cuts result in either a short-term or no response; whereas, girdles that
are too wide can kill trees if they do not close within a reasonable time. Gir-
dling cuts phloem transport, starves roots of photoassimilates and interrupts
auxin transport to roots (Morris and Thomas, 1978; Hegele et al., 2004). These
are detrimental to root development and can alter the bud cytokinin:auxin
ratio due to reduced cytokinin translocation from roots. This results in delayed
shoot initiation, which can impact the level of the age-dependent, putative VP
when shoot initiation occurs. The delay in flushing, therefore, enhances
flowering. Defoliation of trees stimulates flushing, possibly by altering the
cytokinin:auxin ratio in buds because leaves are the primary source of auxin.
Bloom delay is useful where recurring temperatures < 15°C stimulate
flowering, but continued low temperatures hamper pollination, fertilization
and early fruit development (Young and Sauls, 1979; Wolstenholme and Mul-
lins, 1982a, b; Whiley et al., 1988; Galán-Saúco et al., 1991). Low temperatures
cause production of seedless, underdeveloped fruit. Deblossoming stimu-
lates growth of dormant axillary buds, which produce inflorescences if initia-
tion occurs under conditions conducive to floral morphogenesis (Singh et al.,
1974). Late blooms can also be promoted with ethephon (Chadha and Pal,
1986; Galán-Saúco et al., 1993) and cycloheximide (Pal and Chadha, 1982;
Shu, 1993). Sprays of these compounds cause abscission of apical panicles,
thereby releasing dormant axillary buds that will produce inflorescences
under cool, floral-inductive conditions of the subtropics.
5.9 Conceptual Flowering Models
Several conceptual models have been proposed that attempt to explain the
physiological basis of mango flowering. Each model should be viewed as a
collection of integrated ideas, which require rigorous testing for validity
within the context of the models. A useful model should explain how flower-
ing and vegetative growth is regulated in all cultivars and races in both
humid and dry climates in the tropics and subtropics. It should also be sup-
ported by the preponderance of research evidence. The flowering models are
either carbohydrate-regulated or hormone-regulated.
Carbohydrate-regulated flowering models
Cull (1987, 1991) presented a holistic approach for tree crop research and
management to maximize sustainable fruit production. This concept is based
Reproductive Physiology 125
on the axiom of genotype/environment adaptability expressed through the
annual phenological cycle and is an alternative to the traditional reductive-
based approach to crop research and development. He proposed that pro-
ductive cultivars follow ‘normal’ phenological patterns from year to year
due to gene expression in specific environments. A significant departure
from this pattern results in reduced or total crop failure. Annual variations in
climatic conditions that alter tree phenology can be countered by strategic
applications of nutrients, water, plant growth regulators and canopy manip-
ulation. The model does not attempt to explain the intricacies of shoot ini-
tiation or induction, but takes a broader approach in detailing temporal
relationships between reproductive and vegetative growth that lead to reli-
able cropping.
The fundamental principle underlying this model is that yield is the
product of photoassimilate (carbohydrate) accumulation and subsequent
redistribution during the annual growth cycle. Accumulated photoassimi-
lates would drive critical growth events that require higher levels of resources
than are available from current photoassimilate supplies. Cultivars that pro-
ceed with balanced reproductive, vegetative and rest phases are more likely
to have sufficient carbon resources to meet periods of critical demand and
therefore will sustain high yields. The model illustrates floral initiation as
occurring after a 2- to 3-month rest period during autumn/winter when a
critical threshold level of carbohydrate is reached in buds together with a
putative floral stimulus. Bud break during cool weather results in a high per-
centage of flowering stems (> 90%; Searle et al., 1995) with fruit set and reten-
tion suppressing vegetative flushing on individual fruiting stems until after
they have matured and harvested. Shortly after harvest, vegetative buds are
released and a flush of growth occurs during the summer, which is followed
by a period of strong root growth. The regenerated canopy becomes a source
for rebuilding photoassimilate reserves that are stored in the roots, bark and
resting stems. In the tropics, growth events are less orderly, and cultivar and
management skills are of greater importance. The pre-flowering rest period
is usually achieved by drought as temperatures remain above the critical
threshold for shoot growth (15°C) (Whiley et al., 1989). Other practices used
with some success to enforce canopy quiescence are girdling and the applica-
tion of growth retardants.
The principles of phenological modelling have been advanced into work-
ing pheno/physiological models for avocado (Whiley, 1994) and mango
(Searle et al., 1995). The advantage of this approach is that the annual pro-
gression of growth cycles with associated physiological changes is studied
concurrently, adding a further dimension to our understanding of tree growth
and development. Information gathered in this way provides opportunities
to identify and assess critical yield-limiting events, which in the case of man-
goes largely relates to the success or failure of flowering.
Chacko (1991) proposed a flowering model driven by assimilate supply
and diversion to apical meristems (Fig. 5.6). Environmental conditions, such
as water stress, cool temperatures, high evaporative demand, flooding,
girdling and other events that inhibit vegetative growth result in a shift in

T
.
L
.

D
a
v
e
n
p
o
r
t
1
2
6
FLORAL
INDUCTION
KNO
3
(cultivar and location
specific)
High
starch
MISSING
INCREASED
ASSIMILATE supply
to SHOOT APEX
ASSIMILATE
DIVERSION
from SHOOT APEX
GROWTH
STIMULATION
and high
gibberellin
Exogenous
gibberellin
LINK
?
GROWTH
CHECK
Sugar
· Water stress
· Low temperature
· High VPD
· Flooding
· High temperature
· High humidity
· High soil moisture
High
nitrogen
High
gibberellin
levels
JUVENILITY
HEREDITY
Over vigorous
cultivars
e.g. ‘Kensington’
Dwarf/precocious
cultivars
e.g. ‘Irwin’
Frequent
flushing of
roots and shoots
· Stem girdling
· Root pruning
· High reserves
· Efficient
assimilate
partitioning
· Low reserves
· More wood
formation
· Mild nitrogen
stress
· Growth
retardants
· Inhibitors
FLORAL
INHIBITION
Fig. 5.6. Chacko’s Assimilate Supply and Diversion Flowering Model, a carbohydrate-regulated flowering model (Source: Chacko, 1991).
Reproductive Physiology 127
carbohydrate partitioning and a diversion of soluble assimilates to stem api-
ces. The elevated carbohydrate status in buds, together with a floral stimulus,
results in floral induction. Vigorously growing cultivars and juvenile plants
have low starch reserves (Whiley et al., 1988, 1989, 1991) and a diversion of
soluble assimilates from stem apices results in floral inhibition. Conditions
that promote vegetative growth, i.e. high temperature and moisture, high
gibberellins and N, also lead to floral inhibition. Experiments involving
chemical girdling of trees are based on this model (Blaikie et al., 1999).
Hormone-regulated flowering models
Tri-factor Hypothesis of Flowering
Extensive work on movement of the putative floral stimulus across grafts
from donor to receptor stems (Kulkarni, 1986, 1988b) and the inhibitory influ-
ence of fruit on subsequent flowering (Kulkarni and Rameshwar, 1989) form
the basis of a flowering model proposed by Kulkarni (1991): the Tri-factor
Hypothesis of Flowering in mango (Kulkarni, 2004). This theory (Fig. 5.7)
proposes an interactive role for a putative, cyclically produced floral stimu-
lus in leaves, a floral inhibitor in leaves and fruits, and bud activity during
the floral cycle. During dormancy following a vegetative cycle, genetic and
Flowering promoter
synthesized in the
leaves in the floral
cycle
Flowering inhibitor
and vegetative
promoter
synthesized in the
leaves and possibly
other organs
Bud activity
in synchrony with
the floral cycle
Pure panicles Mixed leafy panicles Vegetative flush
Genetic and Environmental Factors
Fig. 5.7. Kulkarni’s Tri-factor Hypothesis of Flowering in mango, a hormone-regulated
flowering model (Source: Kulkarni, 2004).
T.L. Davenport 128
environmental factors determine the level of synthesis of the putative floral
stimulus. Flowering occurs only if certain correlative factors are present, for
example if the receptor bud becomes active. If fruits are or have been recently
present on the stem, vegetative growth will result. Presence of the putative
floral inhibitor in leaves interferes with expression of the floral stimulus
resulting in vegetative growth. The level of the floral stimulus determines
the response: high levels give rise to normal panicles, intermediate levels
give rise to mixed panicles and low levels result in vegetative growth.
Comprehensive Conceptual Flowering Model
This is a model of flowering involving the various classes of phytohormones
(Davenport, 1992, 1993, 2000, 2003; Davenport and Núñez-Elisea, 1997)
(Fig. 5.8) based on many lines of experimental evidence as well as on research
of other tropical and subtropical fruit crops with similar phenological cycles
(Menzel, 1983; Davenport, 1990, 1992; Menzel et al., 1990; Menzel and Simp-
son, 1994; Davenport and Stern, 2005). Focusing on events occurring in indi-
vidual buds, it is applicable to monoembryonic and polyembryonic cultivars
in the tropics and subtropics and attempts to explain the physiological basis
for the annual progression of the phenological cycle.
SHOOT FORMATION. Two distinct events must occur for vegetative or repro-
ductive growth to occur in resting apical or lateral buds of mango: (i) the
Mango flowering model
PHOTOASSIMILATES
FRUIT
MIXED SHOOT GENERATIVE SHOOT
CHILLING TEMP.
OTHER FACTORS?
WATER STRESS
PROMOTER
IN LEAVES
INDUCTION
VEGETATIVE SHOOT
AUXIN
GIBBERELLINS
SHOOT INITIATION
ROOT INITIATION
ROOTS
CYTOKININS
GIRDLING
CHILLING TEMP.
STORAGE CARBOHYDRATES
PRUNING
DEFOLIATION
NITRATE SPRAY
ETHYLENE
FREQUENT
VEGETATIVE
GROWTH
AUXIN
GIBBERELLINS A
3
A
x
GA
1
GA
3
GA
x
Fig. 5.8. Davenport’s Comprehensive Conceptual Hormone-regulated Flowering Model
(Source: Davenport and Núñez-Elisea, 1997; Davenport, 2000). Single lines indicate promotive
impact. Double lines indicate inhibitory impact.
Reproductive Physiology 129
bud(s) must be initiated to grow (shoot initiation); and (ii) at the time of ini-
tiation, shoot development (i.e. vegetative, mixed, or generative) is deter-
mined (induction). Although conditions for floral induction may be present
prior to shoot initiation, determination of that inductive condition in buds is
not made until initiation occurs. Initiation and induction events are regulated
by different signals and each may be manipulated by different stimuli.
Removing the apical whorl of leaves or tip pruning physiologically mature
stems stimulates shoot initiation in apical or lateral buds, respectively. If
containerized plants are maintained in warm temperatures (30°C day/25°C
night) following initiation, vegetative shoot growth is induced. If they are
kept under cool conditions (18°C day/10°C night), initiating shoots are
induced to be generative. In either of the two temperature regimes without
pruning, they do not initiate shoots until the natural flushing event occurs
much later. They become vegetative or reproductive according to the tem-
perature at the time of shoot initiation. If transferred from cool to warm tem-
peratures before shoot initiation, new shoot growth is induced to be vegetative.
Induction is therefore determined at the time of shoot initiation, and plants
rapidly lose their floral inductive potential when removed from the cool envi-
ronment. Determination of shoot type can be reversed during morphogenesis
by transferring containerized trees from warm-to-cool or cool-to-warm con-
ditions (Batten and McConchie, 1995; Núñez-Elisea et al., 1996).
INITIATION CYCLE. The cyclic initiation of vegetative or reproductive shoots
is common to all mango cultivars. Developing vegetative shoots are rich
sources of auxins and gibberellins, which may be inhibitors in an internal
cycle that regulates shoot initiation. Auxins are actively transported basipe-
tally to roots from production sites in stems (Goldsmith, 1968; Cane and
Wilkins, 1970; Wilkins and Cane, 1970; Goldsmith and Ray, 1973), and they
stimulate adventitious root growth in mango and other crops (Hassig, 1974;
Wightman et al., 1980; Sadhu and Bose, 1988; Rajan and Ram, 1989; Núñez-
Elisea et al., 1992). Elevated auxin synthesis in periodically flushing shoots is
likely to form a concentrated pulse of auxin, which inhibits recurring bud
break and moves basipetally to the roots. This pulse of elevated auxin may
stimulate initiation of new root flushes following each vegetative flush.
Alteration of root and shoot growth occurs in mango (Krishnamurthi et al.,
1960; Cull, 1987, 1991; Parisot, 1988) and other tropical and subtropical trees
(Bevington and Castle, 1986; Williamson and Coston, 1989; Ploetz et al., 1991,
1993). Timing of the root flush may depend on the distance from stems to
roots, the physiological condition of the transport path, and environmental
conditions (i.e. temperature or water relations).
New roots that develop following growth stimulation are a primary
source of cytokinins (Davies, 1995). Cytokinins are transported passively to
stems via the xylem sap in all plants and are active in bud break (Went, 1943;
Kende and Sitton, 1967; Sitton et al., 1967; Itai et al., 1973; Haberer and Kieber,
2002). Cytokinins stimulate shoot initiation in mango (Chen, 1985; Núñez-
Elisea et al., 1990) and other plants (Oslund and Davenport, 1987; Belding and
Young, 1989; Williamson and Coston, 1989; Davenport, 1990; Davies, 1995;
T.L. Davenport 130
Henny, 1995). Auxin inhibits shoot initiation (Davies, 1995) and confers api-
cal dominance by preventing axillary bud break. Leaf-produced auxin and
petiolar auxin transport capacity declines as leaves age (Davenport et al.,
1980). Auxin and cytokinins may therefore be involved in the periodic cycle
of bud break.
A critical balance of these two phytohormones, possibly modulated by
GA
3
, may regulate shoot initiation. During a rest period, the inhibitory action
of auxin transported to buds decreases with time; whereas, cytokinin lev-
els in buds increase (Chen, 1987). When a critical cytokinin/auxin ratio is
achieved, the buds are stimulated to grow, thereby resetting the cycle with
initiation of new shoots. The interaction of auxin and cytokinin to control
bud break in plants is a concept that is supported by molecular studies (see
review by Nordstrom et al., 2004). Moreover, initiation of shoot growth occurs
following pruning, defoliation or the application of thidiazuron (Núñez-
Elisea et al., 1990). Vigorous cultivars (Whiley et al., 1989) and young, small
trees under vegetatively promotive conditions flush frequently with only
short periods of rest; however, this cycle slows with age. Old centennial trees
flush infrequently (N. Golez, personal communication, the Philippines, 1989).
Foliar or soil-applied NO
3

stimulates initiation of reproductive shoots
only if applied after resting stems have attained an age to overcome any veg-
etatively inductive influence. In contrast, high N in soils leads to high N lev-
els in leaves resulting in frequent vegetative flushes. The mechanism whereby
NO
3

stimulates shoot initiation is unknown.
Seeds are rich sources of auxin and gibberellins, which contribute to the
strong inhibition of bud break commonly observed on fruit-bearing mango
stems. The longer that fruit are attached to stems, the longer the postharvest
inhibition may last in the stem (Kulkarni and Rameshwar, 1989; Kulkarni,
1991).
Water stress inhibits shoot initiation by its direct impact on cell division
and elongation possibly by interfering with translocation of cytokinins from
roots. There is little evidence that water stress is directly involved in induc-
tive processes. During water stress, roots continue to grow and produce cyto-
kinins (Itai and Vaadia, 1965; Itai et al., 1968; Wu et al., 1994). Reduced xylem
flux due to limited soil hydration, and transpiration due to increased sto-
matal resistance during water stress may reduce the amount of cytokinins
reaching stems. After rewatering, the increased levels of cytokinins in roots
may translocate to and accumulate in buds. Auxin synthesis and transport
from leaves are reduced during water stress (Davenport et al., 1980) and may
require several days for correction after rewatering. This rapid shift in the
cytokinin/auxin ratio of buds may explain the shooting response that occurs
soon after relief of water stress. GA
3
may act with auxin to inhibit shoot ini-
tiation (Davenport et al., 2001b). Early flowering in plants treated with PBZ
may be a response to lowered gibberellin levels, thus lowering the level of
initiation inhibitor.
This model could explain why sectors of tree canopies flush in the trop-
ics. Mango trees flush often and synchronously throughout the canopy when
they are young. With advancing age, the frequency of flushing is reduced
Reproductive Physiology 131
and synchrony is lost, resulting in sporadic flushes of vegetative or reproduc-
tive growth in sections of the canopy. As the distance between stems and
roots increases, the time required for transport of the putative pulses of ele-
vated auxin levels to roots, formed during a vegetative flush, is increased.
Groups of stems exhibiting simultaneous flushing ultimately connect to a
common branch. Dye trace studies indicate that water transport remains in
strict phylotaxic alignment from secondary roots to the canopy, even in large
trees (T.L. Davenport, unpublished results, Florida, 1991). Unless disturbed
by girdling or by pruning of branches or roots, specific branches in the can-
opy communicate only with those roots in phylotaxic alignment with them.
The hormone transport time may vary among sections of the canopy as the
tree grows. This generates individual initiation cycles in sections of the can-
opy that are separately maintained unless resynchronized with the rest of the
tree following a canopy-wide environmental trigger.
Synchronization of growth throughout trees occurs following exposure
to low temperature, water stress, light pruning of the entire tree and any
condition that would increase the postulated cytokinin/auxin ratio in buds
throughout the canopy. An increased ratio may occur by inhibiting auxin
transport from leaves to buds, or increasing cytokinin translocation from
roots to stems. Winter in the subtropics would reduce auxin transport;
whereas, water stress in the tropics may impact the availability of cytokinins
from roots and auxin from leaves. The intensity of the initiation response (i.e.
synchronization of flushes in the canopy) may be regulated by decreased
auxin transport at low temperatures, the base level of which may be deter-
mined by the age of individual stems. Passage of a strong, extended cold
front during subtropical winters produces synchronized flowering. Milder
winters with weak cold fronts result in asynchronous flowering in sections of
trees. The oldest sectors of canopies flower first, followed by sectors bearing
sequentially younger flushes in subsequent cold fronts. Vegetative flushes
occur when night temperatures are > 18°C for significant periods between
cold fronts.
INDUCTION SWITCH. Floral or vegetative induction is possibly governed by the
interactive ratio of a FP that is up-regulated in low temperatures to an age-
regulated VP in leaves at the time of shoot initiation. High FP:VP ratios would
be conducive to induction of generative shoots, low ratios conducive to veg-
etative shoots and an intermediate ratio of the two would be conducive to
mixed shoots. Regardless of the endogenous levels of the two components
perceived in buds at the time of initiation, flowering and vegetative growth
responses can best be explained by the ratio of the two.
Although the putative FP seems to be up-regulated during leaf exposure
to cool temperatures (< 18°C), there appears to be a basal level present at all
times in leaves exposed to higher temperatures. Flowering of mango occurs
in low-latitude tropics lacking cool night temperatures when stems become
sufficiently aged so that the ratio of the basal level of resident FP to decreas-
ing VP increases to a critical threshold to provide floral induction when
shoots are initiated. This could explain how flowering on non-synchronized
T.L. Davenport 132
branches may occur at any time of the year in trees growing in low-latitude
tropics. High proportions of mixed shoots are commonly found in these con-
ditions, indicating the marginally floral-inductive ratios present under these
conditions. In contrast, flowering in younger stems having higher levels of
VP is observed only when initiation occurs in cool, floral-inductive tempera-
tures. More flowering occurs throughout the canopy when stems are exposed
to cool temperatures, attributable to the higher ratio of up-regulated FP to
resident VP.
Genetic differences in base levels of the putative FP and/or VP or the
receptors of these components could explain the range in flowering responses
in tropical and subtropical cultivars and why a cultivar grown in an environ-
ment different from that in which it was selected is less productive. Cultivars
selected in the subtropics usually flower as well in the low-latitude tropics as
those selected in the tropics. Cool temperatures in the subtropics sometimes
cause earlier flowering in tropical cultivars than those selected in the sub-
tropics. Kulkarni (1991) demonstrated that several multi-flowering cultivars
can induce flowering in receptor graft plants and cause a range of the flower-
ing response of the receivers to donors. Some cultivars may produce higher
base levels of putative FP than others. These are the same cultivars that read-
ily flower under warm temperatures and flower early during cool winter
months. The Comprehensive Conceptual Flowering Model suggests that
flowering can occur at any time in any cultivar regardless of origin so long as
stems are sufficiently old to reduce the VP level to below the critical FP/VP
ratio when initiation occurs.
Although the putative FP, perhaps a product of an ortholog of the Arabi-
dopsis FT gene, has not been identified, the VP may be a gibberellin. Triazoles
and other plant growth retardants that inhibit gibberellin synthesis, promote
strong and out-of-season flowering under conditions that would normally
be marginally or non-floral inductive.
PHOTOASSIMILATES. Photoassimilates produced by leaves provide carbohy-
drates essential for development of roots and other plant organs, including
fruit. They are either used immediately by the nearest sinks (Finazzo et al.,
1994) or are stored in locations throughout the tree to be used when demand
for carbon resources exceeds the existing photosynthetic supply (Whiley
et al., 1988, 1989, 1991). A direct role for carbohydrates in shoot initiation or
induction is not part of this model, although they facilitate mass flow in
phloem from leaves to passively carry the FP to buds.
ALTERNATE BEARING. High levels of auxin and gibberellins produced in seeds
possibly inhibit shoot initiation on fruit-bearing stems for weeks or months
following fruit removal. Rapid production of new shoots following light
pruning of fruit-bearing stems after harvest indicates that residual levels of
auxin and gibberellins linger only in the rachis and last intercalary unit. If
fruit are not set on the lingering rachis, there is less inhibition. Heavy fruit set
in 1 year impacts the timing of subsequent shoot initiation on the large num-
ber of fruit-bearing branches. Substantial delays in subsequent vegetative
Reproductive Physiology 133
flushes until close to the normal flowering period impact the flowering abil-
ity of young shoots. This may explain the occurrence of chronic alternate
bearing in some cultivars.
5.10 Floral Management
The Comprehensive Conceptual Flowering Model is consistent with growth
and development patterns of mango trees in the tropics and subtropics. It
provides a reasonable explanation for the various events in the phenologi-
cal model of Cull, predicts what will happen under a defined set of cir-
cumstances and is being used to develop strategies for consistent mango
flowering. Flowering can be obtained at any time of the year in a flowering
management programme (Davenport, 2003).
The flowering management programme begins each season with tip
pruning of the entire canopy of orchard trees. Tip pruning can be done imme-
diately after harvest to move production forward in the following year or
c.1–2 months after harvest, depending upon cultivar, in order to achieve har-
vest at the same time as the previous year. If sufficient soil water is available
at the time of pruning, a vegetative flush will occur on all pruned stems c.1
month later. The number of new shoots that will mature to become stems will
be five- to eightfold greater than the original number of pre-pruned stems
due to initiation of many lateral vegetative shoots on each stem. This increase
in terminal stem number in the canopy will be reflected in a concomitant
increase in yield. The frequent flushes that can cause an early second flush of
vegetative growth tend to be suppressed.
The new stems must not flush a second time until at least 5 months after
pruning (Davenport, 2006). If they flush within 3–4 months after pruning,
they will be induced to be vegetative. Pre-prune leaf N levels in the stems
must be 1.1–1.4% in order to suppress a second flush of vegetative growth
during the rainy season. Mild water stress after the post-prune flush during
the dry season will suppress a second, undesired vegetative flush when leaf
N levels are above the optimum range. Pruning near the end of the dry sea-
son in non-irrigated or furrow-irrigated trees should be avoided. Transition
from dry to wet season 2–3 months after pruning causes a rain-stimulated
vegetative flush prior to achieving sufficient age of stems from the last flush.
Test sprays of 4% KNO
3
on two to three representative trees should be
applied 5 months (for easily induced cultivars) and 6 months (for more dif-
ficult to flower cultivars) after pruning. If no developing shoots occur within
2 weeks, the spray is repeated. A flowering response is usually evident after
the second application. The other trees that were pruned on or near the same
date can then receive the foliar spray and will respond by synchronized flow-
ering. Although Davenport (2003) described the appropriate timing of PBZ
in a flowering management programme, it is not recommended because
flowering can be achieved without it. For orchard trees to be amenable to tip
pruning, efficient spray application of KNO
3
and easy harvesting, they
should be no taller than 4 m. Pruning to rejuvenate large mango trees and
T.L. Davenport 134
properly shape trees for the annual flowering management programme is
recommended (Davenport, 2006).
5.11 Floral Biology
Detailed descriptions of generative shoots were reviewed in Davenport
and Núñez-Elisea (1997). Juliano and Cuevas (1932) described ovary devel-
opment.
Sex ratio
Sex ratio (i.e. the proportion of perfect to staminate flowers) is a variable
component within panicles, trees and among cultivars. This ratio varies with
cultivar, but is usually < 50% (Davenport and Núñez-Elisea, 1997). Most per-
fect and staminate flowers are borne in the proximal portion of panicles due
to their architecture (Musahib-ud-din and Dinsa, 1946; Cobin, 1950). The
variability in the perfect/staminate flower ratio may be governed by physi-
ological and environmental conditions. Most studies indicate that although
the total number of flowers is substantially less in the distal half of panicles,
there is a greater proportion of perfect flowers in this region (Davenport and
Núñez-Elisea, 1997); however, this condition may be reversed in some culti-
vars (Hussein et al., 1989).
Perfect flowers tend to form in the terminals of individual inflorescences
while staminate flowers are displayed in the earlier forming flowers located
closer to the panicle axis. When panicles begin to elongate in the lower inflo-
rescences, only staminate flowers form and the perfect flowers form at the
terminus of each lateral inflorescence. As more distally located lateral inflo-
rescences begin elongation and anthesis, they too first display staminate
flowers before perfect flowers. These inflorescences are progressively shorter
than previously formed proximal inflorescences, and there are fewer stami-
nate flowers. The final vertical spike of the panicle is composed almost exclu-
sively of perfect flowers. Flowers abscise soon after anthesis, thereby shifting
the sex ratio. The sex ratio should include sex type of all flowers in each
panicle; however, sex ratios are normally determined at some arbitrary
moment during panicle development. Thus, the sex ratio is naturally vari-
able, increasing from extremely low to extremely high values so that timing
of observations during panicle development is critical.
Environmental determinants of sex ratio
Tropical cultivars yield poorly in the subtropics due to a small proportion of
perfect flowers on inflorescences (Singh and Singh, 1959; Singh et al., 1965;
Singh, 1971). Cool weather during inflorescence development contributes to
fewer perfect flowers (Naik and Mohan Rao, 1943; Singh et al., 1965, 1966).
Reproductive Physiology 135
Inflorescences that emerge during the middle and end of the flowering sea-
son produce two and seven times more perfect flowers, respectively, than the
early breaking inflorescences (Majumder and Mukherjee, 1961; Singh et al.,
1966). This response correlates with higher temperatures later in the flower-
ing season. In controlled-environment studies, low temperatures (15°C
day/10°C night) reduced the proportion of perfect flowers, particularly in
tropical, polyembryonic cultivars relative to subtropical, monoembryonic
cultivars (Sukhvibul et al., 1999).
Physiological determinants of sex ratio
Endogenous factors affect the ratio of perfect to staminate flowers. Bajwa
et al. (1956), Majumder and Mukherjee (1961) and Joubert et al. (1993) reported
that lateral inflorescences on mixed shoots carried higher proportions of per-
fect flowers. Inflorescences on older trees produce higher proportions of per-
fect flowers than those on young trees (Naik and Mohan Rao, 1943; Majumder
and Mukherjee, 1961; Chacko and Randhawa, 1971; Pandey, 1989). This also
occurs in inflorescences borne on grafted compared with seedling trees
(Musahib-ud-din and Dinsa, 1946). The effect of tree maturity or rootstocks
on sex ratio of flowers is not understood. Panicles carried within the canopy
of some cultivars (Majumder and Mukherjee, 1961; Singh et al., 1966) or on
particular sides of the canopy (Mukherjee, 1953; Majumder and Mukherjee,
1961) have been reported to have higher proportions of perfect flowers.
Application of some hormones and growth regulators alters the sex ratio
of inflorescences. GA
3
, applied at concentrations of 50–100 mg/l just prior to
inflorescence shoot initiation, substantially reduces the proportion of perfect
flowers (Maiti, 1973), as do combination sprays of urea (0, 3 and 6%) and GA
3

(0, 15 and 30 mg/l) (Rajput and Singh, 1989). Soil-applied PBZ (10 g ai/tree)
significantly increases the ratio of perfect/staminate flowers (Kurian and Iyer,
1993a). Increases in floral ratio also occur with daminozide, whereas maleic
hydrazide either had no effect or lowered the ratio (Singh et al., 1965; Subhad-
rabandu, 1986). Foliar application of 50 mg/l BA with 2% calcium ion (Ca
2+
)
increased the proportion of perfect flowers (Singh and Rajput, 1990). Naphtha-
lene acetic acid (NAA) at concentrations of 50, 100 and 200 mg/l increased the
perfect/staminate flower ratio (Mallik et al., 1959; Singh et al., 1965).
Other factors influencing sex ratios of inflorescences include stem age
and mineral nutrients. Gunjate et al. (1983), Desai et al. (1986) and Hussein
et al. (1989) reported that inflorescences from stems that grew at different
times during the previous summer/autumn period had significantly differ-
ent perfect/staminate flower ratios. Singh and Dhillon (1987) found that
boron (B) levels affect sex ratio.
Sex ratios can be manipulated with growth regulators, but has no com-
mercial advantage (A.W. Whiley, personal communication, Queensland,
1996). Increases in fruit yield resulting from chemically increased perfect/
staminate flower ratios have not been observed, suggesting that perfect
flower numbers are not the primary limitation to crop performance (Schaffer
T.L. Davenport 136
et al., 1994). If only one or two fruits were set on each terminal, the tree would
carry an unusually heavy crop. It is unlikely that reduced perfect flower
numbers due to cool temperatures during inflorescence development is
directly responsible for poor fruit set and yields. Pollen viability, growth and
ovule fertilization are probably the main factors contributing to low fruit set
under these conditions.
Anthesis and dehiscence
Floral anthesis generally occurs at night in polyembryonic cultivars (Wagle,
1929; Torres, 1931; Galang and Lazo, 1937; Pimentel et al., 1984) and at night or
early morning in monoembryonic types (Popenoe, 1917; Musahib-ud-din and
Dinsa, 1946; Singh, 1954a; Randhawa and Damodaran, 1961a, b). Subsequent
dehiscence of the four-lobed anthers occurs during the daylight morning hours
revealing pale blue pollen grains (Torres, 1931; Galang and Lazo, 1937; Mallik,
1957; L.B. Singh, 1960). Anthesis and anther dehiscence are delayed by low
temperatures or overcast days (Singh, 1954a; De Wet and Robbertse, 1986a).
Dehiscence is also delayed by high RH, and pollination occurs primarily
around midday (Mallik, 1957; Randhawa and Damodaran, 1961a, b).
Stigmas are receptive from c.18 h prior to anthesis to at least 72 h after
anthesis with optimum receptivity within 3 h from anthesis (Popenoe, 1917;
Wagle, 1929; Sen et al., 1946; Singh, 1954a; Spencer and Kennard, 1956; Rand-
hawa and Damodaran, 1961a, b; Gunjate et al., 1983; Pimentel et al., 1984;
Robbertse et al., 1994). Receptive stigmas are shiny and white-green, whereas
non-receptive stigmas are desiccated and yellow-brown. Pollen germination
generally occurs within 90 min of deposition, although the percentage ger-
mination of pollen deposited on stigmas is relatively poor (Singh, 1954a; S.N.
Singh, 1961). Pollination is initiated by the formation of two unusual protu-
berances that meet to form a bridge or ponticulus connecting the dorsal side
of the ovule with the ovary wall in line with the base of the style (Joel and
Eisenstein, 1980). The ponticulus may guide the elongating pollen tube to the
ovule. Ovule fertilization occurs 48–72 h after pollination (S.N. Singh, 1961;
Ram et al., 1976). Both zygote cell and endosperm nuclear division appear to
rest for about 2 weeks following pollination despite cell division and growth
of the ovary (Sharma and Singh, 1972; Ram et al., 1976). A description of
embryo development is presented by U.R. Singh (1961). Application of B
may improve stigma receptivity, pollen tube germination and growth and
ovule fertilization (De Wet and Robbertse, 1986b; Robbertse et al., 1988; De
Wet et al., 1989) as well as fruit development (Chen, 1979; Robbertse et al.,
1990); however, Jutamanee et al. (2002) could not verify the effect of B.
Pollen
Pollen grains are 20–45 m long and are oblong when dry and more
spherical when hydrated (Popenoe, 1917; Jivanna Rao, 1923; Bijhouwer, 1937;
Reproductive Physiology 137
Mukherjee, 1950; Singh, 1954a; Randhawa and Damodaran, 1961b; S.N.
Singh, 1961). There are generally three equilateral, tapering furrows along
the longitudinal sides of dry pollen that give hydrated grains a roughly tri-
angular shape when viewed on end (Popenoe, 1917; Singh, 1954a; S.N. Singh,
1961; U.R. Singh and A.P. Singh, 1973). Each furrow has a germpore in its
centre (Mukherjee, 1950; S.N. Singh, 1961). Anthers produce c.250–650 pollen
grains with a mean of 410 grains per anther (Popenoe, 1917, 1920; Spencer
and Kennard, 1955).
In vitro germination of mango pollen has been reported (Popenoe, 1917;
Spencer and Kennard, 1955; Young, 1958; Randhawa and Damodaran, 1961b;
S.N. Singh, 1961). Germination on stigmas was c.10% less than that on artificial
media, that is 78% across cultivars (Spencer and Kennard, 1955), although
lower rates of germination have been reported (Mukherjee, 1950). Mango pol-
len is most viable soon after anther dehiscence and rapidly degrades (Sen et al.,
1946; Spencer and Kennard, 1955; Mallik, 1957; S.N. Singh, 1963). Although the
initial percentage of viable pollen is generally ≥ 90% during warm weather
(Popenoe, 1917; Mukherjee, 1949a, b; Singh, 1954a; S.N. Singh, 1961), cool tem-
peratures early in the flowering season result in abnormal, non-viable pollen
grains (Popenoe, 1917; U.R. Singh and A.P. Singh, 1973; Shu et al., 1989; Gazit
et al., 1992; Issarakraisila et al., 1992). The pre-vacuolate stage of meiosis during
microsporogenisis is the most sensitive period to temperatures < 10°C (Issara-
kraisila and Considine, 1994). Germination and pollen tube growth are reduced
by cool temperatures (S.N. Singh, 1961; Mullins, 1986; Robbertse et al., 1988;
Whiley et al., 1988; De Wet et al., 1989) and completely inhibited at tempera-
tures < 15°C (Popenoe, 1917; Young, 1955; Sukhvibul et al., 2000).
Pollination
Pollination is a major yield-limiting constraint, due to the large number of
flowers on trees and low fruit set. Unlike polyembryonic cultivars, which
produce nucellar embryos, pollination is necessary for fruit set with mo-
noembryonic cultivars (Popenoe, 1917; Young, 1942; Spencer and Kennard,
1955; Gunjate et al., 1983; Pimentel et al., 1984; Robbertse et al., 1994). Pollen
compatibility within and between cultivars has been widely investigated.
Complete or partial self-incompatibility has been reported (Mukherjee et al.,
1961; Singh et al., 1962b; Sharma and Singh, 1970, 1972; Ram et al., 1976; De
Wet et al., 1989; Robbertse et al., 1993). Incompatibility is evident by degen-
eration of embryonic and nucellar tissues and excessive loss of fruitlets. Cross
incompatibility between some cultivars has also been noted (Saha and Chhonkar,
1972; Ram et al., 1976; Robbertse et al., 1993).
Wind
Early investigators concluded that the species is wind pollinated (Hartless,
1914). Initially wet pollen dries to a powdery consistency on anthers soon
after anthesis in dry conditions (Pimentel et al., 1984), whence it is likely to be
liberated in moving air or via gravity to adjacent stigmas on the same and
T.L. Davenport 138
nearby flowers (Naik and Mohan Rao, 1943; Mallik, 1957). Singh (1954a) and
S.N. Singh (1961) suggested, however, that the amount of pollen moving in
air streams was too low for wind to be a pollination vector. They did not
report the location of pollen-collecting slides or take into account the close
proximity of flowers within inflorescences or numbers of open flowers in the
canopy. Panicles bagged to exclude pollinating insects were reported to set
fruit (Free and Williams, 1976), which were retained to maturity, thereby con-
firming that mango pollen can be transferred by air movement or gravity
(Bijhouwer, 1937; Mallik 1957). The tacit assumption that open-pollinated
flowers are exclusively crossed is likely to be incorrect, although mango may
favour cross-pollination.
Insect
Popenoe (1917) reasoned that pollen transfer occurs primarily within flowers
by insects. Panicles bagged to exclude insect visitation generally result in less
fruit set than on panicles in the open (Popenoe, 1917; Musahib-ud-din and
Dinsa, 1946; Mallik, 1957; Free and Williams, 1976; Jiron and Hedstrom, 1985).
Insects working mango flowers include Diptera, Hymenoptera, Lepidoptera
and Coleoptera (Popenoe, 1917; Simao and Maranhao, 1959; Randhawa and
Damodaran, 1961b; McGregor, 1974; Anderson et al., 1982; Jiron and Hed-
strom, 1985). Flies of various genera are common on mango flowers (Pope-
noe, 1917; Burns and Prayag, 1921; Bijhouwer, 1937; Singh, 1954a; Spencer
and Kennard, 1955; Eardley and Mansell, 1993). Polistes wasps are observed
on mango flowers but are considered to be ineffectual for pollen transfer
(Spencer and Kennard, 1955; Free and Williams, 1976; Wolfenbarger, 1977).
Honeybees (Hymenoptera) are occasional visitors (Young, 1942; Simao and
Maranhao, 1959; Smith, 1960; Morton, 1964; Jiron and Hedstrom, 1985; Mac-
Millan, 1991; Du Toit and Swart, 1993, 1994; Eardley and Mansell, 1993, 1994),
but only if other more inviting flowers are not present (Spencer and Kennard,
1955; Free and Williams, 1976; McGregor, 1976). They are assumed to be the
most effective pollinators of mango and may be more effective if hives are
placed in orchards during flowering (Du Toit and Swart, 1993, 1994). Ander-
son et al. (1982) recorded actual pollen transfer on mango flowers by insects
and found, in order of importance, the most efficient pollinators to be wasps,
bees, large ants and large flies.
With few exceptions (Mallik, 1957), pollen deposition rates are generally
low (Naik and Mohan Rao, 1943; Mukherjee, 1951). Differences in pollination
rates can be attributed to environmental conditions during flowering, differ-
ing attraction of insects to specific cultivars, proximity of more attractive
flowering species or a combination of the above. Young (1942) observed that
insects visit only 10–12% of available flowers. Depending on weather condi-
tions, insect activity on mango flowers is usually continuous from early
morning to late afternoon, but nocturnal activity of some species has also
been reported (Jiron and Hedstrom, 1985).
The role of insects in cross-pollination is not understood. Anderson et al.
(1982) observed various insects carrying pollen to and from flowers and
noted pollination subsequent to those visits; however, they made no distinction
Reproductive Physiology 139
between actual pollen depositions by visiting insects and pollen transferred
by other means. Wester (1920) considered that pollination is facilitated by
wind and to a lesser extent by insects, and this conclusion is probably correct
in most environments. Self-pollination within flowers by insects while the
pollen is still damp is likely to occur. Use of isozyme (Degani et al., 1990; Rob-
bertse et al., 1993) and microsatellite DNA markers (Adato et al., 1995; Schnell
et al., 2005) to discern ratios of cross- versus self-pollinated fruitlets and off-
spring is the most accurate procedure to confirm self- and cross-pollination.
Initial fruit set of pollinated flowers is inconsequential since most of these
fruitlets abscise before reaching maturity (Lynch and Mustard, 1950; Singh,
1954a; Randhawa and Demodaran, 1961a).
5.12 Fruit Development
Fruit growth is correlated with several growth regulating substances. Enlarge-
ment is sigmoidal reaching a constant size c.2–3 weeks before maturity
(Singh, 1954a; Randhawa and Damodaran, 1961a, b; Ram, 1983; Prakash and
Ram, 1984). The highest rates of fruit growth have been correlated with peak
levels of putative endogenous auxins found in seeds (Chacko et al., 1970a, b;
Singh and Singh, 1974; Chen, 1981; Ram, 1983; Prakash and Ram, 1984).
Baghel et al. (1987a, b) reported increased fruit mass with a combination
spray of NAA and urea to pre-anthesis panicles. Free and bound gibberellins,
especially in seeds (Ogawa, 1963; Ram, 1983), peak similarly to putative aux-
ins during fruit development (Chacko et al., 1970c, 1972a; Ram and Pal, 1979;
Chen, 1981). Cytokinins tentatively identified as zeatin and zeatin riboside
and other active fractions appear to vary in concentration in seeds and peri-
carp with two distinct peaks of activity. No particular relationship to growth
was evident (Ram, 1983; Ram et al., 1983). Seeds produced more cytokinin
activity than did the pericarp. In contrast, Chen (1983) found only one peak
of cytokinin activity in seeds and pulp occurring at about half full size of
both seed and pericarp. Seed tissues contain the highest cytokinin activity.
Levels of endogenous auxin, gibberellins and cytokinins in leaves during
fruit set were compared to production in leaves during other periods with-
out conclusive results (Paulas and Shanmugavelu, 1989).
5.13 Stenospermocarpy
Abscission of non-fertilized and fertilized flowers is normal. Fruitlet abscis-
sion from pea size on is often associated with embryo abortion (Chandler,
1958; U.R. Singh, 1961; Singh, 1964; Lakshminarayana and Aguilar, 1975;
Ram et al., 1976) and is referred to as stenospermocarpy (Soule, 1985). Steno-
spermocarpy in mango is unusual (Chacko and Singh, 1969a) but occurs
regularly in some cultivars (Núñez-Elisea and Davenport, 1983; Whiley et al.,
1988). Stenospermocarpic fruitlets have slower growth rates than seeded
fruit, generally become misshapen and fail to reach full size.
T.L. Davenport 140
Stenospermocarpy in some cultivars has been correlated with low tem-
peratures during flowering and early fruit set in the subtropics (Lakshmina-
rayana and Aguilar, 1975; Young and Sauls, 1979; Whiley et al., 1988; Schaffer
et al., 1994). Núñez-Elisea and Davenport (1983) reported that stenospermo-
carpic fruit often occur distal to seeded fruitlets within panicles and sug-
gested that embryo abortion is associated with high temperatures when these
latter fruit set. Secondary spring flowering of some monoembryonic culti-
vars under high temperatures has resulted in high proportions of steno-
spermocarpic fruit (E.K. Chacko, personal communication, Australia, 1995).
Application of auxins, gibberellins and cytokinins produce seedless fruit in
some cultivars, suggesting that the abscission zone is protected by these hor-
mones despite the loss of the endogenous supply from the aborted seed (Ven-
kataratnam, 1949; Chacko and Singh, 1969a, b; Kulkarni and Rameshwar,
1978).
5.14 Fruit Set and Retention
Fruit set and retention of mango was recently reviewed by Singh et al. (2005).
Abscission of flowers and fruitlets is accomplished by rapid formation of a
separation layer in the abscission zone in the pedicel-peduncle junction (Bar-
nell, 1939). U.R. Singh (1961) described formation of the abscission zone dur-
ing floral ontogeny and of the separation layer during abscission of flowers
and fruitlets. The majority of panicles lose all fruitlets (Núñez-Elisea and
Davenport, 1983). The pattern of fruitlet abscission is asymptotic with the
greatest losses occurring during the first weeks after anthesis (Núñez-Elisea
and Davenport, 1983; Prakash and Ram, 1984; Searle et al., 1995). Except for
the tendency to retain fruit in the distal portion of panicles, abscission of
flowers and fruitlets is random. It can involve fruitlets regardless of size or
location.
Of the 8–13% of perfect flowers setting fruit, < 1% reach maturity (Bijhou-
wer, 1937; Sen, 1939; Naik and Mohan Rao, 1943; Mukherjee, 1949b; U.R.
Singh, 1960; Randhawa and Damodaran, 1961a; Singh, 1978; Gunjate et al.,
1983; Prakash and Ram, 1984). Generally, most fruit are set on the most distal
spike portion of panicles (Chadha and Singh, 1963; Núñez-Elisea and Daven-
port, 1983). Fruit loss has been associated with embryo abortion, resulting in
blackened or shrivelled embryos (Singh, 1954a, 1964; Chandler, 1958; U.R.
Singh, 1961; Sharma and Singh, 1972; Ram et al., 1976) after the fruit is sepa-
rated from the tree (Núñez-Elisea and Davenport, 1983).
Sex ratio
The perfect/staminate floral ratio in panicles may influence fruit set and pro-
ductivity (Naik and Mohan Rao, 1943; Singh, 1954b; Singh and Singh, 1959;
U.R. Singh, 1960). Mallik (1957) noted that more perfect flowers are formed
in ‘on’ than ‘off’ years of alternate-bearing cultivars. Other studies, however,
Reproductive Physiology 141
have demonstrated that the number of perfect flowers does not correlate
with subsequent yield (Randhawa and Damodaran, 1961a) so long as the
proportion of perfect flowers is not < 4% (Singh, 1964, 1971). Most fruit are
borne in the distal portion of panicles (Shawky et al., 1977), which may be
correlated with the high ratio of perfect to staminate flowers there. Schole-
field and Oag (1984) estimated that one mature fruit is harvested for each 169
perfect flowers in the distal half of the panicle; whereas 592 perfect flowers
are required to produce one fruit in the proximal half. Therefore, intrinsic
factors other than sex ratio regulate fruit set.
Mineral nutrients
Boron is one of seven micronutrients required for normal plant growth. The
physiological function of B is unknown (Hu and Brown, 1994), although it is
essential for floral development, pollen germination, pollen tube growth,
embryo development and growth of organs (i.e. fruit) (Vasil, 1963; Agarwala
et al., 1981; Dell and Huang, 1997; Shorrocks, 1997). Deficient soils are com-
monly found in mango-producing areas of Australia, Thailand, Central and
South America and Africa where symptoms are common (Aitken et al., 1987;
Singh et al., 2005). Boron applications to deficient mango trees increase normal
fruit set (Robbertse et al., 1990; Raja et al., 2005). Fruitlet abscission in mangoes
has also been attributed to zinc (Zn) deficiency (Jiron and Hedstrom, 1985).
Hormonal control
Auxin
Research demonstrating improved fruit set and retention following applica-
tion of several auxin analogues to pre-anthesis panicles or to panicles bearing
fruitlets of various sizes has been reviewed (Davenport and Núñez-Elisea,
1997; Singh et al., 2005). NAA is the most effective auxin analogue for improv-
ing fruit retention (Prakash and Ram, 1986; Khan et al., 1993). Initial fruit set
was substantially increased when sprays of 200 mg/l indole acetic acid (IAA)
were applied to developing panicles (Singh et al., 1965). A 300–400% increase
in fruit set resulted when NAA (40 or 50 mg/l) was sprayed at the pre-anthesis
stage (Ram, 1983; Singh and Ram, 1983; Prakash and Ram, 1986). Chen (1981)
reported no effect on fruit retention when 5 mg/l of either naphthaleneacet-
amide or β-naphthoxoyacetic acid were applied three times at 2-week inter-
vals to panicles in which fruit had reached 4 mm in diameter.
Despite increased fruit retention of mango using exogenous applications
of auxins, few studies have examined endogenous auxins in fruit as related
to retention (Chacko et al., 1970a, b; Ram et al., 1983; Prakash and Ram, 1984).
Singh and Singh (1974) were unable to detect significant differences in endog-
enous auxins or inhibitors when comparing alternate and regular bearing
cultivars. Chen (1981) observed lower levels of auxin-like substances in
T.L. Davenport 142
mesocarp and calyx tissues of abscised fruits than those of intact fruits. Simi-
lar decreases in auxin and gibberellins with an increase in abscisic acid as
fruitlets abscised were reported by Bains et al. (1999). The interaction of auxin
in fruit and abscission zones to maintain mango fruit retention is not clear.
Continuous auxin synthesis and basipetal transport to the abscission
zone is critical for maintenance of plant organs, including fruit (Crane, 1964;
Nitsch, 1965; Morgan et al., 1977; Davenport et al., 1980; Roberts and Osborne,
1981). Increased mango fruit set and retention in response to exogenously
applied auxins confirms this requirement; however, other hormonal factors
also appear to be involved. Developing seeds are rich sources of all the known
classes of phytohormones, including auxins (Crane, 1964; Nitsch, 1965; Chacko
et al., 1970a, b, c; Chen, 1981). Hence, exogenous enrichment of auxin in the
presence of other seed-produced phytohormones facilitates increased fruit
retention. In contrast, NAA (10 and 20 mg/l) spray-applied to bagged, self-
pollinated flowers, does not result in development of stenospermocarpic
fruits beyond the marble size (Venkataratnam, 1949; Chacko and Singh,
1969a, b). Similarly, applications of 250 or 500 mg/l GA
3
or 250 mg/l BA
alone to panicles does not promote production of stenospermocarpic fruits
(Chacko and Singh, 1969a, b). Supplying exogenous β-naphthoxyacetic acid
(10 mg/l), BA (250 mg/l) and GA
3
(250 and 500 mg/l) together in multiple
sprays until half grown, however, resulted in retention of several seedless
fruit to maturity. Chen (1983) and Oosthuyse (1995b) observed that gibberel-
lin, cytokinin and auxin reduce fruit drop of open-pollinated fruitlets of some
cultivars. Thus, although auxin is important for maintaining the abscission
zone, the presence of other phytohormones appears to be important for fruit-
let development (Chacko et al., 1970a, b; Ram, 1983; Ram et al., 1983).
Cytokinins
Although cytokinins are not generally thought to be associated directly with
abscission, Ram (1983) and Ram et al. (1983) concluded that low cytokinin
levels during fruit development might contribute to fruit loss. Chen (1983)
observed a correlation of low cytokinin levels in stenospermocarpic fruits
with abscission at the marble stage of growth. Application of 250 mg/l BA to
bagged panicles does not promote production of seedless fruits (Chacko and
Singh, 1969a, b). The synthetic cytokinin, N-(2-chloro-4-pyridyl)-N’-phenylurea
(CPPU) also does not improve fruit set when applied alone at a rate of 10
mg/l to post-anthesis panicles (Oosthuyse, 1995b). The role of cytokinins in
separation events remains inconclusive.
Gibberellins
Gibberellins do not appear to be directly linked to the onset of abscission
(Chacko et al., 1970c, 1972a; Ram and Pal, 1979; Chen, 1981; Ram, 1983). Spray
applications of GA
3
to pre- and post-anthesis panicles to increase fruit set
and retention have been inconsistent. Increased yield (Teaotia et al., 1967;
Singh and Ram, 1983; Rajput and Singh, 1989) and production of seedless fruit
(Kulkarni and Rameshwar, 1978) have been reported from these treatments,
but Chacko and Singh (1969a, b) observed no such effects. Chen (1983) and
Reproductive Physiology 143
Oosthuyse (1995b) investigated the effects of several foliar applications of GA
3

starting at the 4 mm diameter stage, but were unable to improve fruit set.
Several classes of gibberellin-synthesis inhibitors have been tested for
reducing fruit drop. The growth retardants, daminozide and cycocel, increased
fruit set when applied to post-anthesis panicles (Singh and Ram, 1983). The
authors suggested that increased fruit retention might have been mediated
through increased cytokinin-like activity of the growth retardants. Although
initial fruit set was promoted by PBZ, yield was not improved (Kurian and
Iyer, 1993a). It is not clear whether the contrasting results of increased yield
(Kurian and Iyer, 1993b) were due to reduced fruit loss or more intense flow-
ering in response to treatment. Goguey (1990) reported increased fruit set
and retention using soil-applied PBZ at 5 g ai/tree. Spray application of uni-
conazole, a more biologically active triazole (500–2000 mg/l), reportedly
increased fruit set and yield (Galila and El-Masry, 1991). It is difficult to
resolve the contradictory results demonstrating enhancement of fruit reten-
tion by GA
3
and inhibitors of its synthesis.
Inhibitors
Abscisic acid (ABA) is possibly involved in fruitlet abscission. Although cor-
relations exist between certain inhibitors and abscission of mango fruitlets,
no clear cause and effect relationships have been established. Fruit drop was
correlated with levels of an acidic inhibitor, possibly ABA (Chacko et al.,
1970b, 1972a; Singh and Singh, 1974; Ram, 1983; Prakash and Ram, 1984).
Chen (1981) reported similar changes in putative ABA with maximum levels
occurring during early fruit drop and with advancing age of fruits. Putative
ABA levels in abscised and retained fruits were compared and were highest
in the calyx and mesocarp of abscised fruitlets.
Ethylene
Ethylene has the greatest immediate impact on flower and fruitlet abscission.
Van Lelyveld and Nel (1982) reported higher levels of ethylene in abscised
fruitlets compared with those retained on trees. Núñez-Elisea and Davenport
(1983, 1984, 1986) examined the dynamics of ethylene production in intact
and excised fruitlets from onset to separation. Increased production began in
explants about 26 h postharvest and increased logarithmically until fruit sep-
aration. Abscission of the fruitlets began 48 h after the onset of enhanced
ethylene production. Similar results with avocado fruitlet abscission experi-
ments (Davenport and Manners, 1982) indicate that the onset of ethylene
production in intact fruitlets is spontaneous in individual fruitlets followed
by abscission 48 h later. The pericarp provided the bulk of ethylene for induc-
tion of abscission processes; the pedicel produced no ethylene. There was
reduced fruit drop in response to inhibitors of ethylene production and action
(Singh and Ram, 1983; Naqvi et al., 1990, 1992). Whereas increased peroxi-
dase (Van Lelyveld, 1978) and polyphenol oxidase activities have been
reported in abscissed mango fruitlets (Van Lelyveld and Nel, 1982), Núñez-
Elisea and Davenport (1984) observed no changes in peroxidase activity or
protein levels prior to separation of fruitlets.
T.L. Davenport 144
Abscission of stenospermocarpic fruits has been associated with small
increments in ethylene production (Núñez-Elisea and Davenport, 1983). Sen-
sitivity to low levels of endogenous ethylene may reflect the absence of seed-
produced auxins. Protection of the abscission zone depends on a constant
supply of auxin, and ethylene production levels in tissues correlate with
endogenous auxin levels (Roberts and Osborne, 1981).
Despite their roles in cell division, cell enlargement and maintenance of
the abscission zone in developing fruit, a specific recommendation for exog-
enous application of plant growth regulators, either alone or in combination,
to improve yield of mango has not been adopted. Phytohormones have little
residual effect on fruit development, and multiple applications of products
to counteract the short-term responses are prohibitively expensive.
Photoassimilates
Wolstenholme and Whiley (1995) discussed the ecophysiology of the mango
as a basis for preharvest management. They proposed that the adaptive sur-
vival strategies of the mango explain its notoriously poor cropping perfor-
mance. Mechanisms that impart tolerance to heat, drought and flood stresses,
which the tree has developed for survival in harsh environments, have come
at considerable carbon cost with the resultant diversion of photoassimilate
resources away from fruiting.
There is abundant evidence that heavy cropping in tree crops exhausts
stored reserves (Jones et al., 1975; Kaiser and Wolstenholme, 1994; Whiley et al.,
1996) and that current photosynthate is often unable to satisfy the demands
of fruit set and fruit growth after heavy and prolonged flowering (Chacko
et al., 1982). There are significant genotypic differences in photoassimilation
rates between low- and high-yielding cultivars growing in both the tropics
and the subtropics of Australia (Chacko et al., 1995; Searle et al., 1995). At each
location, photoassimilation rates were considerably greater on the higher-
yielding cultivar, and this difference was maintained from flowering through
to fruit maturation.
14
C studies during the fruit set and abscission period also
demonstrated strong discrimination in the movement of assimilates, which
was dominated by randomly located fruit on panicles of the low-yielding
cultivar (Chacko et al., 1995). In contrast, assimilate discrimination to fruitlets
was less severe in the high-yielding cultivar with a more even distribution of
photoassimilates. It was concluded that the availability and distribution of
photoassimilates during the fruit set and establishment stages was largely
responsible for the yield differences between the cultivars.
Supporting evidence for the role of photoassimilates in fruit set and
retention also comes from enrichment studies (Schaffer et al., 1999). Container-
grown plants that flowered in the open were transferred to controlled-
environment glasshouse rooms immediately after the completion of anthesis.
Temperatures were 28°C day/20°C night while the atmospheric carbon diox-
ide (CO
2
) concentrations were 350 or 600 mol/mol. Photoassimilation of
trees in the CO
2
-enriched rooms was approximately 60% greater than those
held at partial pressures of 350 mol/mol CO
2
. Fruit retention and final yield
were significantly higher on those trees grown at the partial pressure of
Reproductive Physiology 145
600 µmol/mol CO
2
. Higher levels of available assimilates during the fruiting
cycle appear to benefit fruit retention and yield.
5.15 Alternate Bearing
Alternate bearing of certain mango cultivars has plagued growers (Hartless,
1914; Rao, 1997). Lack of production in the ‘off’ year is usually a result of lack
of floral initiation (R.N. Singh, 1959), and has been reviewed (Mukherjee,
1953; Gangolly et al., 1957; L.B. Singh, 1960, 1972; Chadha and Pal, 1986; Pan-
dey, 1989). Shoot initiation and induction described in this chapter perhaps
offers a clearer understanding of this phenomenon.
5.16 Conclusions
This chapter has provided a comprehensive review of investigations of vari-
ous factors potentially involved in mango flowering, fruit set and retention.
Many of the reports cited are contradictory. Such variable results reflect: (i)
the different experimental approaches utilized, especially in field experi-
ments; (ii) the range of environments in which experiments have been con-
ducted; and (iii) differences in the responses of cultivars to treatments. As a
consequence, it is difficult to draw unambiguous conclusions with respect to
the role of specific factors on flowering, fruit set and retention. More research
is clearly needed in these areas, particularly in controlled environments. For
example, although KNO
3
is utilized with great success to stimulate flowering
in tropical conditions, confusion remains as to whether it effects initiation or
floral induction. Future studies on flowering research should include results
of the proportion of stems that remain in rest and those that produce vegeta-
tive shoots as well as the proportion of reproductive shoots. Providing this
information allows an analysis of the impact of treatments on initiation and
inductive events.
This chapter also presents several hypothetical models for shoot devel-
opment and flowering. Each of the proposed models consists of a series of
hypotheses that invite further study to test their validity. Future research
should challenge these models so that flowering and crop yield can be better
understood in both the tropics and the subtropics.
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170 (ed. R.E. Litz)
6 Ecophysiology
B. Schaffer,
1
L. Urban,
2
P. Lu
3
and A.W. Whiley
4
1
University of Florida, Florida, USA
2
Centre INRA de Corse, San Giuliano, France
3
EWL Sciences, PO Box 39443, Winnellie, Northern Territory, Australia
4
Sunshine Coast Horticultural Services Pty Ltd, Nambour, Queensland, Australia
6.1 Introduction 170
6.2 Photosynthesis 172
Introduction 172
Light 177
Leaf temperature 179
Elevated atmospheric CO
2
concentration 181
Humidity 182
Flooding 182
Internal factors 184
Photosynthetic contributions by fruit 187
6.3 Plant Water Relations 188
6.4 Tree Growth and Development 190
Light 190
Temperature 192
Drought 193
Flooding 194
Wind 195
Salinity 196
Elevated atmospheric CO
2
concentration 197
6.5 Crop Production 198
Temperature limitations to crop production 198
Light interception and orchard design 199
6.6 Conclusions 200
6.1 Introduction
The genetic composition of mango cultivars is the primary determinant of
yield potential. However, actual yield, as well as tree growth and develop-
ment, are mediated by several endogenous factors including previous fruit
load, postharvest vegetative growth, preflowering maturity of terminal shoots,
Ecophysiology 171
production and mobilization of carbohydrates, nutritional status, plant
growth substances and carbon to nitrogen ratios (Schaffer et al., 1994). These
factors are either directly or indirectly affected by environmental variables
such as light, temperature and water availability. Environmental conditions
outside the range for optimum growth may also impose stress which results
in physiological changes that reduce growth or cause permanent damage to
mango trees. For example, major climatic events (i.e. extended drought,
floods, wind storms, heat waves and freezes) can cause severe damage to
crops due to development of excessive stress. However, mediated stress and
the release from stress imposed by normal seasonal changes provide condi-
tions that result in the progression of cropping cycles due to phenological
changes in the plants. An example of beneficial stress in mango is the
improved synchrony and reliability of flowering in subtropical climates due
to cool winter temperatures. Thus, understanding the impact of the environ-
ment on tree physiology and growth, and the particular adaptive strategies
developed through the processes of evolution, can provide a framework to
manage the crop to maximize genetic yield potential (Schaffer et al., 1994).
Physiological responses of mango to environmental variables can be
related to the evolutionary centre of origin of a specific cultivar. Mango culti-
vars are classified into two ecotypes based on embryony (see Mukherjee and
Litz, Chapter 1, this volume). A race with a single zygotic seed, monoembry-
onic types, evolved in the dry subtropical, monsoonal regions of the Indian
subcontinent with very hot summers but cooler winters. The polyembryonic
types, produced through nucellar embryony, largely evolved in the consis-
tently hot, humid tropics of South-east Asia where the monsoonal pattern
still predominates but the dry season is shorter than that of the Indian sub-
continent (Mukherjee, 1972). Hybridization occurs freely within and between
the two ecotypes and has led to a proliferation of cultivars of widely varying
genetic composition. Differences in growth and flowering responses to tem-
perature have been observed between the two embryonic ecotypes (Whiley
et al., 1989) and selection and breeding offer potential for increasing the crop-
ping performance of this notoriously low-yielding and recalcitrant tree across
a wider range of environmental conditions (Schaffer et al., 1994).
Inevitably, the many unique features of the mango tree represent its evo-
lutionary response to an indigenous environment that is particularly hostile,
with sustained extreme heat and high evaporative demand for much of the
year. This chapter provides an overview of the impact of environmental fac-
tors on physiology, growth and productivity of mango. Plant responses will
be considered in the context of the evolutionary origin and adaptability of
the mango tree. Responses to light, temperature and water are emphasized,
while the effects of atmospheric carbon dioxide (CO
2
) concentration, wind
and salinity are also discussed.
Photosynthesis and plant water relations are closely associated with
environmental conditions and directly affect plant growth and productivity.
The principles of leaf gas exchange and plant water relations are discussed to
provide a theoretical basis for interpreting physiological responses to the
environment. The impact of photosynthesis and tree water relations on
B. Schaffer et al. 172
growth and development under different environmental conditions is dis-
cussed. Pollination, fertilization, flowering and fruit set, which are strongly
influenced by environmental factors, have been addressed elsewhere (see
Davenport, Chapter 5, this volume; Schaffer et al., 1994). In the final sec-
tion of this chapter, mango crop production is integrated with aspects of
ecophysiology.
6.2 Photosynthesis
Introduction
The net CO
2
assimilation rate (A
net
) in C
3
plants is a function of the carboxyla-
tion rate (V
c
), the oxygenation rate (V
o
) and the rate of CO
2
evolution in light
that results from processes other than photorespiration, sometimes called
‘day respiration’ (R
d
):
A
net
= V
c
– 0.5V
o
– R
d
(6.1)
R
d
is usually inferred from measurements of leaf CO
2
exchanges after 5 min
in the dark (i.e. ‘night respiration’ R
n
). However, it has been repeatedly
shown that R
d
is lower than R
n
(see Atkin et al. (2000) for review), so that light
is known to inhibit respiration, with a R
d
/R
n
value ranging from 30 to 100%
(see Peisker and Apel (2001) for review). Urban et al. (2008) established the
following linear regression for R
d
of mango leaves: R
d
= 0.35R
n
– 0.21, which
may be used to infer R
d
from R
n
for photosynthetic photon flux (Q) values
above 170 mol photons/m
2
/s.
Currently, modelling of A
net
often uses the Harley et al. (1992) version
of the Farquhar et al. (1980) model. According to this model, A
net
can be
expressed as:
A
net
= (1 – 0.5O/(C
i
))min(W
c
, W
j
, W
p
) – R
d

(6.2)
where O represents the partial pressure of oxygen (O
2
) in the intercellular air
spaces (Pa), the specificity factor of ribulose-1,5-bisphosphate carboxylase/
oxygenase (Rubisco). C
i
is the partial pressure of CO
2
in the intercellular air
spaces (Pa), W
c
the carboxylation rate limited by the amount, activation state
or kinetic properties of Rubisco (mol CO
2
/m
2
/s), W
j
the carboxylation rate
limited by the rate of ribulose bisphosphate regeneration (mol CO
2
/m
2
/s),
and W
p
the carboxylation rate limited by triose phosphate utilization in
sucrose and starch synthesis (mol CO
2
/m
2
/s).
Usually O is set as 21 kPa (21%). The variable , which characterizes the
ratio of the affinities of CO
2
and O
2
for ribulose-1,5-bisphosphate in the active
site of Rubisco, can be calculated from the CO
2
compensation point * (the
CO
2
concentration at which photosynthesis equilibrates with respiration):
= 0.5O/* (6.3)
where = 2220 mol CO
2
/mol O
2
at 25°C for ‘Cogshall’ mango leaves (Urban
et al., 2008), which is lower than those given by Epron et al. (1995): 2100–2900
Ecophysiology 173
mol CO
2
/mol O
2
. Rubisco’s large subunit is encoded by a single gene in the
chloroplast genome, and no post-transcriptional modifications have been
discovered so far. It is thus very unlikely that can change in the short term
(Spreitzer and Salvucci, 2002).
The internal partial pressure of CO
2
(C
i
) is one of the two major variables
of photosynthesis (with the photosynthetically active photon flux density).
It may be calculated from the supply function:
C
i
= C
a
– A
net
/g
b
– A
net
/g
s
(6.4)
where C
a
is the partial pressure of CO
2
(Pa) in ambient air, g
b
represents the
leaf boundary layer conductance (mol H
2
O/m
2
/s), and g
s
is the stomatal
conductance of water (H
2
O) (mol H
2
O/m
2
/s).
Stomatal conductance is the major factor controlling A
net
. It ranges from
c.0.02 to c.0.4 mol H
2
O/m
2
/s in ‘Cogshall’ mango leaves and may be linearly
related to A
net
(Urban et al., 2002, 2003, 2006). The slope of the relationship
between g
s
and A
net
however is affected by drought (Fig. 6.1). Variations in
the slope of this relationship reflect changes in photosynthetic water use effi-
ciency and are not well understood.
It must be stressed that using C
i
as the driving variable of photosynthesis
is much debated. It has been advocated that C
c
, the partial pressure of CO
2
at
the site of carboxylation, should be utilized instead. Using C
i
implies that the
following assumptions have been made: C
c
= C
i
and g
m
= 0, where g
m
repre-
sents mesophyll conductance, also called liquid phase resistance, which
Dry
y = 0.009x + 0.024
R
2
= 0.695
0
0.05
0.10
0.15
0.20
0.25
0.30
0.35
0.40
0.45
0.50
0 5 10 15 20
A
net
(µmol CO
2/
m
2
/
s)
g
s

(
m
o
l

H
2
O
/
m
2
/
s
)
Wet
y = 0.025x + 0.028
R
2
= 0.86
Fig. 6.1. The relationship between stomatal conductance (g
s
) and net photosynthesis
(A
net
) in mango leaves from well-irrigated (■) and drought-stressed (▲) 12-year-old
‘Cogshall’ trees (Source: redrawn from Urban et al., 2006).
B. Schaffer et al. 174
encompasses diffusion from the intercellular leaf spaces to the carboxylation
sites in the chloroplasts. There is a growing body of evidence that g
m
is not
negligible in most species. The average value of g
m
in unstressed mango
leaves (0.21 mol CO
2
/m
2
/s) (Urban et al., 2008), calculated using the method
of Epron et al. (1995), is within the range of values for broadleaf species sur-
veyed by Ethier and Livingston (2004) and Manter and Kerrigan (2004). The
carboxylation rate (in Eqn 6.2) limited by the amount, activation state or
kinetic properties of Rubisco (W
c
) can be calculated as:
W
c
= V
cmax
C
i
/(C
i
+ K
c
(1 + O/K
o
)) (6.5)
where V
cmax
represents the maximum rate of carboxylation (mol CO
2
/
m
2
/s), and K
c
(Pa CO
2
) and K
o
(Pa O
2
) are the Michaelis constants of Rubisco
carboxylation and oxygenation, respectively. The V
cmax
values of well-
exposed mango leaves at a leaf temperature of 30°C are typically in the range
of 80–100 mol CO
2
/m
2
/s (Urban et al., 2006). Specific values of K
c
and K
o
for
mango leaves have not been estimated and are approximated using data
from other species (i.e. cotton or tobacco).
The carboxylation rate limited by the rate of ribulose bisphosphate regen-
eration (W
j
) is controlled by the rate of electron flow J (mol electrons/m
2
/s):
W
j
= JC
i
/(4(C
i
+ O/)) (6.6)
with
J = Q/(1 +
2

2
Q
2
/J
max
2
)
0.5
(6.7)
where Q is the photosynthetically active photon flux density (mol quanta/
m
2
/s), represents leaf absorbance (no units), is the apparent efficiency of
light energy conversion (mol electrons/mol photons) and J
max
is the light-
saturated rate of electron transport (mol electrons/m
2
/s). Leaf absorbance
of mango leaves, measured from 390–760 nm using an integrating sphere,
was found to be close to 0.81 (Urban et al., 2008) and is in the normal range of
values of the literature (Bauerle et al., 2004). Leaf absorbance, which is pos-
itively correlated with leaf chlorophyll content, may increase as a conse-
quence of paclobutrazol treatments (Gonzalez and Blaikie, 2003). The apparent
efficiency of light energy conversion in mango reaches 0.32 mol electrons/
mol photons (Urban et al., 2004b), in the absence of photoinhibition or pho-
todamage. This value corresponds to the mean value of operational (Sin-
gaas et al., 2001). The J
max
values of well-exposed mango leaves at a leaf
temperature of 30°C are typically in the 120–150 mol CO
2
/m
2
/s range. The
J
max
as well as the V
cmax
values are rather low when compared to values from
other species and partly explain why maximal rates of leaf photosynthesis
(A
max
) are rather low, typically 12–15 mol CO
2
/m
2
/s.
The carboxylation rate limited by triose phosphate utilization during
sucrose and starch synthesis (W
p
in Equation 2), can be calculated by:
W
p
= 3TPU + V
o
/2 = 3TPU + V
c
0.5C
i
/(C
i
) (6.8)
where TPU is the rate of phosphate release in triose phosphate utilization
during starch and sucrose production. The TPU is usually not included in
Ecophysiology 175
most studies on photosynthetic capacity because of methodological difficul-
ties. However, Urban et al. (2003) found that TPU = 8–12 mol CO
2
/m
2
/s at
a leaf temperature of 30°C in well-exposed ‘Cogshall’ mango leaves.
The variables V
cmax
and J
max
are temperature dependent and their depen-
dency is described by:
Parameter (V
cmax
, J
max
, TPU) = exp(c – H
a
/(RT
l
))
/(1+exp((ST
l
– H
d
)/(RT
l
))) (6.9)
where c is a scaling factor, H
a
(J/mol) the activation energy of the given
parameter, R the gas constant (8.3143 J/°K/mol), T
l
(°K) the leaf temperature,
S (J/mol) an entropy term and H
d
(J/mol) the deactivation energy of the
given parameter.
Similarly, the temperature dependency of R
d
, , K
c
and K
o
is described
by:
Parameter (R
d
, , K
c
, K
o
) = exp(c – H
a
/(RT
l
)) (6.10)
Proteins of the Calvin cycle and thylakoids represent the majority of leaf
nitrogen (N). Therefore, photosynthetic capacity is strongly related to leaf N
content expressed on an area basis (N
a
) (Field and Mooney, 1986; Evans, 1989;
Kellomäki and Wang, 1997; Walcroft et al., 1997). To account for the relation-
ship commonly observed between the parameters defining photosynthetic
capacity (V
cmax
, J
max
, TPU and R
d
mainly) and N
a
(Field and Mooney, 1983;
Harley et al., 1992) (Fig. 6.2), scaling factors c of V
cmax
, J
max
, TPU and R
d
may
be related to N
a
, either linearly or slightly non-linearly.
In summary, leaf net photosynthesis depends on five major classes of
factors, either variables (external or internal factors) or parameters (more or
less constant factors), provided that plants are not exposed to too extreme
conditions; we may consider the internal factors as genetic factors. The five
classes of factors are:
The photosynthetically active photon flux density ( 1. Q), which is the major
driving variable of photosynthesis. Gross photosynthesis is determined by
Q while C
i
determines the proportion of photorespiration, and thus net
photosynthesis. One of the major environmental factors affecting C
i
is water
availability in the root zone through its effect on g
s
.
Leaf nitrogen concentration ( 2. N
a
), which is not a rate-determining factor
of photosynthesis, unlike Q, but may be considered as a rate-limiting factor.
In other words, N
a
sets the photosynthetic potential of a leaf (i.e. photosyn-
thetic capacity). We shall see below which factors influence N
a
in mango
leaves.
Leaf temperature influences leaf photosynthesis. Net photosynthesis is 3.
positively correlated with leaf temperature in a normal range. Leaf tempera-
ture (T
l
) is not a driving variable of photosynthesis but it is the single most
important rate-determining factor after Q. In addition, extreme temperatures
may influence photosynthesis through their damaging effects. Kinetics of
enzymes involved with photosynthetic reactions collectively comprise an
additional set of factors that influence leaf net photosynthesis.
B. Schaffer et al. 176
Several parameters related to enzymes include the specificity factor of 4.
Rubisco (), the Michaelis constants of Rubisco carboxylation and oxygen-
ation, K
c
and K
o
, the activation and deactivation energies of the different pa-
rameters H
a
and H
d
, the entropy terms S, c factors and leaf absorbance
(). With the exception of K
c
and K
o
, the specific values of all these parame-
ters have been estimated for ‘Cogshall’ mango (Urban et al., 2003).
The apparent efficiency of light energy conversion ( 5. ). This factor be-
longs to a category of its own since it should theoretically not differ from one
y = 41.52x – 15.52
R
2
= 0.87
y = –201.64x
–1
+ 173.41
R
2
= 0.88
0
20
40
60
80
100
120
140
(a)
(b)
1.0 1.5 2.0 2.5 3.0 3.5 4.0
N
a
(g N/m
2
)
V
c
m
a
x


(
µ
m
o
l

C
O
2
/
m
2
/
s
)
0
50
100
150
200
250
1.0 1.5 2.0 2.5 3.0 3.5 4.0
y = 66.94x – 15.40
R
2
= 0.83
y = –330.44x
–1
+ 291.55
R
2
= 0.86
J
m
a
x


(
µ
m
o
l
/
m
2
/
s
)
N
a
(g N/m
2
)
Fig. 6.2. Relationship between (a) the maximum rate of carboxylation (V
cmax
) and (b)
the light-saturated rate of electron transport (J
max
), and nitrogen concentration per unit
leaf area (N
a
). Measurements were performed on mango leaves of 3-year-old ‘Cogshall’
trees (●), standard leaves () and leaves close to developing fruits () of 11-year-old
‘Cogshall’ trees. Best fit lines for pooled data correspond to the linear (
_
) and the ax
–1
+
b (

) models (Source: redrawn from Urban et al., 2003).
Ecophysiology 177
species to another and may be considered as a constant in the absence of
photoinhibition and photodamage.
Light
Light exposure
Plants allocate nitrogen resources within the canopy to enhance photosyn-
thetic capacity at locations exposed to high incident light levels, thus maxi-
mizing whole plant carbon gain (Field and Mooney, 1983; Hollinger, 1996;
Carswell et al., 2000). For leaves of a given age and for a given nitrogen sup-
ply, leaf N per unit leaf area appears to be strongly related with light expo-
sure (DeJong and Doyle, 1985; Le Roux et al., 1999, 2001; Rosati et al., 1999,
2000). Photosynthetic light acclimation of leaves may result from changes in
either leaf nitrogen concentration (N
m
) or mass-to-area ratio (M
a
) because
N
a
= M
a
N
m
. Lynch and González (1993) observed a negative correlation
between N
m
and light exposure in the tropical fruit tree Borojoa patinoi, but
such a behaviour is rare; positive correlations between N
m
and light exposure
are more commonly observed. In addition, photosynthetic light acclimation
of leaves may result from changes in partitioning of total leaf N among the
different pools of the photosynthetic machinery (Evans, 1989). In mango,
light acclimation of photosynthesis results mainly from changes in M
a
, and
to a lesser extent from changes in allocation of total leaf N at low irradiance;
whereas changes in N
m
play only a minor role (Fig. 6.3). Light acclimation of
mango leaves thus follows a pattern similar to peach leaves (Le Roux et al.,
1999; Walcroft et al., 2002).
Light intensity
Photosynthesis of ‘Cogshall’ mango trees increases with increasing levels of
light intensity to reach a maximum at Q = 1200 mol photons/m
2
/s (L.
Urban, unpublished data). Whiley et al. (1999) measured Q at 1284 mol pho-
tons/m
2
/s for field-grown ‘Kensington Pride’ trees growing in subtropical
Queensland, Australia, which is well below full sunlight (full sunlight ≥ 2000
mol photons/m
2
/s). Such a high threshold is a typical feature of sun plants.
Individual leaves are rarely able to utilize full sunlight; whole trees consist of
many leaves that shade each other, so that only a small fraction of a tree’s
leaves are exposed to full sun at any given time of the day, while the rest of
the leaves receive subsaturating photon fluxes in the form of small patches of
light that penetrate through gaps of the leaf canopy. Because the photosyn-
thetic response of whole trees is the sum of the photosynthetic activity of all
the leaves, only rarely is photosynthesis saturated with light at the whole-
tree level.
While most leaves experience subsaturating light intensities, well-exposed
leaves of the upper-crown may receive excessive quantities of light. Those
leaves must dissipate the absorbed light energy in excess to prevent damage to
the photosynthetic apparatus. Moderate decreases in maximal quantum
efficiency (i.e. quantum efficiency of dark-adapted leaves F
v
/F
mPredawn
) are
B. Schaffer et al. 178
typical features of moderate photoinhibition and should be interpreted in
terms of non-photochemical quenching, an adaptative mechanism involving
the xanthophyll cycle and allowing excess energy to be dissipated in the form
of heat (Adams et al., 2005). Such small decreases in F
v
/F
mPredawn
are com-
monly observed in mango leaves even from well-irrigated trees (Urban and
Alphonsout, 2007).
When temperature (30°C) and water vapour pressure deficit (VPD < 1
kPa) are non-limiting, and in the absence of photoinhibition, maximal rates
of net leaf photosynthesis (A
max
) may reach 12–15 mol CO
2
/m
2
/s at saturating
0.0
1.0
2.0
3.0
0.0 0.2 0.4 0.6 0.8 1.0
Gap fraction
N
m

(
g

N
/
g

d
r
y

m
a
t
t
e
r
)
Tree # 1
y = 1.42x + 1.43
R
2
= 0.90
0.0
0.5
1.0
1.5
2.0
2.5
3.0
0.0 0.2 0.4 0.6 0.8 1.0
Gap fraction
N
a

(
g
/
m
2
)Tree # 2
y = 1.33x + 1.44
R
2
= 0.79
(a)
(b)
Fig. 6.3. Relationship between (a) leaf nitrogen concentration per unit mass (N
m
) and
(b) leaf nitrogen concentration per unit leaf area (N
a
) and the gap fraction for mango
leaves measured in the crown of two 3-year-old ‘Cogshall’ trees. Gap fractions were
measured as an indicator of light exposure. Measurements were performed on leaves
< 2 months old (●), 8 months old (■), 12–14 months old (▲) and 17–20 months old
(♦) (Source: Urban et al., 2003).
Ecophysiology 179
Q, on ‘Cogshall’ trees. Whiley et al. (1999) measured A
max
of 15.2 mol CO
2
/
m
2
/s for ‘Kensington Pride’ trees growing in a subtropical climate in Queen-
sland, Australia. However, values > 16 mol CO
2
/m
2
/s were observed on
field-grown trees of ‘Tommy Atkins’, ‘Haden’ and ‘Irwin’ on sunny days
during the wet season in tropical regions of Australia (P. Lu, unpublished
data). This is much higher than citrus (< 10 mol CO
2
/m
2
/s), but substan-
tially lower than plum (approx. 26 mol CO
2
/m
2
/s). Whiley et al. (1999) esti-
mated the light compensation point to be 29 mol photons/m
2
/s for leaves
of non-stressed, field-grown mango trees, which is much higher than that
attributed to shade-tolerant species (< 10 mol photons/m
2
/s) (Harvey,
1979). The data show that mango trees are basically sun-adapted plants.
Leaf temperature
Effect of temperatures in a normal range on leaf photosynthetic capacity
Medlyn et al. (2002) calculated optimal temperatures for V
cmax
and J
max
to be
35–41°C and 30–38°C, respectively. Tree species native to cold climates had
the lowest temperature optima for both V
cmax
and J
max
. For ‘Cogshall’ mango
trees, calculated temperature optima for V
cmax
and J
max
are 44 and 45.5°C,
respectively. They demonstrated that mango photosynthesis increases with
temperature well above 40°C (Fig. 6.4). Although there is a clear lack of refer-
ences for other tropical trees, it is tempting to attribute the temperature
response of mango photosynthesis to its tropical origin.
Estimates of H
a
, H
d
and S are 7.0695, 17.0799 and 536 J/mol for V
cmax
,
and 3.8782, 10.2211 and 317 J/mol for J
max
, respectively. Both H
d
and S are
within the range of published values for V
cmax
and J
max
(Dreyer et al., 2001).
In addition, H
a
for V
cmax
is within the 60–80 kJ/mol range for many species,
including crop species as well as deciduous and evergreen trees (Medlyn
et al., 2002). For mango, H
a
for J
max
is consistent with data published for
evergreen species (Medlyn et al., 2002), which is consistent with the fact that
mango leaves commonly are 2–4 years old before senescence and abscision.
However, the J
max
/V
cmax
at 25°C for mango is about 1.86, approximately 11%
higher than the mean value calculated over the whole range of species stud-
ied by Medlyn et al. (2002). Lower activation energies for J
max
than V
cmax

result in a temperature-induced decrease in J
max
/V
cmax
, confirming previous
observations by Walcroft et al. (1997) and Dreyer et al. (2001). The estimate of
H
a
for R
d
is 4.5710 J/mol. H
a
is higher than the range of published values
for R
d
(Dreyer et al., 2001).
Chilling temperatures
F
v
/F
mPredawn
decreases in mango leaves with decreasing temperature, while
chilling reduces quantum efficiency (Whiley et al., 1999; Sukhvibul et al.,
2000; Weng et al., 2006a, b). The decrease in F
v
/F
mPredawn
may be interpreted
to reflect sustained engagement of zeaxanthin in photoprotective energy dis-
sipation. Decreases in quantum efficiency correspond to a decrease in the
rate of electron flow. It may be argued that sustained zeaxanthin-dependent
B. Schaffer et al. 180
energy dissipation reduces the risk of formation of singlet oxygen
1
O
2

in the
antennae, while decreases in J lower the risk of electrons reducing O
2
to anion
superoxide O
2

in the photosynthetic electron transport chain (Adams et al.,
2005). In other words, decreases in F
v
/F
mPredawn
and quantum efficiency cor-
respond to adaptative mechanisms against the effect of cold, when photosyn-
thesis is low and there is an imbalance between the quantity of light energy
absorbed and the quantity of energy used in the photochemical reactions of
0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
4.0
(a)
(b)
15 20 25 30 35 40 45 50
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
1.8
2.0
15 20 25 30 35 40 45 50
T
l
(°C)
J
m
a
x
/
J
m
a
x

a
t

2
5
°
C
V
c
m
a
x
/
V
c
m
a
x

a
t

2
5
°
C
T
l
(°C)
Fig. 6.4. Temperature response functions adjusted to the (a) maximal rate of car-
boxylation (V
cmax
) and (b) the light-saturated rate of photosynthetic electron flux (J
max
),
normalized to the mean value at 25°C in leaves from ‘Cogshall’ mango seedlings.
The data scatter represents the real scatter at each temperature. Reference values at
25°C were computed for each of eight leaves, taken from young trees from two
origins (● and ), and a unique temperature response was adjusted over the range of
normalized data. T
l
, the leaf temperature; the dotted lines correspond to Equation 9;
the solid lines correspond to Equation 10 (no deactivation energy component).
Ecophysiology 181
photosynthesis (Adams et al., 2005). Interestingly, Weng et al. (2006b) found
that mango leaves transferred from warm and dark to chilling conditions
showed only slight down-regulation of PSII efficiency when compared to
leaves moved from dim light to chilling conditions. Of course, long-term
exposure to cold and very low temperatures (≤ 10°C) may eventually
result in true photodamage, not just photoinhibition. Very low values of F
v
/
F
mPredawn
, decreases in chlorophyll content and slow recovery kinetics are all
indicators of photodamage. Sukhvibul et al. (2000) observed that susceptibil-
ity to cold-induced photodamage was more pronounced in polyembryonic
cultivars than in monoembryonic cultivars, possibly reflecting their different
eco-evolutionary development.
Elevated atmospheric CO
2
concentration
The CO
2
concentration in the earth’s atmosphere has been increasing rapidly
since the early 20th century and is continuing to rise, primarily due to burn-
ing of fossil fuels (Houghton, 2005). Earth’s atmospheric CO
2
concentration
is currently about 370 mol CO
2
/mol (Houghton, 2005) and is projected to
reach 600 mol CO
2
/mol by 2050. Elevated ambient CO
2
levels will undoubt-
edly affect cropping systems since atmospheric CO
2
concentrations can sig-
nificantly affect plant growth and productivity (Idso and Kimball, 1991;
Houghton, 2005). There is little published information concerning the effects
of elevated ambient CO
2
levels on physiology, growth and production of
tropical fruit trees, including mango.
Schaffer et al. (1997) exposed leaves of field- and container-grown ‘Kens-
ington’ (syn. ‘Kensington Pride’) trees to short durations (several minutes) of
varying ambient CO
2
concentrations. They found that under saturating light
levels for photosynthesis, net photosynthesis increased as ambient CO
2
con-
centration increased up to 1200 mol CO
2
/mol. At ambient CO
2
concentra-
tions > 1200 mol CO
2
/mol, net photosynthesis stabilized, probably due to
leaves reaching their maximum biochemical capacity to fix carbon. Studies
with ‘Cogshall’ mango trees indicated that when C
a
increases, stomata close
swiftly and C
i
may become very unpredictable (L. Urban, unpublished data).
Therefore, using C
i
may be preferable to C
a
for quantifying short-term effects
of elevated CO
2
concentations on A
net
of mango. Saturating CO
2
levels may
often be reached at C
i
= 800 mol CO
2
/mol air.
Long-term (6–12 months) exposure of ‘Kensington’ mango trees to an
atmospheric CO
2
concentration of 700 mol/mol resulted in higher net CO
2

assimilation rates than in leaves of plants grown at atmospheric CO
2
concen-
trations of 350 mol/mol when net CO
2
assimilation was measured at the
same CO
2
concentration as the growth environment. However, carboxylation
efficiency (the amount of CO
2
fixed per mole of ambient CO
2
) was lower for
plants in the CO
2
-enriched environment compared to plants in the ambient
(350 mol CO
2
/mol) environment (Schaffer et al., 1997). Although further
studies are needed to determine the effects of long-term exposure to elevated
CO
2
concentrations on mango growth and productivity, it appears that
B. Schaffer et al. 182
mango will benefit from increases in atmospheric CO
2
concentrations. How-
ever, the effects of increased atmospheric CO
2
concentrations associated with
global warming on mango production may be offset by higher respiratory
losses and increased assimilate partitioning to shoot growth in highly vegeta-
tive cultivars (Schaffer et al., 1999). Therefore, responses of mango cultivars
to elevated atmospheric CO
2
concentrations need to be evaluated over a
range of temperatures to ascertain their likely performance under changing
atmospheric conditions.
Humidity
Although mango production occurs in the tropics and subtropics in areas of
high and low relative humidity (RH) (Campbell, 1984), there are very few
published reports on the effects of RH or VPD on physiology and tree
growth.
In a study with container-grown ‘Kensington’ plants, Pongsomboon et al.
(1992) reported that stomatal conductance was inversely correlated with
VPD. Differences between cultivar responses to VPD have been observed in
field-grown ‘Irwin’ (monoembryonic) and ‘Kensington’ (polyembryonic)
mango trees during the wet and dry season in tropical Australia. During the
wet season and for well-irrigated trees during the dry season, both ‘Irwin’
and ‘Kensington’ showed decreasing stomatal conductance with increasing
leaf-to-air vapour pressure deficit (LAVPD) but ‘Kensington’ showed a more
rapid decrease than ‘Irwin’ (Fig. 6.5) (P. Lu, unpublished data). It was also
observed that daytime leaf xylem water potential was lower in ‘Irwin’ than
in ‘Kensington’ while predawn water potentials were similar for both culti-
vars (P. Lu, unpublished data). These results indicate that under similar soil
water conditions, ‘Kensington’ tends to close stomata much more rapidly
than ‘Irwin’ to conserve water under dry atmospheric conditions. This water
conservation strategy is probably a reflection of ‘Kensington’s’ adaptive
responses to the hot and dry seasonal tropical environment under which it
evolved (Wolstenholme and Whiley, 1995). Other studies in tropical Austra-
lia revealed that polyembryonic ‘Nam Doc Mai’ behaved like ‘Irwin’ (P. Lu,
unpublished data). However, ‘Nam Doc Mai’ in Thailand has comparatively
low vigour compared to ‘Kensington’ when grown in the tropics.
Further research is required to determine if differences in photosynthetic
or stomatal responses of mango to VPD are indeed based on embryonal char-
acteristics. Clarification of the reasons for variation would undoubtedly facil-
itate breeding and selection of cultivars for dry and humid areas.
Flooding
The primary effect of flooding on plants is due to a reduction in soil oxygen
concentration. Oxygen levels in the soil can decrease from 20% to < 5% within
1–2 days of flooding (Crane and Davies, 1988) and soils eventually become
Ecophysiology 183
anoxic (no oxygen). Mango is considered to be a moderately flood-tolerant
species (Schaffer et al., 1994, 2006) and waterlogging or flooding of trees peri-
odically occurs in many of the regions where the crop is grown (Plate 41).
Mango trees have evolved a mechanism to cope with temporary flooding (see
‘Flooding’ section under Tree Growth and Development, this chapter).
Typically, the first easily measurable responses of fruit trees to flooding
are reductions in A
max
, g
s
and transpiration, which occur within 2–3 days fol-
lowing flooding (Larson et al., 1991c; Schaffer et al., 1992, 2006). Short-term
anoxia results in a decrease in net photosynthesis which cannot be related to
a g
s
-associated decrease in C
i
(Zude-Sasse et al., 2001). Removing trees from
flooded conditions after 28 days reversed the flooding-induced decrease in
leaf gas exchange, resulting in a gradual increase in photosynthesis and tran-
spiration to preflooded rates.
Although flooding adversely affects mango trees, short-term flooding of
trees in limestone soils can result in increased micronutrient availability with
improved plant nutritional status. In calcareous soils of south Florida, in
which iron (Fe) was withheld from the fertilizer programme, short-term
flooding (10–20 days) of polyembryonic ‘Peach’ mango trees resulted in an
increase in net photosynthetic rates to above preflooding levels following the
release of trees from flooding (Larson et al., 1992). This increase in photosyn-
thesis has been correlated with improved Fe and manganese (Mn) uptake as
1 2 3 4 5 6 7 8
0.0
0.1
0.2
0.3
0.4
0.5
0.6
‘Kensington’: R
2
= 0.635
‘Irwin’: R
2
= 0.594
S
t
o
m
a
t
a
l

c
o
n
d
u
c
t
a
n
c
e

(
m
o
l
/
m
2
/
s
)
LAVPD (kPa)
Fig. 6.5. Correlation between leaf stomatal conductance and leaf-to-air vapour
pressure deficit (LAVPD) during the dry and wet season for ‘Irwin’ (closed circles)
and ‘Kensington’ mango trees (open circles). Trees were well irrigated during the dry
season and all measurements were taken when the Q > 300 mol photons/m
2
/s
(n = 12) (Source: P. Lu, unpublished data).
B. Schaffer et al. 184
a result of these elements becoming more soluble when calcareous soils are
flooded (Larson et al., 1991b, 1992).
Internal factors
Leaf age
Leaf characteristics (i.e. photosynthetic capacity and the amount of N per
unit area) are generally strongly influenced by leaf age, with maximum val-
ues being observed when leaves have just completed full expansion (Con-
stable and Rawson, 1980; Marshall and Biscoe, 1980; Dwyer and Stewart,
1986; Field, 1987; Wilson et al., 2000; Frak et al., 2001). Chlorophyll content is
three to four times lower in young than in mature mango leaves (Zude and
Ludders, 1997). Similarly, the concentration of Rubisco is lower in young
than in mature, green leaves (Nii et al., 1995). In contrast to many other plant
species, once mango leaves are mature the relationship between N
a
and irra-
diance does not seem to be affected by leaf age (Urban et al., 2003). The N
a

values may remain high in old leaves experiencing high irradiance. This
indicates that changes in N
a
in mango leaves are influenced by irradiance
and not age, at least during the first year.
Carbohydrate accumulation and source-sink balance
Source-sink imbalances can exert feedback down-regulation or repression of
leaf photosynthesis through carbohydrate accumulation in leaves (Azcon-
Bieto, 1983; Foyer, 1988; Koch, 1996; Schaffer et al., 1997; Whiley et al., 1999;
Paul and Foyer, 2001; Paul and Pellny, 2003). Transient accumulations of car-
bohydrates in leaves, as they have been observed during the diurnal period,
may impair the rate of electron transport (Pammenter et al., 1993). Changes
in photosynthetic capacity, not just assimilation rates, are more likely to be
observed in association with lasting source-sink imbalances. One hypotheti-
cal mechanism is that high levels of carbohydrates repress the expression of
genes coding for several photosynthetic enzymes (Krapp and Stitt, 1995;
Koch, 1996; Drake et al., 1997). Alternatively, carbohydrates may interact with
hormonal signals to control gene expression (Thomas and Rodriguez, 1994).
There is also some evidence that photosynthetic capacity is related to leaf
carbohydrate status through the effect of the latter on phosphate availability
(Riesmeier et al., 1993; Sun et al., 1999). In the long term, carbohydrate accu-
mulation may eventually lead to cell death. High sugar concentration has
been associated with senescence in leaves of several species (Noodén et al.,
1997; Wingler et al., 1998; Quirino et al., 2001). Reduced energy utilization by
CO
2
assimilation, like the one resulting from carbohydrate accumulation, in
combination with high energy capture is potentially dangerous and can
result in over-reduction of the electron transport chain, photoinhibition and
oxidative stress caused by photoreduction of oxygen to superoxide O
2

in the
Mehler-ascorbate peroxidase reaction (Badger, 1985). Moreover, reactive sin-
glet oxygen
1
O
2
can be formed through reaction of oxygen with triplet chlo-
rophyll released by the breakdown of the chlorophyll-protein complexes in
Ecophysiology 185
thylakoids (Merzlyak and Hendry, 1994). Formation of reactive oxygen spe-
cies can lead to membrane damage and eventually cell death.
Whiley et al. (1999) observed that A
max
, which is closely related to photo-
synthetic capacity, the quantum yield and F
v
/F
mPredawn
are substantially
lower in mango trees grown in containers (root-restricted) when compared
to field-grown trees. These observations were confirmed by Urban and
Alphonsout (2007) who studied the effect of the removal of a 1 cm-wide band
of bark on leaf photosynthesis and leaf N content of 3-year-old and 11-year-
old ‘Cogshall’ mango trees. Girdling is a common horticultural practice used
to manipulate tree growth and development in many fruit species. Its most
immediate effect is to stop the basipetal movement of assimilates through the
phloem, which results in an accumulation of carbohydrates above the girdle
(Roper and Williams, 1989; Schaper and Chacko, 1993; Di Vaio et al., 2001).
Girdling can promote floral induction in mango (Chacko, 1991), but it has
also been shown to reduce net photosynthesis. The major effect of girdling is
a dramatic increase in leaf carbohydrate concentration and a concomitant
decrease in photosynthetic electron transport and net photosynthesis (Fig. 6.6)
(Gonzalez and Blaikie, 2003; Urban and Alphonsout, 2007). Urban and
Alphonsout (2007) observed that A
net
was reduced by 77% within 28 days
from girdling and remained at about 2 mol CO
2
/m
2
/s until the beginning
of flowering. The decrease in photosynthetic electron transport rate (J) and
sustained photoprotection (reflected by the decrease in F
v
/F
mPredawn
) pro-
tected leaves of girdled branches effectively from photodamage, as shown by
the vigorous recovery of A
net
and J observed immediately after the appear-
ance of inflorescences. This increase in A
net
and J was associated with no
Q = 2000 µmol photons/m
2
/s
y = 155.4e
–0.0401x
R
2
= 0.74
0
50
100
150
200
250
0 10 20 30 40
Starch (g/m
2
)
J
(
µ
m
o
l

e
l
e
c
t
r
o
n
s
/
m
2
/
s
)
A
Q = 1200 µmol photons/m
2
/s
y = 120.3e
–0.0405x
R
2
= 0.74
Q = 400 µmol photons/m
2
/s
y = 84.5e
–0.0313x
R
2
= 0.69
Fig. 6.6. The relationship between the total photosynthetic electron flux (J) measured
at photosynthetic photon flux (Q) = 400 (), 1200 (●) and 2000 () mol photons/
m
2
/s, and the amount of starch per unit leaf area. Best fit lines at each Q were assessed
from measurements performed on both girdled and non-girdled mango leaves before
flowering. Data were used to establish the following relationship: J = (0.0434Q +
72.8)*e
–0.0412[starch]a
(Source: Urban and Alphonsout, 2007).
B. Schaffer et al. 186
decrease in leaf carbohydrate content during the first month following the
onset of flowering, suggesting that the effect of carbohydrate accumulation
on photosynthesis is mediated by sink activity. Apart from its negative effect
on the carbon budget of mango trees, girdling appeared to be rather harm-
less. However, leaf N concentration decreased, which indicates that there
may indeed exist long-term negative effects of girdling on photosynthetic
capacity. The width of bark (phloem) removed may be critical with respect to
the intensity of the effect of girdling on the tree. Whiley et al. (2006) girdled
the trunks of ‘B74’ (‘Calypso’™) mango trees in the Northern Territory of
Australia in autumn (as soon as they had come out of the wet season). The
girdles were no more than the thickness of a pruning saw (about 1 mm) and
healed within 6 weeks. In the first and third years after girdling, the trees had
significantly higher yields than non-girdled trees on which, coincidentally,
fruit matured early, thus giving market advantage. There was no significant
difference in yield in the second year of treatment between girdled and non-
girdled trees. The first and third years had strong natural induction while the
second year gave poor flowering across all varieties in the district. Thus, dur-
ing years of strong induction, this type of girdling most likely provided extra
carbohydrate reserves to drive flowering and support fruit set and retention
while in the off-flowering year there was sufficient carbohydrate reserves to
support reproductive activity. In contrast to observations with ‘Cogshall’
mango trees (Urban and Alphonsout, 2007), there was no evidence of long-term
effects of narrow girdles on leaf N of ‘B74’ mango trees (Whiley et al., 2006).
However, when wider girdles are made, tree recovery may take much longer
leading to sustained physiological disruption.
Proximity of inflorescences
While the effects of water stress and high light, temperature and atmospheric
CO
2
concentration on photosynthesis are increasingly well described, very
little is known about the effect of phenology, and especially of flowering on
photosynthesis of mango. There is some evidence that flowering may have
an effect on photosynthesis. Flowering-associated decreases in A
net
and g
s

were observed in sweet cherry (Roper et al., 1988) and mango (Shivashankara
and Mahai, 2000; Urban et al., 2004a). Lack of precise knowledge about the
effect of flowering on photosynthesis may impair our ability to adequately
simulate photosynthesis, especially for tropical fruit trees for which flower-
ing often extends over a long period of time. Mango flowering can last for >
2 months. Therefore, its effect on photosynthesis should not be overlooked.
Urban et al. (2004a) showed that the decrease in A
net
in mango leaves close to
inflorescences is not attributable to a g
s
-associated decrease in C
i
or to an
increase in R
d
. R
d
was lower in leaves close to inflorescences than in standard
leaves. If any, the effect of R
d
on A
net
was a positive one. This study suggested
strongly that the decrease in A
net
was due to a decrease in the electron flow in
photosystem II, but failed to provide direct evidence for it as well as the ele-
ments for interpretation. Using a modelling approach, Urban et al. (2008)
confirmed that there is a decrease in the total light-driven photosynthetic
electron flux in leaves close to inflorescences and showed that the decrease
Ecophysiology 187
in A
net
is also attributable to an increase in photorespiration. The latter
appears to be the consequence of a g
m
-associated decrease in C
c
, while the
former results from an increase in electron flow towards alternative sinks, a
decrease in the amount of leaf N per unit leaf area, and, hypothetically, either
a decrease in leaf N allocation to the bioenergetic pool of the photosynthetic
machinery, inorganic phosphorus depletion in leaves, or feedback inhibition
of photosynthesis. The latter hypothesis is least probable in the absence of
carbohydrate accumulation in leaves close to inflorescences. Both of these
hypotheses need to be tested to further our understanding of the inhibiting
effect of inflorescences on photosynthesis of nearby leaves. Interestingly, net
photosynthesis measured on leaves close to panicles bearing set fruits are
intermediary between those measured on standard leaves and those mea-
sured on leaves close to inflorescences, suggesting that changes in photosyn-
thesis associated with flowering are reversible. Urban et al. (2008) also showed
that processes other than temperature or light acclimation, and acclimation
to reduced sink activity, may cause leaf N concentration and photosynthetic
capacity to vary in mango. Whiley (unpublished data) observed that the
mango leaves immediately adjacent to inflorescences lose colour intensity as
the inflorescence grows out. Although leaf N over this period was not mea-
sured in mango, it has been measured in avocado (Whiley, 1994) which has a
similar intense burst of flowering in which a large biomass is produced in a
short time. Leaf N declines rapidly in avocado leaves as the inflorescences
break from buds and grow and then N stabilizes (at a lower concentration)
by mid-bloom. This can be reversed by N applications during flowering and
the additional application of a growth retardant (paclobutrazol (PBZ)) giving
leaf N and A a significant boost. Similarly to avocado, it is likely that the
reduction in A close to mango infloresences is related to reduced leaf N.
Photosynthetic contributions by fruit
Fruit of many species have chlorophyll and photosynthetic activity, particu-
larly during the early stages of growth (Jones, 1981; Whiley et al., 1991). How-
ever, for most crops, respiratory losses from fruit exceed photosynthetic gains
throughout ontogeny (Kriedemann, 1968; Whiley et al., 1991). An exception
to this is blueberry (Vaccinium spp.) fruit in which there is a net photosyn-
thetic gain from petal fall through to colour break, with an estimated 15% of
the total fruit carbon requirement contributed from fruit photosynthesis
(Birkhold et al., 1992). Studies with monoembryonic ‘Dashehari’ mangoes
showed that when fruit were approximately 10 mm in diameter, the photo-
synthetic rate of fruit was 2.7% that of leaves and declined to 1.2% of leaf
photosynthesis at fruit maturity (Chauhan and Pandey, 1984). However, even
this comparatively small carbon contribution may be important during the
critical fruit set period when trees rely on stored carbohydrates and a rela-
tively inefficient canopy to supply current photosynthates. Further studies
with mangoes to establish optimum light regimes for fruit photosynthesis at
different stages of ontogeny are warranted.
B. Schaffer et al. 188
6.3 Plant Water Relations
In this section, theoretical concepts of plant water relations are briefly out-
lined to help interpret the effects of environmental factors on mango water
relations (see also Nobel (1983) and Baker (1984)).
An important concept in plant water relations is water potential (Ψ),
which is a measure of the free energy of water. For pure water, Ψ = 0. As sol-
utes are added to water, its free energy decreases and becomes more nega-
tive. Water moves along a gradient from higher to lower (more negative) .
can be expressed as:
=

+
p
+
m
(6.11)
where is osmotic (or solute) potential which refers to the effect of solutes
on the change in free energy of water;
p
is the hydrostatic or pressure poten-
tial also referred to as the turgor pressure; and
m
is the matric potential,
which is generally negligible in plant cells.
In plant cells,
p
is generally positive or equal to 0. However, in xylem
tissue of transpiring plants,
p
is negative (under tension). The driving force
for transpiration is the vapour pressure difference between the leaf (consid-
ered to be water saturated) and the surrounding air. Thus, water moves from
a greater to a lower (or more negative)
p
and hence along a decreasing
gradient. The cohesive forces of the H
2
O molecules allows the xylem water to
remain in a continuous column even though there is a negative
p
.
Plant water stress can be determined from Eqn 6.11 and from changes in
. The components of , such as

and
p
, can often be used to define the
sources of water stress. The drought tolerance of mango highlights some
unique aspects of physiology of this tree with respect to its water manage-
ment. Typical mango environments in the tropics impose extreme water
stress and high evaporative demand for prolonged periods. Adaptive strate-
gies of mango trees include a deep root system (Sukonthasing et al., 1991),
desiccation-tolerant surface feeder roots and drought avoidance mechanisms
thought to be mediated by a comprehensive system of resin canals distrib-
uted throughout the tree (Venning, 1948; Joel, 1980; Joel and Fahn, 1980a, b;
Pongsomboon, 1991) and rapid stomatal closure. Plants with laticfers or resin
ducts/canals have been reported to be drought tolerant due to extended
maintenance of turgor following the withdrawal of water (Downton, 1981;
Kramer, 1983; Kallarackal et al., 1990). While the mechanism of turgor main-
tenance remains unresolved, it is believed that the latex or resin is probably
involved in the modulation of plant water status (Kallarackal et al., 1990).
The differentiation, structure and distribution of resin canals in mango has
been described by Venning (1948), Joel (1980) and Joel and Fahn (1980a, b, c).
Resin canals are present in trunks, shoots, leaves and fruit (exocarp) of mango
in close association with the vascular tissues. The resin contains mainly ter-
penes, but phenols and protein-carbohydrate mucilage are also present (Joel
and Fahn, 1980c). In well-watered trees, the resin is under positive pressure
and freely exudes from damaged or cut surfaces (Pongsomboon, 1991). In
studies of the development of water deficit in container-grown ‘Kensington’
Ecophysiology 189
mango trees, loss of turgor occurred in expanding leaves when leaf water
potential (
l
) reached –1.2 Mpa. In mature leaves turgor was not lost until
l

reached –1.75 MPa (Pongsomboon, 1991). Necrotic leaf areas appeared when

l
reached approximately –3.2 MPa with permanent wilting developing at
−3.45 MPa. This is high (less negative) compared with a
l
of –6.6, –5.0 MPa
for orange (Citrus sinensis) and macadamia (Macadamia integrifolia), respec-
tively (Fereres et al., 1979; Stephenson et al., 1989). Thus, mango leaves toler-
ate less internal water stress than woody perennial fruit trees from more
mesic environments. With mango, however, the permanent wilting point
was reached 36 days after withholding water compared to 10 days for simi-
larly sized macadamia trees (Stephenson et al., 1989). The higher critical
threshold of
l
and the longer period of survival for ‘Kensington’ mango
indicates that drought tolerance is based on more effective water regulation
to prevent desiccation and on maintenance of leaf turgor rather than resis-
tance by tissues to damage.
Reich and Borchert (1988) observed that stomatal regulation in mango
significantly reduced the rate of development of internal water deficit when
compared with four other tropical tree species. In water-withholding studies,
the radial expansion of mango trunks continued when most of the other
species were shrinking, indicating that mango trees could better tolerate
drought conditions and maintain photoassimilation rates. This is consistent
with the decrease in g
s
/A
net
observed by Urban et al. (2006) as a consequence
of drought (Fig. 6.1). A decrease in g
s
/A
net
indicates that there is an increase
in photosynthetic water use efficiency.
In containerized ‘Kensington’ mango trees, there was a linear correlation
between stomatal conductance and
l
during the development of water
stress (Pongsomboon, 1991). In contrast, with avocado and macadamia,
stomatal response was much more rapid with a curvilinear relationship
between
l
and stomatal conductance, and stomatal closure reached at –1.2
and 3.0 MPa, respectively. The slower response of stomatal conductance to

l
in

mango trees appears to be related to a more effective mechanism for
the mediation of water deficit development compared with avocado and
macadamia.
Pongsomboon (1991) monitored leaf water potential, osmotic potential of
resin (
r
) and osmotic potential of the whole leaf tissue (

) in container-
grown ‘Kensington’ mango trees when water was withheld for a 45-day
period. When tissues were fully hydrated,
l
and
r
were higher than

. For
40 days of the drying cycle,
l
and
r
declined at a similar rate; however,

declined to about –1.2 MPa within 4 days where it remained stable until 18
days into the drying cycle. There was a subsequent decline in

for 28 days
after withholding water when it stabilized at –2.0 MPa, remaining constant
for another 12 days. Pongsomboon (1991) suggested that osmotic adjustment
occurred, probably mediated through the resin as water deficit developed. It
appears, therefore, that the energy investment by the tree in a resin canal sys-
tem is justified by the vital drought-avoidance benefits conferred by main-
taining turgor and preventing wilting under prolonged periods of water
stress. Further investigations are required to substantiate these conclusions.
B. Schaffer et al. 190
In a recent experiment of 2-year-old potted ‘Cogshall’ mango trees
grafted on ‘Maison Rouge’ rootstock, midday minimum leaf water potential
remained relatively constant, about –1.0 MPa, for the range of predawn leaf
water potential (a surrogate of soil water potential) from 0 to –0.5 MPa, but
when the predawn leaf water potential dropped below –0.5 MPa, leaf water
potential fell rapidly to about –1.8 MPa (Gaelle Damour, Centre de coopéra-
tion Internationale en Recherche Agronomique pour le Développement
(CIRAD), personal communication). This ‘isohydric’ behaviour seemed to be
associated with rapid stomatal closure in response to decrease in water avail-
ability (Urban and Jannoyer, 2004).
Development of novel techniques to study xylem integrity have pro-
vided some unique insights into stomatal function (Tyree and Sperry, 1988;
Cochard et al., 2002). Xylem sap is transported under negative pressures in
plants and therefore is susceptible to cavitation events that render xylem
conduits non-conductive. Cavitation occurs when the negative sap pressure
exceeds a threshold value defined by anatomical characteristics (Tyree and
Sperry, 1988). Many species function very close to the point of embolism.
Therefore, stomata control both plant water losses and sap pressure and thus
may actively control the risk of xylem embolism (Jones and Sutherland,
1991). Xylem vulnerability to cavitation was studied in twigs of 13-year-old
‘Cogshall’ mango trees. Xylem vessels started to cavitate at a xylem water
potential of about of –1.5 MPa and underwent a rapid and substantial loss
(90%) of hydraulic conductivity when xylem water potential dropped from
–2.0 to –3.0 MPa (Fig. 6.7) (H. Cochard, unpublished data). This result is con-
sistent with the observation that leaves lose turgor when
l
reaches –1.75
MPa and permanent wilting appears when
l
is above –3.0 MPa (Pongsom-
boon, 1991) when Fig. 6.7 predicts 90% of xylem hydraulic conductivity is
lost. It also confirms that mango, like many other trees, operates close to the
point of embolism via effective stomatal control.
Direct measurement of xylem water potential using a pressure chamber
is problematic due to the presence of latex (Castro Neto et al., 2004). It is more
problematic in some cultivars, for example ‘Kensington’, than in others, such
as ‘Irwin’. Alternative indicators of plant water status such as measurements
of whole-tree water use (sap flow) and stem/branch shrinkage were found to
be sensitive and integrated indicators of whole-tree water status in mango
and were successfully used to schedule irrigation in a mango orchard in the
seasonal dry-wet tropical region of northern Australia (Lu, 2002, 2006).
6.4 Tree Growth and Development
Light
In an orchard, light distribution within and between tree canopies can have
a profound effect on growth and development of the fruit. We have previ-
ously discussed the effect of light on photosynthesis and defined the opti-
mum light levels required for mango leaves. When light levels fall below the
Ecophysiology 191
threshold required for light saturation of photosynthesis, the subsequent
reduction in available photoassimilates will affect growth of the tree. In many
tree fruit crops, flower-bud induction, fruit size and fruit colour are reduced
when low light levels occur due to crowding within and between tree cano-
pies (Jackson, 1980; Flore, 1994; Whiley and Schaffer, 1994). There is no pub-
lished information on the effect of light levels on mango fruit size, although
fruit are photosynthetically active and a reduction in size under low Q could
be expected.
Fruit skin colour is an important feature of mango with fruit of many
cultivars developing attractive pink to red coloration. Fruit colour is geneti-
cally determined and the reddish blush is generally more developed in
monoembryonic cultivars, while fruit from most polyembryonic cultivars
remain green/yellow at maturity. Skin coloration of mature fruit is partly
due to anthocyanins which develop when tissues are exposed to light. While
this subject is well researched in other fruit crops (Proctor and Creasey, 1971),
light levels required for skin coloration of mango fruit have not been quanti-
fied. Studies in Australia with the polyembryonic cultivar ‘Kensington’,
which develops a blush only on the exposed side of the fruit, indicated that
the position of fruit on trees had a significant effect on the development of
colour due to differences in the penetration of light into the canopy during
fruit ontogeny (Schaffer et al., 1994). The intensity of redness was greatest on
fruit from the eastern side of the tree followed by fruit from the south-western
100
80
60
40
P
L
C
20
0
–4 –3 –2
Xylem pressure (MPa)
–1 0
2006 light
2006 shade
2005
Fig. 6.7. Vulnerability of xylem of 13-year-old ‘Cogshall’ mango trees to cavitation
in twig segments from sun-exposed (light) or shaded sides of the tree. Cavitation is
expressed as percentage loss of conductivity (PLC) with decreasing xylem water
potential. Symbols represent means and error bars represent one standard error.
Vulnerability curves were obtained with the centrifuge technique (Source: Cochard
et al, 2005; and from H. Cochard, unpublished data, with permission).
B. Schaffer et al. 192
and northern sides of the tree. This information establishes an important con-
cept with respect to the light regime but does not quantify the absolute light
levels required for anthocyanin development. Further research is necessary
to establish physiological parameters from which pruning and orchard man-
agement strategies can be developed.
Temperature
Mango is a predominantly tropical species although the tree will usually
grow and produce more successfully in frost-free subtropical latitudes with
a marked dry season and high heat accumulation. Under optimum tempera-
tures with non-limiting nutrients and water, the tree remains vegetative with
growth flushes occurring at regular intervals. The large size and poor crop-
ping of trees in the humid lowland tropics are well known, and there is a
direct relationship between temperature and the frequency of vegetative
flushes. Trees grown at 20°C days/15°C nights (20/15°C) required 20 weeks
(mean of ten cultivars) to complete a growth/dormancy cycle while at
30/25°C the same cycle was completed in 6 weeks (Whiley et al., 1989). There
are marked differences between cultivars with respect to their tendency
towards vegetative growth. For instance, in controlled temperature studies
over a 20-week period, at 30/25°C, ‘Irwin’ produced 2.0 growth flushes with
approximately 45 days of dormancy between active growth periods while
‘Kensington’ produced 4.7 growth flushes with only 5 days of quiescence
between flushes (Whiley et al., 1989). Dry matter accumulation over the 20
weeks was similar for the two cultivars; however, starch accumulation in the
woody trunk tissues of ‘Irwin’ and ‘Kensington’ was 13 and 3.6% of dry mat-
ter, respectively. The response differences between these two cultivars may
be the contributing factor to their performance at tropical latitudes where
temperature is non-limiting for growth. Under these conditions ‘Irwin’ has
more reliable cropping than ‘Kensington’, suggesting that the genetically
determined low-vigour trait is more sensitive to environmentally precipi-
tated stresses that induce flowering.
The number and size of leaves which develop on each growth flush are
also influenced by temperature. Whiley et al. (1989) reported that on trees
growing at 20/15°C an average of 7.1 leaves per flush were produced while
at 30/25°C there were 13.6 leaves on each growth flush (data are mean values
from ten cultivars). At 30/25°C the mean leaf size was 300% greater than
those on trees growing at 20/15°C. Soil temperatures have also been reported
to have a strong effect on the growth of mangoes. In studies with ‘Irwin’
grafted on ‘Turpentine’ rootstocks, episodic shoot growth occurred when
soil temperatures were held at 27°C or 32°C for 120 days but an extended
dormant period developed when soil temperatures were held at 21°C (Yusof
et al., 1969). These results indicate that environmental control over shoot
growth of mangoes may in part be related to soil temperatures.
From controlled temperature studies it has been calculated that the median
daily temperature (mean of the maximum and minimum daily temperatures)
Ecophysiology 193
at which shoot growth ceases is approximately 15°C (mean value for ten cul-
tivars) (Whiley et al., 1989). Subsequent studies (Issarakraisila et al., 1991)
have confirmed that 15°C is the critical minimum growth temperature for
shoots of ‘Kensington’.
Stress-inducing temperatures which prevent shoot growth have been
shown to promote floral induction in mangoes, but this is outside the scope
of this discussion. For further information of the effects of temperature on
pollination, floral initiation and fruit development, see Davenport, Chapter
5, this volume and Schaffer et al. (1994). We again emphasize that although
mango is a ‘heat-loving’ crop well adapted to the hot, semi-arid subtropics
and monsoonal tropics, in these environments it experiences extremes of heat,
drought and evaporative demand that may cumulatively reduce potential
production capacity.
Drought
Although mango is considered to be drought tolerant and may survive with-
out rain or irrigation for > 8 months (Gandhi, 1955), water deficits during the
reproductive cycle can have severe effects on the retention and early growth
of mango fruit. In studies with bearing, container-grown ‘Irwin’ trees, pre-
dawn
l
levels were maintained at either less than –0.3 MPa (non-stressed)
or –1.2 MPa (water stressed) for the first 2 months after fruit set. For the first
5 days following fruit set, all trees lost a similar percentage of fruit, but there-
after fruit abscission was greater on water-stressed trees. After 1 month,
drought-stressed trees had retained approximately 4% of their initial fruit set
compared with approximately 8% on non-stressed trees. During the first 30
days following fruit set, the rate of fruit growth for non-stressed trees was
twice that of drought-stressed trees, and final fruit size (measured 60 days
after fruit set) of non-stressed trees was 20% greater than on water-stressed
trees. In a separate study, another group of ‘Irwin’ trees was maintained
stress-free (pre-dawn water potential above –0.3 MPa) during the first month
following fruit set with water-stress (–1.2 MPa) imposed on some trees dur-
ing the second month of fruit development (Pongsomboon, 1991). There was
no effect of drought on fruit retention but final fruit size was 34% smaller for
stressed compared with non-stressed trees.
In field studies, Singh and Arora (1965) compared fruit drop of monoem-
bryonic ‘Dashehari’ trees irrigated at 1-week intervals with trees irrigated at
3-week intervals. During the first 6 weeks of fruit growth, weekly irrigation
reduced fruit drop compared with the irrigation at 3-week intervals. During
the latter stages of fruit development, these gains were lost as more fruit
dropped from the weekly irrigated trees. In another study, field-grown
monoembryonic ‘Tommy Atkins’ trees were managed under different irriga-
tion regimes from early fruit set until the start of the rainy season (approxi-
mately 43 days) (Larson et al., 1989). Trees were irrigated on a 7- or 14-day
schedule or received no irrigation. Pre-dawn
l
was –0.3 MPa for trees irri-
gated on the 7-day schedule and decreased to –0.5 MPa for the non-irrigated
B. Schaffer et al. 194
trees. Irrigation at 7-day intervals resulted in the greatest yield with the larg-
est fruit, especially during the early harvest period (Larson et al., 1989).
Irrigation, therefore, particularly during the first 4–6 weeks following
fruit set, increases individual fruit size and yield. This is a critical period of
fruit development since it is when cell division is most rapid and cell walls
are developed. Even slight reductions in plant water status during this period
can have adverse effects on fruit growth and retention (Pongsomboon, 1991).
Although drought tolerance of the mango tree is well known, this comes at
considerable cost to tree performance, particularly in areas with prolonged
dry seasons that extend through flowering and fruiting. Irrigation is there-
fore one of the most powerful tools to alleviate non-lethal yet potentially
yield-reducing drought stress.
Flooding
Studies with container-grown mango trees have reported variable responses
with respect to tree survival. Larson et al. (1991c) observed that as many as
45% of trees died after their roots were submerged in water for 4–10 days, but
in the surviving population no further mortality occurred when flooding
was extended for up to 110 days. In other experiments, there was no tree
mortality after container-grown mango trees were flooded from 1 to several
months although tree growth was significantly reduced (Larson et al., 1991c;
B. Schaffer unpublished data, Homestead, Florida, 1993).
The ability of mango trees to survive prolonged flooding appears to be
dependent on the development of hypertrophic (swollen) stem lenticels
immediately above the water line (Plate 42). The initial stages of lenticel
hypertrophy are characterized by the development of intercellular spaces in
the phellem tissue and production of additional phellem tissue by increased
phellogen activity. Later stages of hypertrophy are characterized by the
development of intercellular spaces in the phellem tissue and cortex (Larson
et al., 1991a). Observations vary among studies whether or not trees devel-
oped hypertrophic lenticels or how quickly after flooding they formed. These
anomalies have been attributed to environmental differences at the time of
root submersion (Larson et al., 1991c). In trees that died as a result of flooding
stress there was no lenticel hypertrophy; however, stem lenticels hypertro-
phied within 4–10 days on mango trees that survived flooding (Larson et al.,
1991a, c, 1993). Sealing hypertrophic lenticels of mango trees with silicone
grease or petroleum jelly resulted in tree death within 3 days of flooding,
thereby demonstrating their necessity for tree survival. The role of hypertro-
phic lenticels in flood-tolerant species is not clear, although they are thought
to eliminate potentially toxic metabolites such as ethanol, acetaldehyde and
ethylene which result from anaerobic respiration in the roots (Chirkova and
Gutman, 1972; Larson et al., 1993). They may also confer flood tolerance by
enhancing O
2
diffusion to the roots (Kozlowski, 1984).
In some instances, adventitious roots have developed above the water
line when container-grown mango trees have been flooded for long periods
Ecophysiology 195
(Schaffer et al., 1994). It is likely that these roots facilitate the absorption and
translocation of O
2
to submerged roots and their development is a common
morphological response of many woody plants to root anoxia. The develop-
ment of adventitious roots has not been reported for flooded, field-grown
trees and they may only form on young trunks after extended flooding peri-
ods, which usually do not occur under normal production conditions.
Vegetative growth of mango trees generally declines when trees become
flooded for > 2–3 days. When trees in a limestone soil in containers were
flooded for > 110 days, there was a 94% reduction in shoot extension growth,
while flooding for approximately 10 days resulted in a 57% reduction in
shoot extension growth (Larson et al., 1991c). In a subsequent study, the stem
radial growth (a more sensitive indicator of tree growth than shoot extension
growth) of mango trees decreased 2 weeks after roots were submerged.
Flooding for > 14 days also significantly reduced root dry weight, resulting
in an increased shoot to root ratio (Larson et al., 1991c). These adverse effects
of flooding on the growth of mango trees are expected as reduced net photo-
synthesis and presumably higher root respiration limit the availability of
carbon-based assimilates required for growth.
Wind
Most fruit trees benefit from wind protection, particularly during the estab-
lishment years when the disruption of physiological processes results in a
significant depression of growth in young trees. In addition, wind also causes
abrasions to the skin of fruits, particularly when they are small, which
develop into unsightly blemishes by the time they are fully grown thereby
reducing quality and market value. However, the cost of windbreaks may
not be offset by higher returns. In some mango-producing regions, winds are
not sufficiently strong to justify the cost of wind protection. Until recently,
wind protection in South Africa was not recommended for mangoes due to
the loss of potential cropping space by ‘living’ windbreaks, their potential to
create frost pockets, and the likelihood of promoting the incidence of flower
and fruit diseases through increased humidity (Van der Meulen et al., 1971);
however, the value of windbreaks is well appreciated today in South Africa
(B.N Wolstenholme, personal communication, Pietermaritzburg, South
Africa, 1995).
In studies with ‘Kensington’ mangoes in Australia, where artificial wind-
breaks were constructed to shelter trees from the prevailing summer south-
easterly winds, a 600% increase in yield was recorded in the first year
following wind protection (Mayers et al., 1984). This significant improvement
in tree performance was a result of better growth of trees which set and held
more fruit per panicle, suffered less damage to leaves (cuticle fracturing) and
had reduced fruit loss from bacterial black spot (caused by Xanthomonas camp-
estris pv. mangiferaeindicae) compared to the wind-exposed trees. These results
indicate that wind can have a significant effect on mango productivity from
the reduction of both growth and yield through undisclosed physiological
B. Schaffer et al. 196
mechanisms, and a decreased level of bacterial black spot infection. The pro-
vision of windbreaks in orchards is expensive with decisions to be made on
the use of either ‘living’ or artificial shelters. Requirements for wind protec-
tion will vary depending on site circumstances, and all factors pertaining to
crop performance will require careful consideration.
Salinity
Salt stress in mango trees produces symptoms similar to those described for
other species (Harding et al., 1958; Ehlig, 1960; Kadman, 1964; Bingham et al.,
1968). Mild symptoms of chloride toxicity are scorched leaf tips and margins
and leaf curling, while in more severe cases growth ceases, leaves abscise and
trees die. Necrotic areas develop on leaves of trees exposed to high sodium
levels (Jindal et al., 1976; Kadman et al., 1976; Gazit and Kadman, 1980). Irri-
gation water concentrations of 20–60 mM sodium chloride (NaCl) or sodium
sulfate (Na
2
SO
4
) reduced leaf area and changed the branching structure of
container-grown mango trees, suggesting that salinity resulted in reduced
leaf cell elongation, and affected the activity of the terminal meristem
(Schmutz and Lüdders, 1993). As the duration of the exposure to saline con-
ditions increased, transpiration decreased exponentially (Schmutz and Lüd-
ders, 1993). In a later study, Schmutz (2000) found that following a gradual
increase in salinity of the nutrient solution applied to potted polyembryonic
‘13-1’ rootstocks (from 0 to 120 mM NaCl over 15 days), A
max
significantly
declined despite there being no visible leaf symptoms of salt toxicity. The
decline in A
max
occurred within 6 days of beginning the salinity treatment,
which was 15 mM NaCl for 3 days followed by 30 mM NaCl for 3 days. This
indicates that photoassimilation in mango is quite sensitive to exposure of
trees to salinity.
There is considerable variation in salinity stress of mango, both within
and between populations of mono- or polyembryonic mango ecotypes. Based
on the results of limited studies, there appears to be greater salt tolerance in
polyembryonic than in monoembryonic populations (Jindal et al., 1975; Kad-
man et al., 1976). In seedling populations from mono- and polyembryonic
cultivars irrigated for 2 years with water containing approximately 10 mM
chloride, most plants developed leaf scorching after 6 months which gradu-
ally became more severe, culminating in degeneration and death. However,
some seedlings which had no damage or only slight toxicity symptoms were
mostly of the polyembryonic ‘13-1’ rootstock cutivar or related types (Kad-
man et al., 1976). Leaf analyses revealed that the chloride concentration in
tolerant seedlings (0.68–0.77%) was greater than in susceptible seedlings
(0.43–0.55%). In addition, tolerant plants had lower leaf concentrations of
potassium, calcium and magnesium than saline-sensitive seedlings, possibly
a result of comparative nutrient dilution since vegetative growth was greater
for saline-tolerant than for saline-sensitive seedlings. Kadman et al. (1976)
also suggested that the mechanism of chloride tolerance in ‘13-1’ was based
on greater physiological tolerance of chloride concentrations in leaf tissues,
Ecophysiology 197
rather than ion exclusion or a selective uptake mechanism common in other
species (Collander, 1941; Walker, 1986). However, the relative sodium toler-
ance of ‘13-1’ was due to exclusion of sodium from shoots and its accumula-
tion in root cell vacuoles (Schmutz and Lüdder, 1993). More recently Hoult
et al. (1996) reported significant cultivar differences within a population of 21
polyembryonic mango cultivars exposed to saline (480 mg/l NaCl) irrigation
water for 10 months. Differences were measured in leaf Na (0.37–1.34%) and
Cl (0.39–1.07%) concentrations but these were poorly correlated with toxicity
symptoms on leaves.
Salinization of agricultural land is increasing and in many areas salinity
management is critical. There appears to be sufficient genetic diversity within
Mangifera indica to enable the selection and development of saline-tolerant
rootstocks (Kadman et al., 1976; Gazit and Kadman, 1980; Hoult et al., 1996).
However, quantitative data on the critical limits of soil and water salinity
which mango trees will tolerate without reductions in yield and fruit quality
are needed.
Elevated atmospheric CO
2
concentration
Growing ‘Kensington’ mango trees for 6 months in a controlled atmosphere
glasshouse with an ambient CO
2
concentration of 600 mol/mol resulted in
more dry matter partitioned to the roots compared to plants grown in an
ambient CO
2
environment of 350 mol/mol (Schaffer et al., 1999) (Fig. 6.8).
Fruit dry weight was greater for mango trees grown in an atmospheric CO
2

concentration of 600 mol/mol compared to trees grown at 350 mol/mol
(Schaffer et al., 1999) (Fig. 6.9). Most of the increased total fruit dry weight at
Old
leaves
Old
branches
New
leaves
Plant part
New
branches
Trunk Roots
0
100
200
300
400
500
D
r
y

w
e
i
g
h
t

(
g
)
600 µmol CO
2
/mol
350 µmol CO
2
/mol
Fig. 6.8. Partitioning of dry matter in ‘Kensington’ mango trees grown for 6 months in
atmospheric CO
2
concentrations of 350 or 600 mol/mol. Bars represent means (n = 6
trees) ± standard error (Source: redrawn from Schaffer et al., 1999).
B. Schaffer et al. 198
the higher atmospheric CO
2
concentration was a result of increased amount
of pulp; whereas, there were no significant effects of increased atmospheric
CO
2
concentration on dry matter accumulation in the skin, testa or seed
(Schaffer et al., 1999) (Fig. 6.9). Thus, increasing the atmospheric CO
2
assimi-
lation rate increased growth and partitioning to the mesocarp. Therefore,
increased atmospheric CO
2
concentration (at least to 600 mol/mol) as a
result of global climate change may increase the economic yield of mango
(Schaffer et al., 1997). However, under actual environmental conditions result-
ing from global climate change, water and nutrient availability may be limit-
ing and is likely to offset any increases in biomass or economic crop yield
resulting from elevated ambient CO
2
concentrations (Gitay et al., 2001).
6.5 Crop Production
Temperature limitations to crop production
Temperature is probably the most important environmental variable to con-
sider when selecting mango cultivars for particular sites. The mean tempera-
ture range for optimum growth of mango is about 24–30°C (Mukherjee, 1953;
Whiley et al., 1989). However, mango trees can tolerate temperatures up to
48°C for short periods (Mukherjee, 1953). Mango trees have limited tolerance
to cold and trees are usually severely damaged or killed after a few hours at
temperatures < 0°C (Carmichael, 1958; Campbell et al., 1977). Although
mature trees have withstood temperatures of −4°C for a few hours with lim-
ited damage, juvenile trees were killed after 13 h at −4°C to −6°C (Campbell et al.,
1977).
Fruit tissues
Skin Flesh Testa Seed Total fruit
D
r
y

w
e
i
g
h
t

(
g
)
0
10
20
30
40
50
600 µmol CO
2
/mol
350 µmol CO
2
/mol
Fig. 6.9. Partitioning of dry matter in ‘Kensington’ fruit of trees grown for 6 months
in atmospheric CO
2
concentrations of 350 or 500 mol/mol. Bars represent means
(n = 33–49 fruit) ± standard error (Source: redrawn from Schaffer et al., 1999).
Ecophysiology 199
Monoembryonic mango cultivars tend to be more cold tolerant than
polyembryonic cultivars, probably due their probable subtropical origin.
There are also differences in fruit setting capacity between mono- and poly-
embryonic cultivars at subtropical latitudes, where monoembryonic culti-
vars crop more successfully when minimum temperatures fall below 12°C
during flowering (Searle et al., 1995). Differences in low temperature toler-
ance have important implications for the selection of suitable cultivars for
production under specific climatic conditions.
Light interception and orchard design
Light interception and utilization within tree canopies is a primary consider-
ation in orchard design. Thus, tree spacing as well as pruning practices in
orchards are primarily based on maximizing light for photosynthesis (see
previous section on ‘Light’ under Photosynthesis, this chapter). The follow-
ing equation describes the relationship between light interception, photosyn-
thesis and yield in fruit tree orchards:
Biological Yield = (Light Available) (% Light Intercepted) (Photosynthe-
sis) − Respiration (6.12)
where biological yield refers to dry matter production, including that of fruit
(Lakso, 1994). The amount of light available is a function of climate and can-
not be manipulated, and potential net photosynthetic efficiency of a crop is
inherent and cannot be altered without genetic manipulation. Thus, optimiz-
ing biological yield is based on maximizing the percentage of light inter-
cepted by the orchard canopy and minimizing stress so that photosynthetic
potential is not compromised. With respect to light, the mango tree presents
special problems due to long-lived leaves, dense canopies, and the potential
for vigorous growth in some cultivars.
Maximizing light interception by the photosynthetic surface of an orchard
is a function of tree spacing, canopy density and height. For some fruit tree
trees, for example apple (Malus domestica) and citrus (Citrus spp.), the use of
low vigour rootstocks and pruning provides opportunities for an array of
management options. However, the lack of dwarfing rootstocks and the com-
plications in pruning due to the floral morphology of mango (inflorescences
are predominantly borne terminally on the most recently produced shoots)
limit the efficient harvest of light with respect to maintenance of productiv-
ity. Improved productivity can be obtained by grafting, which either short-
ens the juvenility phase and/or exerts some control over vigour. Compared
to temperate fruit orchards, canopies of commercial mango orchards have a
higher proportion of ‘shade’ to ‘sun’ leaves (Schaffer et al., 1994). Mango trees
are rarely selectively pruned (Young and Sauls, 1981), although novel ideas
of ‘heading back cuts’ after harvest have been investigated (Oosthuyse, 1992).
In some areas, trees are mechanically topped and hedged to control tree size.
Hedging encourages increased complexity (ramification), resulting in a dense
outer canopy. Where costs are not prohibitive, strategically timed, selective
B. Schaffer et al. 200
pruning to increase the percentage of leaves exposed to > 60% of full sunlight
will increase the photosynthetic efficiency of the canopy with a potential
improvement in yield. Schaffer and Gaye (1989) increased light interception
of mango by removing 25% of the canopy. This resulted in higher chlorophyll
concentrations in leaves of pruned canopies later in the year. Although net
photosynthesis was not measured in that study, it is likely that the higher leaf
chlorophyll concentrations in the pruned canopies resulted in higher photo-
synthetic rates.
Light utilization of mango can be enhanced by pruning, but the timing of
such treatments is critical. For example, in the subtropics, shoots produced
following harvest generally flower 3–5 months after being exposed to induc-
tive (cool) temperatures. Therefore, trees should be pruned immediately after
harvest to improve light penetration and contain tree size. However, in the
tropics there is a shorter period between the cessation of summer growth and
flowering and summer-grown shoots of many cultivars fail to induct that
year (Scholefield et al., 1986; see Davenport, Chapter 5, this volume). There-
fore, due to the removal of potential flowering sites, summer pruning of
mango trees in the tropics generally reduces yield in the following season.
Growth control of mango trees through the development of dwarfing
rootstocks, low-vigour cultivars and pruning strategies to increase light pen-
etration within tree canopies and by entire orchards, may improve yield per-
formance of this crop. Improved information on light interception, critical
leaf to fruit ratios and relationships between shoot maturity and floral induc-
tion with respect to genotype/environmental interactions will enhance the
development of improved cultural practices for mango production. It is per-
tinent to emphasize that few evergreen fruit trees are as precocious as mango,
or as large at maturity. Orchardists should take advantage of the precocity
and of light interception principles by initial high-density planting, with
hedgerows perhaps the best option. As trees become crowded, however, effi-
ciency of light interception is compromised and remedial action before this
occurs is required to maintain productivity.
6.6 Conclusions
Although much literature in the past stressed problems associated with low
temperature on mango growth and production, sustained high temperatures
with associated soil moisture stress during fruit ontogeny and high evapora-
tive demand are perhaps the major reasons for relatively low yields in mango
orchards worldwide, especially in the tropics. Although mango trees have a
number of survival mechanisms that allow them to cope with stressful envi-
ronments, these come at a considerable energy cost thereby potentially reduc-
ing the availability of carbon-based inputs for fruiting. It is likely that annual
assimilation gains and resource availability during critical developmental
periods are inadequate for sustained high yields of quality fruit. These prob-
lems can be alleviated by development of improved germplasm with adapta-
tion to specific environments (Whiley et al., 2006). In the future, greater effort
Ecophysiology 201
is required for rootstock development and understanding the manipulative
effect on whole tree physiology. Expansion of human populations and cli-
mate change scenarios predict lesser and poorer quality water available for
cropping systems across tropical and subtropical latitudes. Hence, greater
salinity tolerance within the species should be identified. Knowledge of
mango physiology, particularly in relation to the tree’s responses to varying
environmental conditions remains basic and must be expanded. A greater
understanding of these principles together with their application will assist
in the development of more productive cropping systems.
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© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
210 (ed. R.E. Litz)
7 Fruit Diseases
D. Prusky, I. Kobiler, I. Miyara and N. Alkan
Agricultural Research Organization, Bet Dagan, Israel
7.1 Introduction 210
7.2 Anthracnose 211
The pathogenesis strategy 212
Epidemiology – the disease cycle 213
Mechanisms of fungal pathogenicity and fruit resistance 214
Management 216
7.3 Alternaria Rot (Black Spot) 219
Pathogenesis 219
Mechanisms of fungal pathogenicity and fruit resistance 220
Management 220
7.4 Stem-end Rots 221
Pathogenesis 221
Epidemiology 222
Management 223
7.5 Other Minor Diseases 224
7.6 Conclusions 224
7.1 Introduction
Postharvest diseases can reduce fruit quality and cause severe economic
losses, due to decay resulting in completely unmarketable and blemished
fruits that are often sold in less demanding local markets, where the prices
are considerably lower than export prices. It is clear to the producer that
quality at the time of harvest cannot be improved but merely maintained for
a limited period of time. Harvesting fruits at the optimal stage, with respect
to size and maturity, can, therefore, ensure peak quality and maximum shelf-
life potential. Thus, managing total tree health can contribute to reducing
postharvest losses. It is known that older and neglected orchards may become
a profuse inoculum source for postharvest diseases. Furthermore, preharvest
Fruit Diseases 211
stress factors such as excess or shortage of water, fluctuating or extreme envi-
ronmental conditions and high nitrogen levels (Hawthorne, 1989) can render
fruit more susceptible to postharvest diseases. After harvest, improper treat-
ment of fruits through storage at non-optimal temperatures accelerates fruit
deterioration as a result of enhancement of normal physiological processes
such as respiration and ethylene production (Thompson et al., 2002). More-
over, excessive temperatures in the field during harvesting, in transit and in
the packing house can render tropical fruit more susceptible to chilling injury
and can contribute to the development of disease. Combinations of high tem-
peratures and high relative humidity (RH) favour the growth of postharvest
pathogens, and can contribute to the development of disease at the retail end
(Banik et al., 1998). Proper, uninterrupted cooling, therefore, protects quality
and extends the shelf life of produce. Precooling of products (Thompson et
al., 2002), and application of top or liquid icing, vacuum, hydrocooling and
forced air cooling (Thompson et al., 2002) are examples of effective alterna-
tive methods that can be used to remove field heat and to restrict pathogen
growth. Effective cold-chain management is crucial to ensuring product
integrity and preventing postharvest pathogens from spoiling produce in
transit or shortly after arrival (Lizada, 1993). Furthermore, low-temperature
storage conditions are generally not conducive to disease development, a
fact that can be exploited to ensure quality and extend shelf life (Banik et al.,
1998). However, improper cooling during shipping, or interrupted cooling
will also promote microbial growth, resulting ultimately in product spoilage.
In practice, several breaks in the cold chain might have significant impacts on
decay development.
Among the postharvest diseases of mango, anthracnose is the most prev-
alent in humid growing regions. The incidence of this disease can reach
almost 100% in fruit produced under wet or very humid conditions. Other
common postharvest diseases of mango in humid areas are stem-end rot,
caused by Lasiodiplodia theobromae (Pat.) Griffon & Maubl. (Sangchote, 1998)
and Dothiorella dominicana Petrak. et Cif. (Johnson and Sangchote, 1994), and
black mould rot, caused by Aspergillus niger van Tieghem. Mango black spot,
caused by Alternaria alternata (Fr.:Fr.) Keissl. (Prusky et al., 1983), is prevalent
in dry countries.
7.2 Anthracnose
Mango anthracnose is caused by Glomerella cingulata (Stoneman) Spauld. &
H. Schrenk (anamorph: Colletotrichum gloeosporioides (Penz.) Penz. & Sacc. In
Penz.), C. gloeosporioides Penz. var. minor J.H. Simmonds (Fitzell and Peak,
1984) and Colletotrichum acutatum J.H. Simmonds (Freeman et al., 1998). The
pathogen also causes blossom blight, leaf blight and, in severe cases, tree
dieback (Ploetz, 1994; Ploetz and Prakash, 1997). The form-genus Colletotri-
chum Corda (form-order Melanconiales; form-class Coelomycetes; subdivi-
sion Deuteromycotina) comprises imperfect fungal species which exist as
Glomerella (subdivision Ascomycotina) in their sexual, teleomorphic or perfect
D. Prusky et al. 212
state. These fungal pathogens occur worldwide, and the genus is synony-
mous with anthracnose. Leaf anthracnose appears as irregular black necrotic
spots on both sides of the mango leaves. Lesions often coalesce and form
large necrotic areas, frequently along the leaf margins. Lesions develop pri-
marily on young tissue, and conidia are formed in lesions of all ages. Under
favourable conditions, the fungus can invade the twigs and cause dieback
(Ploetz et al., 1996). Panicle anthracnose or blossom blight can affect both the
inflorescence stalk and the individual flowers; in the stalk, elongated dark
grey to black lesions appear; the blighted flowers are dry, and their colour
ranges from brown to black. Fruits smaller than pea-size can be infected and
abort; whereas, larger fruits that are aborted because of normal self-thinning
or other physiological causes are usually mummified. The resulting mummi-
fied fruit are invaded saprophytically by C. gloeosporioides, and the fungus
sporulates abundantly on them.
Although field anthracnose causes considerable damage, the vast losses
inflicted by postharvest anthracnose are of far greater economic importance.
Postharvest anthracnose appears on the fruit surface, as rounded brown to
black lesions with an indefinite border. Lesions > 2 cm in diameter are fairly
common. Lesions of various sizes can coalesce and cover extensive areas of
the fruit, typically in a tear-drop pattern that develops from the basal towards
the distal end of the fruit. Lesions are usually restricted to the peel, but in
severe cases the fungus can invade the pulp. In advanced stages of the dis-
ease, the fungus produces acervuli, and abundant orange to salmon-pink
masses of conidia appear on the lesions.
The pathogenesis strategy
Colletotrichum gloeosporioides is regarded as a hemibiotrophic species that
commences its invasion with a transient post-penetrative asymptomatic bio-
trophy, characterized by a temporary confinement with a localized mode of
intracellular hemibiotrophy. This is succeeded by a phase of destructive
necrotrophy that culminates in the appearance of disease symptoms and pro-
duction of conidiomata by the pathogens (Prusky and Plumbley, 1992; Latunde-
Dada et al., 1999). Quiescent species pass through a prolonged phase of
pre-penetrative growth that is arrested in synchrony with the physiological
state of the infected organ.
Following its landing on the fruit or host tissue, the ungerminated
aseptate conidium differentiates to a melanized appressorium. In the case of
Colletotrichum lindemuthianum, appressorial adhesion is mediated by the man-
nose- and galactose-rich glycoproteins of the extracellular matrices, which
are secreted during appressorial differentiation (Pain et al., 1996). Both the
surface wax of the specific fruit and the hard surface of the tissue are recog-
nized as host signals that selectively trigger germination and appresso-
rium formation solely by the conidia of C. gloeosporioides (Podila et al., 1993;
Liu and Kolattukudy, 1998). The melanin of the melanized appressoria pro-
tects the fungus and behaves as a permeability barrier, and the appressoria
Fruit Diseases 213
contain osmolytes (i.e. glycerol) that are needed for generating the osmoti-
cum from which high turgor pressure will drive the invasive forces of the
penetrating hyphae through the small appressorial pores (0.5 m in Col-
letotrichum sublineolum). The osmolyte is generated by the metabolism of
stored glycogen, trehalose and lipids and, at an in vivo concentration of at
least 3.3 M in mature-melanized appressoria (de Jong et al., 1997), this osmo-
lyte is capable of generating turgor pressures as high as 8 MPa (Howard et al.,
1991; Money and Howard, 1996). Melanized appressoria appear to be quite
capable of a forcible, non-enzymatic penetration of an intact host surface.
However, Dickman et al. (1982) suggested that attack on papaya by C. gloeo-
sporioides depended on cutinase production by the pathogen. The germi-
nated appressoria in Colletotrichum develop single infection hyphae that
grow and extend into the waxy cuticle, reach the first layers of pericarp
cells, and then become quiescent for long periods of time (Coates et al.,
1993; Prusky, 1996).
In recent years, the taxonomy of Colletotrichum has been clarified by the
adoption of molecular biological techniques that involve the use of poly-
merase chain reaction amplification, alignment of nucleotide sequences, and
the construction of dendograms, phylogenetic trees and similarity matrices
from the data generated. For example, Sherriff et al. (1994) and Sreenivasap-
rasad et al. (1996) used homologies between the nucleotide sequences from
amplified non-transcribed regions (internally transcribed spacers 1 and 2
(ITS1, ITS2)) and the large subunits (domains 1 and 2) of ribosomal DNA
extracted from a wide range of isolates to both justify and resolve the taxo-
nomic status of Colletotrichum species. Bailey (1997) proposed the adoption of
species aggregates based on these and other data.
Epidemiology – the disease cycle
Conidia are formed abundantly in the mango canopy (Fitzell and Peak, 1984)
which, therefore, is considered to be the primary source of inoculum. In the
field, C. gloeosporioides produces conidia on lesions on leaves, twigs, panicles
and mummified fruits (Arauz, 2000). Conidia can be rain-splashed onto other
leaves or flowers, where they can cause secondary infections. Developing
fruit can be infected, and some isolates can cause preharvest fruit loss (Gan-
totti and Davis, 1993). In the case of postharvest anthracnose, developing
fruits are infected in the field, but the infections remain quiescent until the
onset of ripening, which occurs after harvest. Once the climacteric period of
the fruit starts, lesions begin to develop but there is no fruit-to-fruit infection.
In this context Prusky and co-workers (Guetsky et al., 2005) suggested that
the capability of C. gloeosporioides to cause early disease symptoms in unripe
fruits depends on the activation of laccases by specific strains of the fungus.
Details of these systems are discussed in the following section.
Colletotrichum gloeosporioides requires free water or RH > 95% to enable
conidial germination and appressorium formation (Fitzell et al., 1984; Dodd
et al., 1991). However, conidia can survive for 1–2 weeks under low RH and
D. Prusky et al. 214
then germinate if exposed to 100% RH (Estrada et al., 1993). In general, infec-
tion is favoured at temperatures ranging from 20 to 30°C.
Mechanisms of fungal pathogenicity and fruit resistance
Unripe mango fruits are reservoirs of extremely high concentrations of pre-
formed antimicrobial compounds. This arsenal of constitutive resistance
weapons may accumulate in the immature pericarp, but not in the mesocarp,
at concentrations in fruit fresh weight of up to 220 g/g. They include mix-
tures of 5-substituted resorcinols such as resorcinol-5-(12-heptadecadienyl)
and resorcinol-5-(pentadecyl) (Droby et al., 1986, 1987; Prusky and Plumbley,
1992; Prusky, 1996) (Fig. 7.1). Faced by such a chemically adverse environ-
ment, the fungus usually postpones its development until the concentrations
of the compounds in the host decline. The capability to penetrate the host
without encountering the arsenal of passive chemical resistance or triggering
the weapons of active resistance is inherent in a number of fruit-infecting
Colletotrichum species such as C. gloeosporioides, C. acutatum and A. alternata
(Prusky and Keen, 1993). The conidia, melanized appressoria and penetra-
tion pegs of these quiescent species have evolved mechanisms of physiologi-
cal inactivity that enable them to pause until the host-ripening process and
the decline of antifungal compounds starts, thus enabling the avoidance of
host defences that exist within unripe, pre-climacteric fruits. Colletotrichum
gloeosporioides is a field-to-store pathogen whose conidia infect during the
preharvest growth of fruits. In fruits such as mango and avocado, termina-
tion of fungal quiescence is associated jointly with the climacteric production
of ethylene during fruit ripening and the onset of dramatic reductions in the
levels of preformed fungitoxic resorcinols (Droby et al., 1986, 1987). Flaishman
HO HO HO HO
(CH
2
)
11
CH
CH
(CH
2
)
3
CH
3
(CH
2
)
14
CH
2
CH
3
Resorcinol-5-(pentadecyl) Resorcinol-5-(12-heptadecenyl)
Fig. 7.1. Chemical structure of the 5-substituted resorcinols isolated from mango
fruits.
Fruit Diseases 215
and Kolattukudy (1994) hypothesized that ethylene is involved in terminat-
ing quiescence, by inducing appressoria formation and hyphal growth,
which strongly suggests that Colletotrichum spp. must have coevolved with
their hosts, to develop a mechanism that uses the host’s ripening hormone as
a signal to reactivate the infection process. However, when Prusky and co-
workers (Prusky et al., 1996) applied ethylene to unripe fruits they could not
enhance the termination of quiescence as had been suggested, possibly indi-
cating that ethylene is a signal that induces appressoria formation in vitro,
but that does not enhance fungal growth in the fruit. Furthermore, since fun-
gal infection and appressoria formation occur in the orchard, it is difficult to
conceive how ethylene produced much later, during ripening and storage,
could enhance appressoria formation on the fruit. Regardless of the mecha-
nisms that may be involved in the onset and termination of quiescence, qui-
escent infection appears to be a case of coevolution; it is advantageous to
both pathogen and host in a natural ecosystem to allocate chemical defences
to the immature fruit but not to the ripe fruit (Prusky and Keen, 1993).
Quiescence also may result from a localized host response that is often
associated with an oxidative burst, i.e. production of reactive oxygen species
(ROS). Localized generation of ROS during quiescence was found to be one
of the earliest (within 2–3 h) detectable cytological defence responses to
attempted penetration of unripe, resistant avocado fruits by C. gloeosporioides.
Beno-Moualem and Prusky (2000) proposed that fungal infection of unripe
fruits leads to production of ROS during quiescence, at the infection loci
surrounding the germinated appressoria and their penetration hyphae. This
localized ROS production would activate the synthesis of antifungal com-
pounds or compounds that inhibit fungal metabolism (Ardi et al., 1998) at the
infection loci, thereby enhancing and/or preserving the levels of antifungal
compounds and, in turn, inhibiting fungal development and imposing
quiescence.
Guetsky et al. (2005) suggested that initiation of quiescence could result
from the fungal capability to induce laccase activity that modulates the metab-
olism of preformed antifungal compounds and the activation of the quies-
cent C. gloeosporioides that occurs during fruit ripening. Early activity of
aggressive isolates in unripe fruits included increased laccase activities that
resulted in early metabolism of preformed antifungal compounds leading to
early appearance of symptoms.
When the levels of toxic compounds in the fruit peel decline, C. gloeospo-
rioides can also enhance its colonization of ripening fruits dynamically by
locally altering the pH of the fruit at the infection site to suit the increased
expression of pathogenicity factors and the enzymatic arsenal (Yakoby et al.,
2000; Prusky et al., 2001; Eshel et al., 2002; Prusky and Yakoby, 2003). In the
pathogen C. gloeosporioides, the gene pelB encoding for the enzyme pectate
lyase, a key factor for virulence, is expressed when the pH is > 6.0, a value at
which decay is initiated in the tissue (Yakoby et al., 2000, 2001). Also in the
case of A. alternata, another pathogen of mango fruits, the expression of the
endoglucanase gene AaK1 is maximal at pH levels > 6.0 (i.e. values that are
characteristic of decayed tissue). AaK1 is not expressed at the lower pH
D. Prusky et al. 216
values at which the pathogen is quiescent (Eshel et al., 2002). Ambient alkal-
ization by Colletotrichum is achieved by active secretion of ammonia, which is
produced as a result of protease activity and deamination of amino acids.
The pathogenicity of C. gloeosporioides and expression of the virulence factor
PL-B both depend on raising the ambient pH (Drori et al., 2003). This modu-
lation of environmental pH has been used as the basis for a new approach to
disease control in mango fruits, and is discussed below.
Management
Control of postharvest anthracnose can be achieved by field management,
postharvest treatments or a combination of both. Management of mango
anthracnose in the field involves cultural and chemical practices, as well as
cultivar selection.
Cultural control
Since the development of mango anthracnose is dependent on moisture or
high RH, orchards ideally should be established in areas with a well-defined
dry season, to allow for fruit development under conditions unfavourable
for disease development. In the tropics, mango flowering usually occurs dur-
ing dry seasons, but anthracnose incidence of > 90% is common in fruits that
develop during the rainy season (Arauz, 1999). In contrast, the incidence and
severity of mango anthracnose can be close to zero in fruits that develop
completely in the dry season, without the application of any other control
measure (Arauz, 2000).
Considerable effort has been invested in understanding and managing
mango flowering. Flowering can be advanced by several weeks by applying
potassium nitrate sprays to mature foliage (Núñez-Elisea, 1985). The growth
retardant paclobutrazol, alone or followed by potassium nitrate sprays, can
also be used to advance flowering (Núñez-Elisea et al., 1993). Both treatments
could contribute to the manipulation of the flowering season to a less sensi-
tive period. Field sanitation of the tree itself is difficult to practise. Elimina-
tion of dry panicles and mummified fruits is time consuming. Bagging results
in reduced anthracnose severity, but it also reduces the red colour of the fruit,
which could reduce consumer appeal (Hofman et al., 1997).
Resistant cultivars
Although all commercial mango cultivars are susceptible to anthracnose,
some are less susceptible than others; ‘Tommy Atkins’ and ‘Keitt’ are less
susceptible than ‘Irwin’, ‘Kent’, ‘Haden’ and ‘Edward’ (Campbell, 1992).
Chemical control
In extreme situations, in which fruit develops entirely under disease-favouring
conditions, seasonal applications of up to 25 sprays of protective and sys-
temic fungicides have been reported (Dodd et al., 1997). However, few fungi-
cides are approved in importing countries for use on mango. Therefore, the
Fruit Diseases 217
choice of fungicides depends on the intended destination of the fruit. Dithio-
carbamate fungicides are highly effective for anthracnose control but can be
used for only a few specific countries, whereas copper fungicides are recom-
mended, but their efficacy is lower (Arauz, 2000). Fungicides with erradicant
activity for mango anthracnose include benzimidazoles and the imidazole
prochloraz. Benomyl has been used under calendar-based schedules, usually
in a mix with protectant fungicides, to delay the build-up of resistance in the
pathogen population. Prochloraz has been used as a protectant or as an erad-
icant spray (Estrada et al., 1996), but is mainly used only as a postharvest
treatment.
Biological control
A number of microorganisms with in vitro or in vivo activity against C. gloeo-
sporioides have been isolated (Jeffries and Koomen, 1992), but few examples
had been used commercially in the field until Korsten (2004) isolated Bacillus
spp. from leaf and fruit surfaces, and effectively controlled anthracnose of
mango. Postharvest control was achieved with semi-commercial preharvest
sprays or postharvest packing house dips, sprays or ultra-low-volume appli-
cations. Integrated treatments involving antagonists combined with quarter-
strength or recommended dosages of fungicides such as prochloraz or sodium
hypochlorite also effectively suppressed postharvest anthracnose of mango.
Commercializing the antagonists in South Africa (Korsten, 2004) and in the
Philippines proved to be difficult because of the limitations set by the local
registration guidelines, and the effect of product formulation on antagonist
performance in commercial applications.
Postharvest control
Traditionally, postharvest control of mango anthracnose has aimed at eradi-
cation of quiescent infections on the fruit. Such eradication is achieved com-
mercially by thermal and chemical treatments, or a combination of both
(McMillan, 1987). Dipping fruit in hot water alone is moderately efficient;
temperatures of 50–55°C for 3–15 min have been recommended, with the
higher temperatures corresponding to the shorter exposures. Fruit from Latin
America entering the USA market must undergo a quarantine hot water
treatment to eliminate fruit fly (Ceratitis capitata and Anastrepha spp.) larvae;
the fruit is immersed in water at 46°C for 90–120 min, depending on variety
and fruit size. The efficacy of the fruit fly quarantine treatment varies from 60
to 85% for elimination of anthracnose infections (McGuire, 1991). Hot water
treatments leave no chemical residue on the fruit and could be a good anthra-
cnose control option for organically-produced mangoes or for mangoes tar-
geted for markets in the USA, where no fungicides are currently approved
for postharvest use. Temperature and time controls are critical, because fruits
can easily be damaged by overexposure to heat. Several fungicides have been
applied after harvest to control mango anthracnose. Benomyl, at rates vary-
ing from 500 to 1000 g/ml, was used as a postharvest dip in the past, but its
use is no longer permitted. Thiabendazole at 1000–2000 g/ml is also effec-
tive, and there is interest in its registration for postharvest use with mango,
D. Prusky et al. 218
as it is currently used on other fruits such as citrus. Prochloraz can be used

with fruit shipped to the European Union (EU), at rates up to 1000 g/ml; its
efficacy ranges from 65% under very high disease pressure to 94% under
moderate disease pressure (Arauz, 2000). One advantage of benzimidazole
fungicides (i.e. benomyl or thiabendazole) is that they are also effective in
controlling stem-end rot caused by L. theobromae, which is the second most
important postharvest disease of mango in tropical areas. Imidazoles such as
prochloraz and imazalil are not effective against L. theobromae on mango
(Estrada et al., 1996). The combination of hot water and fungicides is the most
effective commercial postharvest treatment for the control of mango anthra-
cnose; the rate of fungicide application and the duration of exposure to hot
water are both lower, and efficacy is higher, than either treatment used sepa-
rately. The hot water and the fungicides can be applied sequentially or together.
Irradiation of fruit to control anthracnose has been attempted: gamma irra-
diation was not successful (Spalding and Reeder, 1986), but a short-wave
infrared radiation treatment developed in South Africa resulted in anthra-
cnose levels similar to those resulting from the commercial hot-water treat-
ment, and was much faster and less expensive (Saaiman, 1996a).
Prusky and Keen (1993) suggested that it might be possible to prolong
the period of fruit resistance and to delay the onset of anthracnose develop-
ment until the fruit ripens. The climacteric rise occurs simultaneously with
the production of ethylene and with changes in external and internal colour,
flavour, aroma and firmness, and with a reduction in fruit resistance to fun-
gal attack (Prusky and Keen, 1993). Postharvest practices such as cold stor-
age and controlled atmosphere storage maintain resistance to decay by
delaying the ripening process, but there are two limitations to the potential
benefit of this approach in mango. First, mangoes, similarly to many other
tropical and subtropical fruits, are sensitive to chilling and are injured at tem-
peratures < 10–13°C, depending on the cultivar and the duration of expo-
sure. Second, once the fruits are allowed to ripen under ambient conditions,
disease develops normally. Nevertheless, some progress has been made
towards anthracnose control through maintaining fruit resistance beyond
the onset of ripening. In the last decade, a better understanding of the physi-
ological basis of quiescent infections on tropical and subtropical climacteric
fruits has been achieved, mainly through the work of D. Prusky and his co-
workers (1993). The decline in antifungal compounds, which is brought
about by oxidative processes, can be delayed so that it occurs closer to full
ripeness. In avocadoes and mangoes, the concentrations of antifungal com-
pounds were enhanced and, consequently, the decline in concentration was
delayed, by exposure of fruit to an atmosphere containing 30% carbon diox-
ide (CO
2
) for 24 h, or by treatment of fruit with the antioxidant compound
butylated hydroxy anisole (Prusky, 1988; Kobiler et al., 1998). The delay
resulted in less disease in ripe fruit.
Postharvest biological control of anthracnose in mangoes has been
attempted. In an investigation with a strain of a Bacillus sp. that exhibited in
vitro activity against C. gloeosporioides, it was found that disease control in
vivo was obtained when fruits were inoculated with the bacterium 24 h prior
Fruit Diseases 219
to inoculation with the fungus, but not when fruit were inoculated with the
pathogen first (Korsten et al., 1992), which indicated that the quiescent phase
of the fungus was not affected by the antagonist. Other approaches to disease
control using biological methods included the use of a non-pathogenic strain
of Colletotrichum magna that colonizes the fruit endophytically and prevents
infection by C. gloeosporioides (Prusky et al., 1993); and the expression of an
antifungal peptide in the yeast Saccharomyces, which controlled postharvest
diseases caused by C. coccodes (Jones and Prusky, 2001).
7.3 Alternaria Rot (Black Spot)
Alternaria alternata (Fr.:Fr.) Keissl. causes black spot of mango. Conidiophores,
arising singly or in small groups, are simple or branched, straight or bent,
and sometimes geniculate, and pale- to mid-olivaceous or gold in colour, and
smooth in texture. The conidia are oboclavata; they are borne in long chains
in culture, and most have three to five septa.
Pathogenesis
Germinated conidia penetrate mainly through wounds and specifically
through lenticels of the fruits, and then become quiescent.
Symptoms
Alternaria rot of mango has been increasingly reported as an important
pathogen that causes blossom disease and postharvest fruit rot in ripening
fruits in Australia, Egypt, India, Israel and South Africa. The symptoms are
small, black circular spots that develop around the lenticels. Initially, the
spots are concentrated around the stem end of the fruits, where there are
large numbers of lenticels. The spots can grow and coalesce to become a sin-
gle spot that covers a significant part of the fruit surface. At first, the decay is
firm and does not penetrate the pulp more than 1–2 mm, but later the disease
progresses into the flesh, which darkens and becomes soft (Prusky et al.,
1983). Symptoms of alternaria rot are more limited, darker and firmer than
those of anthracnose. The former pathogen also attacks mango leaves, and
symptoms can be observed throughout the year. The pathogen may also
attack mango inflorescences, resulting in a significant decrease in fruit set.
Epidemiology
The main sources of inoculum are conidia released from infected leaves,
twigs and inflorescences; however, Alternaria spores easily can be found in all
the dry tissues of mango trees in the orchard. Conidia are transferred to the
fruit by air currents and in dew runoff (Ploetz et al., 1994). Germination of
conidia depends on the RH in the orchard during fruit growth. The area of
quiescent Alternaria infection on mango fruit at harvest increased as the num-
ber of hours of exposure to RH ≥ 80% increased over 320 h (Prusky et al.,
D. Prusky et al. 220
1983). Regions with the highest potential for disease incidence are located
close to the 85–90% RH isolines during the fruit growth period. The interme-
diate regions lie between 75 and 85% RH, and the lowest potential risk is in
the dry regions, where the prevailing RH is < 75% (Prusky et al., 1992).
Mechanisms of fungal pathogenicity and fruit resistance
As in the case of anthracnose, the mechanism of resistance against Alternaria
is also related to the presence of high concentrations of preformed antimicro-
bial compounds (Droby et al., 1986, 1987). Susceptibility was found to depend
on the decline of the compound to subfungitoxic concentrations, which
occurred faster in ‘Tommy Atkins’ than in ‘Keitt’.
Alternaria alternata can also alter the local fruit pH at the infection site
dynamically, to match the increased expression of pathogenicity factors and
the enzymatic arsenal (Eshel et al., 2002; Niem et al., 2007). In the case of A.
alternata, the expression of the endoglucanase gene AaK1 was found to be
maximal at pH levels > 6.0, which are characteristic of decayed tissue; it was
not expressed at the lower pH values at which the pathogen was quiescent
(Eshel et al., 2002). Alkalization of the ambient environment by Alternaria is
achieved by active secretion of ammonia (Eshel et al., 2002), probably as a
result of protease activity and deamination of amino acids. This modulation
of environmental pH has been used as the basis of a new approach to disease
control in mango fruits, and is discussed below.
Management
The losses caused by alternaria rot can be minimized by a regular field-
spraying programme and by postharvest fungicide treatments.
Field and postharvest chemical control
Preharvest treatments with dithiocarbamate fungicides inhibit the develop-
ment of latent infection. Three sprays with the protectant fungicide maneb,
starting 2 weeks after fruit set, are most effective for disease control (Prusky
et al., 1983). However, since quiescent infections do not develop until after
harvest and ripening, the application of a postharvest treatment by spraying
the fruits on the packing line with 900 g/ml prochloraz is simpler and more
efficient than the preharvest fungicide treatment.
Control of alternaria rot is significantly improved by a combination of
physical and chemical treatments. The physical treatment includes a 15–20 s
hot water spraying and brushing (HWB) treatment at temperatures between
50 and 55°C (Prusky et al., 1999). This new approach improved fruit quality
at the same time as it reduced disease incidence. If this physical treatment is
followed by a 250 g/ml prochloraz spray it can further improve disease
control. Prusky et al. (1999) concluded that the type and strength of the post-
harvest treatment should be varied according to the level of quiescent infection
Fruit Diseases 221
of A. alternata at the time of harvest. Although the fungicide prochloraz is
very effective for the control of postharvest disease, a milder postharvest treat-
ment, such as chlorine at 500 g/ml, can be applied to fruit in which a low
incidence of quiescent infections is found at harvest (Prusky et al., 2002).
This postharvest physical-chemical treatment has been further improved
in light of the recent finding that A. alternata pathogenicity may modulate the
pH of the host environment to promote colonization (Eshel et al., 2002; Prusky
and Yakoby, 2003; Prusky and Lichter, 2007). Application of a combination of
HWB for 15–20 s, followed by spraying with 50 mM hydrochloric acid (HCl),
effectively controlled alternaria rot in stored mango fruit. Similar HWB treat-
ments followed by spraying with reduced concentrations of prochloraz at
45 g/ml in 50 mM HCl inhibited alternaria rot development better than
treatment with HCl alone (Prusky et al., 2006). The enhancement of prochlo-
raz activity in acidified solutions was attributed to its enhanced solubility
under acidic conditions, which resulted in an increase in the fungicidally
active ingredient in the solution. This technology represents the latest devel-
opment in the control of postharvest diseases, and provides a simple treat-
ment for the control of diseases that alkalinize the host environment,
including both alternaria rot and anthracnose.
7.4 Stem-end Rots
Stem-end rots of mango fruit present one of

the most serious postharvest
problems that affect this industry

worldwide. They become more prominent
as orchards become older. Losses increase when fruits are stored for pro-
longed periods at low temperatures or when fruits ripen at temperatures
> 28°C. The stem-end rot diseases are caused

by a variety of fungal patho-
gens including: D. dominicana (anamorph of Botryosphaeria dothidea), Dothi-
orella mangiferae, L. theobromae (Botryodiplodia theobromae), Phomopsis mangiferae
and Pestalotiopsis mangiferae, among which Botryosphaeria

is the dominant
pathogen (Darvas 1991; Johnson et al., 1992; Sangchote, 1998). Stem-end rot
diseases cause heavy losses in mangoes during storage (Prusky et al., 1992;
Mitra, 1997; Kobiler et al., 2001).
Pathogenesis
Johnson and co-workers (1993) have suggested that spores of the pathogen
may germinate and penetrate into the host tissue through wounds, and
remain as endophytes in the branches of mango trees.
Symptoms
Depending on the fungus involved, a variety of symptoms may develop at
the stem and as the fruit ripens. Infections by L. theobromae (B. theobromae), D.
dominicana synonym Fusicoccum aesculi (anamorph of B. dothidea) and D.
mangiferae, cause the formation of diffuse, water-soaked tissue that spreads
D. Prusky et al. 222
from the stem end as fingerlike projections, which darken and coalesce into
circumpedicular lesions or lobed margins (Johnson et al.,

1992; Slippers et al.,
2004, 2005). Necrosis remains beneath the cuticle and may penetrate through-
out the fruit flesh. Superficial mycelia may appear around the pedicel and
ruptures of the skin or, in some cases, may penetrate through the epidermis.
The ascomata of D. mangiferae are initially embedded, either separately or
grouped in complex stromata, and they finally erupt through the epidermis
and open. The spore wall is dark and thick, and becomes thinner towards the
end. Conidiophores are hyaline, cylindrical, smooth and branched at the
base. A watery fluid may drain from the stem or from surface ruptures (Kor-
sten, 2004). Diseased fruit could infect a whole box of fruits by direct contact,
and thereby spread the pathogen in the box. In the case of injured fruit,
lesions could appear at some distance from the stem. Stem-end rot diseases
also have a major effect on the flavour of the fruit.
The disease may also cause tip and branch dieback and cankers on mango
trees (Ramos et al., 1991). Anamorph morphology is commonly used to iden-
tify Botryosphaeria spp. (Jacobs and Rehner, 1998; Slippers et al., 2004), but the
morphological

distinctions among the anamorphs of some of the closely
related

species are not clear. Recent studies based on DNA sequence

data have
highlighted taxonomic groups and relationships in

Botryosphaeria (Jacobs and
Rehner, 1998; Slippers et al., 2004). These data, combined with morphological

characteristics, could clarify the current taxonomic confusion.
Phomopsis mangiferae is a weak parasite of less economic importance than
the species above; it produces a dark, circumpeduncular lesion with defined
edges that spreads relatively slowly but penetrates deeply into the flesh.
Fruiting bodies may develop on the affected surface. Phomopsis mangiferae
can also be distinguished by a dark pinhead-size pycnidial fruiting body
(Johnson et al., 1992). The lesion caused by Phomopsis may be similar to the
stem-end symptoms cause by C. gloeosporioides and A. alternata. However, the
lesions of the latter two diseases penetrate only to a depth of 10–20 mm.
Lesions of L. theobromae can be distinguished by their wrinkled black
edges which have a velvety appearance. In the dark zone, pycnidial masses
are formed (Johnson et al., 1992). In affected plants, twigs die back from the
tips and into old wood, giving a scorched appearance to the limb with abun-
dant gum secretions from branches, stems and the main trunk. Pestalotiopsis
mangiferae appears as silvery grey spots that vary in size, and are usually
sharply outlined by a dark border. The fungus may appear as a member of
the complex of stem-rot pathogens.
Epidemiology
The pathogens causing stem-end rots initiate inoculation in the orchards as
the trees age. They are enabled to do so by mismanagement or neglect of
orchards, and are affected by periods of rain and high RH at the beginning
and end of the dry season (Johnson et al., 1991; Lonsdale, 1993; Johnson and
Sangchote, 1994; González et al., 1999). Hyphae of the fungi colonize the
Fruit Diseases 223
floral parts of mango trees, develop endophytically in healthy tissue of all
parts of mango plants, especially in the fruit pedicel, and remain quiescent
until the fruit matures, at which stage the parasite is ready to infect through
the stem end by developing in the xylem vessels of the maturing fruit (John-
son et al., 1992). Pathogens can colonize flower parts, remain inactive pend-
ing button abscission and then penetrate the stem end, as in the case of
Diplodia natalensis in citrus (Nadel, 1944), of which no sexual stage has been
found in nature. Fruits can be also infected at the stem by the soilborne patho-
gen L. theobromae (anamorph of B. theobromae), if the fruits are placed on the
ground (Johnson et al., 1992) after harvest. It is also possible that infection can
result from transfer of spores by insects; decayed fruits produce volatiles,
which are hypothesized to be attractants of an insect that could be a vector of
the fungal spores (Nago and Matsumoto, 1994).
The range of pathogens that cause stem-end rot is influenced by tempera-
ture, moisture stress and the nutrition status of the host. The initial systemic
infection plays a crucial role in establishing blossom-blight infection. How-
ever, secondary infection is apparently an even more important factor in devel-
opment and incidence of soft stem-end rots. Secondary infections occur when
rain washes spores away from various inoculum sources such as leaves and
stems (Saaiman, 1996b). Endophytic Botryosphaeria spp. were found to be espe-
cially prominent in trees continually exposed to water shortage and salt stress
(Grobler et al., 2002), and Botryosphaeria spp. were found to move into develop-
ing fruit, resulting in postharvest stem-end rot development (Lonsdale, 1993).
Management
Field and postharvest control
A variety of postharvest practices can be used to delay disease develop-
ment. These include low-temperature storage and modified- or controlled-
atmosphere storage; however, the management of stem-end rots is far from
being perfected.
Cultural control
Johnson et al. (1992) demonstrated that infection of mango fruit before har-
vest occurred through endophytic colonization of pedicel tissue by Botry-
osphaeria spp. present from previous growth flushes, and the possibility of
pruning to promote growth flushing was tested as a means to reduce inocu-
lum in the stem tissue from which new-season inflorescences emerged. Cooke
et al. (1998) reported that the levels of endophytic organisms such as Botry-
osphaeria spp. were reduced significantly when a pruning programme was
implemented in mango orchards as a preharvest control measure. Korsten
(2006) found that prevention of water stress during fruit development and
maturation, and avoidance of placing fruits on the ground suppressed dis-
ease development. He also suggested that harvesting of immature fruit
should be avoided; fruit should be cooled to 13°C immediately after harvest
and chemical treatment, and stored in a well-ventilated place.
D. Prusky et al. 224
Chemical control
Postharvest control of Botryosphaeria spp. was achieved by postharvest dip-
ping, spraying or ultra-low-volume application of benomyl (where possible).
Prochloraz or sodium hypochlorite also effectively suppressed postharvest
stem-end rot of mango (Plan et al., 2002; Korsten, 2006). A combined treat-
ment of wax and hot water (55C) provide very effective control of most
postharvest pathogens (Sangchote, 1998), but in some cases partial-vacuum
infiltration improved disease control, which suggests that control efficiency
may have been reduced because the fungicide did not reach the pathogen
(Plan et al., 2002).
A treatment consisting of HWB and prochloraz followed by 2,4-dichloro-
phenoxyacetic acid (2,4-D) diluted in wax, reduced side and stem-end decay
by 50–70%, and improved fruit quality during prolonged storage (Kobiler et al.,
2001). The best control was obtained by concentrations of 2,4-D ranging from
75 to 175 g/ml; this efficiently controlled side rots caused by A. alternata and
the stem-end rots caused by A. alternata, Phomopsis spp. and Lasiodiplodia
spp. The combination of HWB, prochloraz application and 2,4-D treatment
reduced the incidence of stem-end rot after 4 weeks of storage at 14°C and 7
days of shelf life at 20°C from 86 to 10% in ‘Tommy Atkins’ and from 63 to
12% in ‘Keith’ (Kobiler et al., 2001).
Biological control
Bacillus licheniformis, on its own or alternated with copper oxychloride, has
been evaluated as a preharvest spray treatment to control mango fruit dis-
eases. Preharvest applications of B. licheniformis at 3-week intervals from
flowering until harvest controlled moderate levels of anthracnose, and of soft
rot caused by Botryosphaeria, which suggests a potential treatment for com-
mercial preharvest applications (Silimela and Korsten, 2006).
7.5 Other Minor Diseases
Other pathogens may attack mango fruits after harvest through occasional
wounds, and cause severe diseases, such as black mould caused by Aspergillus
spp. and transit rot caused by Rhizopus spp. Disease control begins in the field,
and is followed by postharvest sanitation, and avoidance of latex burn (stain)
and mechanical injury. A hot water treatment consisting of 46°C for 60–120
min and fungicides can be used, depending on the cultivar (Spalding and
Reeder, 1986). HWB at 55°C for 20 s shows good control (Prusky et al., 1999).
7.6 Conclusions
Long periods in transit, new marketing approaches for mangoes (e.g. ‘Ready
to Eat’) and stringent international standards and requirements have raised
the need for improved approaches to disease control, in order to preserve
fruit quality. Integrated postharvest treatment has provided a more durable,
Fruit Diseases 225
consistent, sustainable and practical solution than previously available to
enable producers to control postharvest diseases. Ensuring product quality
and safety starts in the orchard, with proper handling conditions and is fol-
lowed by optimal postharvest treatments; however, the combination of post-
harvest treatments with the proper use of the cold-chain concept may provide
the ultimate in favourable conditions for preserving fruit quality.
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© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 231
8 Foliar, Floral and Soilborne Diseases
R.C. Ploetz
1
and S. Freeman
2
1
University of Florida, Florida, USA
2
Agricultural Research Organization, Bet Dagan, Israel
8.1 Introduction 232
8.2 Foliar and Floral Diseases 232
Algal leaf spot 232
Alternaria leafspot 233
Anthracnose 235
Apical necrosis 240
Bacterial black spot (black canker) 241
Black-banded disease 245
Black mildew, sooty blotch and sooty mould 246
Blossom blight 248
Decline disorders 249
Galls and scaly bark 256
Grey leafspot 259
Leaf blight 260
Malformation 261
Parasitic plants 270
Phoma blight 270
Phoma leafspot 270
Pink disease 271
Powdery mildew 271
Scab 274
Seca and sudden decline 274
Stigmina leafspot 279
8.3 Soilborne Diseases 281
Black root rot 281
Nematode damage 281
Phytophthora diseases 282
Root rot and damping off 284
Sclerotium rot 285
Verticillium wilt 286
White root disease 286
8.4 Conclusions 289
R.C. Ploetz and S. Freeman 232
8.1 Introduction
Diseases affect all tissues and developmental phases of mango. Mango dis-
eases have been reviewed in the first edition of this book (Litz, 1997) and by
Singh (1968), Cook (1975), Snowdon (1990), Ploetz et al. (1994) and Ploetz
(2003). Lim and Khoo (1985), Prakash and Srivastava (1987) and Ridgeway
(1989) have also reviewed this subject. This chapter focuses on foliar, floral
and soilborne diseases of mango. Each is discussed with respect to signifi-
cance, geographical distribution, history, symptoms, causal agent(s), epide-
miology and management. These diseases are caused mainly by eukaryotes
(Domain Eukaryota) among which the true fungi, Eumycota (Ascomycota
and Basidiomycota), predominate. Other less important eukaryotes include
the fungus-like oomycetes (Oomycota), nematodes (Metazoa), and parasitic
plants and green algae (Plantae). With one exception, relatively minor pathogens
of mango are prokaryotes in the Domain Eubacteria; all are Gram-negative
-proteobacteria. None of these diseases is caused by other plant-pathogenic
eukaryotes (protozoa), prokaryotes (- and -proteobacteria, Mollicutes and
Firmicutes), or the nucleic acid-based pathogens, the viruses and viroids.
8.2 Foliar and Floral Diseases
Diseases above ground are the most important and conspicuous problems on
mango. Since many of the pathogens that incite foliar diseases also affect
panicles, diseases on each are combined in this section, as are a few diseases
that affect the branches and trunks of mature trees.
Algal leaf spot
Algal leaf spot, also known as red rust, is caused by a parasitic green alga,
Cephaleuros virescens Kunze (synonyms: Cephaleuros parasiticus Karst, Ce-
phaleuros mycoidea Karst) (family Trentepohliaceae, division Chlorophyta)
(Lim and Khoo, 1985). It is a common problem on mango and many other
tropical and subtropical plants (Joubert and Rijkenberg, 1971). Although
algal leaf spot can cause serious problems on tea Camellia sinensis (L.) O.
Kuntz, black pepper, Piper nigrum L., and other important crops, it is usually
debilitating on mango only in poorly managed orchards (Lim and Khoo,
1985). In the latter cases, abiotic or biotic stresses, such as mites, insects and
other foliar diseases, can increase the severity of this disease.
Conspicuous symptoms of algal leafspot are orange to rust-coloured,
velutinous patches on both leaf surfaces (Plate 43) (Lim and Khoo, 1985).
They are initially 5–8 mm in diameter, but can merge to involve large, irregu-
lar sections of the leaf. They later assume a dull, greyish green colour, and
eventually become bleached patches. The alga can also affect twigs and
branches, causing the bark to crack as the pathogen’s filaments extend into
the host cortical tissues. The orange tufts produced by C. virescens are the
algal thallus located beneath the host cuticle. They produce erect cells, some
Foliar, Floral and Soilborne Diseases 233
of which enlarge to produce stalked, terminal or ovoid, 30 × 24 m, sporangia
(Fig. 8.1). Sporangia produce biflagellate zoospores that are dispersed by
rainsplash and wind and are the primary infective propagules. Flask-shaped
gametangia in the thallus are responsible for sexual reproduction. In free
water, they release 8–32 biflagellate gametes, pairs of which fuse and give
rise to dwarf sporophytes. The sporophytes produce microsporangia that
bear quadriflagellate zoospores; their role is not known.
Cephaleuros virescens requires a humid environment to establish and
spread (Lim and Khoo, 1985). Pruning the canopy, mowing beneath trees and
wider row spacings to increase air circulation and sunlight penetration all
help combat the disease. Proper fertilization and irrigation, and control of
insect pests, mites and other foliar diseases increase the tree’s ability to cope
with algal leafspot. Algacides or fungicides (i.e. fentin acetate or those con-
taining copper (Cu)) can be used to control the disease in severe cases.
Alternaria leafspot
Alternaria alternata (Fr.) Kreissler (synonyms: Alternaria fasciculata (Cooke
and Ellis) L. Jones and Grout, Alternaria tenuis Nees, and Macosporium
Fig. 8.1. Branch of the thallus of Cephaleuros virescens that has terminated in several
oval sporangia (Photograph courtesy of T.-K. Lim).
R.C. Ploetz and S. Freeman 234
Fig. 8.2. Conidia and conidiophores of Alternaria alternata (Source: Ellis, 1971).
fasciculatum Cooke and Ellis; no teleomorph is known) causes black spot on
mango, alternaria leaf spot and lesions on inflorescences (Prusky et al., 1983;
Cronje et al., 1990). Although the fungus is cosmopolitan and has a large
number of host plants (Neergaard, 1945; Domsch et al., 1980), the diseases it
causes on mango are most prevalent in arid environments. In Israel, it is a
more important disease on fruit than leaves.
Symptoms on leaves are round, dark spots, 1–3 mm in diameter; they are
most prevalent on the underside of leaves (Prusky, 1994). Similar spots that
occur on the rachis of inflorescences can reduce fruit set (Cronje et al., 1990).
Conidiophores of the causal fungus originate alone or in small groups, and
are smooth and pale to mid-olivaceous or golden brown. They are sometimes
geniculate, and vary between simple and branched, and straight to flexuous.
Conidia are obclavate, 20–36 × 9–9.5 m, three- to five-septate and borne in
long chains (Fig. 8.2).
Foliar, Floral and Soilborne Diseases 235
Certain antifungal compounds are associated with the latency of A. alter-
nata infections on fruit (Droby et al., 1986, 1987). Whether the same com-
pounds are formed in leaves and inflorescences has not been reported. Infected
leaves, twigs, inflorescences and leaf litter are significant sources of inocu-
lum for fruit infection. Conidia of A. alternata are dispersed by air currents
(Prusky, 1994). Several different fungicides and combined treatments are
effective against the diseases that are caused by this pathogen. Postharvest
development of alternaria fruit rot during storage is usually prevented by a
combination of hot water brushing in combination with prochloraz at
225 g/ml (Prusky et al., 2002).
The finding that A. alternata alkalinizes the host pH (Eshel et al., 2002)
prompted investigations into modulating environmental pH with acid treat-
ments to reduce fungal colonization. Spore germination and germ-tube elon-
gation of A. alternata in vitro were inhibited by, respectively, 95 and 65% by
exposure to 1.25 mM hydrochloric acid (HCl). Germination was completely
inhibited by 2.5 mM HCl (Prusky et al., 2006). Acidified solutions containing
45 g/ml prochloraz inhibited alternaria rot development better than treat-
ment with HCl alone (Eshel et al., 2002). These results have not been extended
to control foliar and floral diseases that are caused by this pathogen.
Anthracnose
Anthracnose is the most important disease of mango (Cook, 1975; Lim and
Khoo, 1985; Ploetz, 2003), particularly on fruit in humid, high rainfall envi-
ronments. It is usually replaced by, and is less important than, other diseases
in dry production areas. Anthracnose can also be a serious problem on foli-
age and flowers. Crowded and moist conditions in nurseries can result in
significant damage to young leaves and, in extreme cases, new orchards have
been devastated (Bose et al., 1973). Blossom blight, which has been attributed
to one of the anthracnose agents but is also caused by other fungi, is covered
separately below.
Symptoms
On panicles, necrotic flowers abscise leaving persistent peduncles. Small, cir-
cular dark spots also develop on pedicles and peduncles. Lesions may enlarge
and coalesce to form large patches of necrotic, dark brown tissue. With suf-
ficient rainfall, salmon- to orange-coloured fructifications of the pathogen
develop on the affected tissues. On leaves, lesions are dark brown and sur-
rounded by chlorotic haloes, have irregular, rounded margins, and are not
delimited by veins (Fig. 8.3). Lesions are 0.5–1.0 cm in diameter on mature
leaves, but can expand on young leaves. Eventually, large, necrotic patches
can develop that deteriorate and fall from the leaf giving it a tattered appear-
ance. Although different mango cultivars are known to vary considerably in
their resistance to anthracnose on fruit, reports on the foliar and floral resis-
tance of different cultivars and whether resistances of the various organs are
related have not been published.
R.C. Ploetz and S. Freeman 236
Aetiology
In most production regions, anthracnose is caused by the ascomycete fungus,
Colletotrichum gloeosporioides Penz. (teleomorph: Glomerella cingulata (Stonem.)
Spauld. and Schrenk.) (Cook, 1975; Snowdon, 1990; Ploetz, 2003). In Austra-
lia, Florida USA, India, Japan and Taiwan, Colletotrichum acutatumSimmonds
(teleomorph: Glomerella acutata) plays a minor role (Fitzell, 1979; Prakash,
1990; Weng and Chuang, 1995; Taba et al., 2004; Tarnowski, unpublished). In
Colombia, Colletotrichum boninense J. Moriwaki, Toy. Sato and T. Tsukiboshi
has been reported (Afanador-Kafuri et al., 2003).
Colletotrichum spp. cause diseases on several subtropical and tropical
hosts (Jeffries et al., 1990; Freeman et al., 1998). Cultures of the fungus on
potato dextrose agar (PDA) are greyish white to dark grey and usually pro-
duce an aerial mycelium ranging from a thick mat to sparse tufts (Holliday,
1980). Conidia are hyaline, unicellular and either cylindrical with obtuse ends
or ellipsoidal with a rounded apex and a narrow, truncate base. They are
7–20 × 2.5–5 m, and are formed on hyaline to faintly brown conidiophores in
Fig. 8.3. Foliar symptoms of anthracnose (Photograph courtesy of S. Freeman).
Foliar, Floral and Soilborne Diseases 237
acervuli that are irregular in shape and about 500 m in diameter. Setae are
4–8 × 200 m, one- to four-septate, brown and slightly swollen at the base and
tapered at the apex. Hyphopodia have been used to distinguish isolates of C.
gloeosporioides and C. acutatum (Du et al., 2005), but provided ambiguous
results in Florida (Palmateer et al., 2007).
Characterization and taxonomic identification of Colletotrichum spp. has
relied on morphology and host range (Freeman et al., 1998; Du et al., 2005). In
general, C. gloeosporioides (Fig. 8.4) produces longer and narrower conidia
than C. acutatum (Fig. 8.5), as well as circular vs lobed hyphopodia. However,
variability in culture and host range has made these criteria unreliable for
species identification (Adaskaveg and Hartin, 1997; Freeman et al., 1998). Col-
letotrichum gloeosporioides and C. acutatum are species complexes with a large
number of hosts that are so broadly defined that the names are of ‘limited use
to the taxonomist or plant health practitioner’ (Du et al., 2005). Several molec-
ular tools have been implemented to differentiate within and among Col-
letotrichum spp., including: species-specific polymerase chain reaction (PCR)
primers, random amplified polymorphic DNA (RAPD) and arbitrarily
primed PCR (Freeman and Rodriguez, 1995; Afanador-Kafuri et al., 2003);
A+T-rich DNA analyses (Freeman et al., 1993); sequence analyses of the inter-
nal transcribed spacer (ITS) regions of ribosomal DNA (rDNA) (Sreenivasap-
rasad et al., 1996; Freeman et al., 2001; Afanador-Kafuri et al., 2003) and
MAT1-2 mating type sequences (Du et al., 2005).
(a)
(b)
(c)
(d)
(g)
(e)
(f )
10µ
50µ
Fig. 8.4. (a) Acervulus and emergent setae, (b) conidiophores, (c) conidia, (d) perithecium, (e)
asci, (f) ascospores and (g) appressoria at hyphal termini of Glomerella cingulata (anamorph:
Colletotrichum gloeosporioides) (Source: from Commonwealth Mycological Institute (CMI)
description no. 315).
R.C. Ploetz and S. Freeman 238
Relatively few studies have examined isolates of C. gloeosporioides from
mango. Gantotti and Davis (1993) reported that isolates from different organs
produced different pectin-degrading enzymes, whereas Alahakoon et al.
(1994b) indicated that > 80% of isolates from mango leaves, inflorescences
and fruit in Sri Lanka were identical, based on restriction fragment length
polymorphisms (RFLPs). Rivera-Vargas et al. (2006) reported that isolates of
C. gloeosporioides from leaves, fruit, flowers and panicles caused similar dis-
ease on detached leaves. Work is needed to clarify whether, and the extent to
which, isolates from different mango organs differ, and whether there is a
relationship among disease reactions on various organs.
Analyses of RFLPs and RAPDs indicated that isolates from mango are
genetically uniform and differ from those recovered from avocado, caram-
bola and papaya (Hodson et al., 1993; Alahakoon et al., 1994a; Hayden et al.,
1994). Isolates genetically similar to those from the latter crops occur infre-
quently on mango, and mango isolates were not found on the other plants
and were usually virulent only on mango. Alahakoon et al. (1994b) suggested
(c)
(d)
10 µ
(a)
25 µ
(b)
10 µ
Fig. 8.5. (a) Acervulus, (b) conidiogenous cells and setae, (c) conidia and (d) appres-
soria of Colletotrichum acutatum, anamorph of Glomerella acutata (Source: from CMI
description no. 630).
Foliar, Floral and Soilborne Diseases 239
that the mango isolates comprise a C. gloeosporioides population that was dis-
seminated worldwide from a single source, perhaps as an endophyte.
A recent study identified Colletotrichum spp. that infect mango, passion-
fruit (Passiflora spp.) and tamarillo (Solanum betacea cav. Sendt.) in Colombia
and assessed whether cross-infection between the host species occurred
(Afanador-Kafuri et al., 2003). With species-specific PCR primers, most of the
mango isolates were identified as C. gloeosporioides. However, DNA of the
passionfruit isolates and single isolates from tamarillo and mango were not
amplified by either C. acutatum- or C. gloeosporioides-specific primers; they
were identified later as C. boninense (Freeman, unpublished data). Further
molecular analyses determined that the isolates of C. gloeosporioides from
mango were heterogeneous, but that the population of C. boninense from pas-
sionfruit, tamarillo and mango was uniform; it may not be host specific.
The origins and diversity of C. gloeosporioides on mango require more
study. Furthermore, whether distinct populations of this diverse species
occur on different mango cultivars and organs, and whether disease reac-
tions on one mango organ could be used to predict those on another should
be determined. These results would be relevant to host resistance and
improvement of the crop, as well as the chemical and cultural control of this
important disease.
Epidemiology and management
Fitzell and Peak (1984) determined that conidia were the most important
inoculum in Australia. They are produced on branch terminals, mummified
inflorescences, flower bracts and leaves. New leaf flushes are the most signifi-
cant source of inoculum, and this was corroborated by Dodd et al. (1991) in the
Philippines. Optimum production of conidia occurred between 25 and 30°C
when free moisture was available, but also formed at 95–97% relative humid-
ity (RH). The pathogen’s teleomorph played no apparent role in the spread of
the disease (Fitzell and Peak, 1984). The threat posed by C. gloeosporioides to
fruit production is greatest in areas with heavy rainfall. In general, this begins
with the onset of flowering (Lim and Khoo, 1985; Jeffries et al., 1990). During
commercial production these diseases are managed with diverse chemicals.
Registration of the various pesticides varies in different production areas
(Jeffries et al., 1990).
Many of the most effective fungicides are systemic. They have selective
modes of action, but are prone to lost or reduced efficacy due to the develop-
ment of resistance in the targeted pathogens. Resistance in C. gloeosporioides
to benomyl is now common (Maymon et al., 2006), and there is inherent toler-
ance in C. acutatum to this fungicide (Freeman et al., 1998; Peres et al., 2002).
Newer strobilurin fungicides that are also susceptible to resistance problems
must be used properly (no more than three applications per season and alter-
nated with fungicides with different modes of action) to ensure that their
efficacy is not lost.
Although fungicide applications usually focus on reducing damage to
fruit, disease control on foliage is indicated in some, and on inflorescences
in most situations. Trees in nurseries usually require protection if they are
R.C. Ploetz and S. Freeman 240
crowded or receive overhead irrigation. Since infected foliage and branch
terminals represent important reservoirs of inoculum for blossom and fruit
infection, fruit set and anthracnose control on fruit are enhanced if disease
control is exercised prior to flowering (Jeffries et al., 1990). Off-season control
measures are beneficial in production environments that receive significant
rainfall.
Apical necrosis
Apical necrosis was first reported in Spain in 1991, and now occurs in Israel,
Italy, Portugal and possibly Egypt (Cazorla et al., 1998, 2006; F.M. Cazorla,
Málaga, 2007, personal communication). The disease can be quite damaging,
and limits production when panicles are affected. Apical buds, leaves and
panicles are susceptible (Fig. 8.6a–c), but fruit are not (Cazorla et al., 1998).
Vegetative and floral apices are affected by a dark-brown to blackish necrosis
(Fig. 8.6a, c). Necrosis on leaves begins as blackened, water-soaked lesions,
1–3 mm in diameter, that can coalesce and expand to cover large areas (Fig.
8.6c). Necrosis also extends from affected buds to petioles, through the leaf
midrib, and can cover large areas (Fig. 8.6a). A milky bacterial exudate often
develops on affected apical buds, but infrequently on petioles (Fig. 8.6b).
Large portions of the canopy and high numbers of flowers can be killed.
Fig. 8.6. Symptoms caused by the apical necrosis pathogen, Pseudomonas syringae
pv. syringae. (a) Extensive necrosis on young stem, apical bud, petioles and leaves;
(b) bacterial exudate on necrotic stem; and (c) death of a developing floral panicle and
associated leaf necrosis (Photographs courtesy of F.M. Cazorla).
(a) (b) (c)
Foliar, Floral and Soilborne Diseases 241
The causal bacterium, Pseudomonas syringae pv. syringae van Hall, affects
many perennial fruit crops (Hirano and Upper, 1983; Kennelly et al., 2007). It
is an epiphyte and is generally not an aggressive pathogen. Disease usually
develops after high populations of the bacterium develop in host tissues as a
result of host predisposition. Strains from mango produce an antimetabolite
toxin, mangotoxin, which plays a role in pathogen virulence and symptom
development (Arrebola et al., 2007).
Cold, wet weather and host genotype are primary factors in the develop-
ment of apical necrosis (Cazorla et al., 1998, 2006). ‘Tommy Atkins’, ‘Lippens’
and ‘Manzanillo’ are very susceptible whereas ‘Keitt’ and ‘Sensation’ are less
so. Apical necrosis is managed in commercial orchards with Cu-containing
pesticides, although there has been a recent increase in control failures and
Cu resistance in Spain and Portugal (Cazorla et al., 2002, 2006). These out-
breaks have been associated with several different Cu-resistance plasmids in
the causal bacterium. Carzorla et al. (2006) determined that the plant resis-
tance activator acibenzolar-S-methyl and the phosphonate derivative fosetyl-Al
provided control comparable to Bordeaux mixture, and that the latter treat-
ment might protect plants due to the protective film it provides against
wound entry for the pathogen.
Bacterial black spot (black canker)
Bacterial black spot is a destructive leaf, stem and fruit disease in many pro-
duction areas (Gagnevin and Pruvost, 2001). In India, the disease is called
bacterial canker or black canker due to the cankers it causes on the stems of
some cultivars (Prakash et al., 1994). It can be the most important disease
where fungal-induced diseases are well controlled (Gagnevin and Pruvost,
2001). Bacterial black spot has been identified in Australia, Myanmar, the
Comoros, India, Japan, Kenya, Malaysia, Mauritius, New Caledonia, Paki-
stan, the Philippines, Réunion, Rodrigues, South Africa, Taiwan, Thailand
and the United Arab Emirates (UAE) (Fukuda et al., 1990; Pruvost et al., 1992;
Prakash et al., 1994; Gagnevin and Pruvost, 1995, 2001; Kishun, 1995; Ah-You
et al., 2007b). Given the ease with which the pathogen is disseminated in
propagation materials and its wide, confirmed distribution, bacterial black
spot may be more widely spread than is currently recognized.
Symptoms
Mango leaves, stems and fruit are affected (Manicom and Pruvost, 1994;
Gagnevin and Pruvost, 2001). On leaves, water-soaked spots are initially 1–3
mm in diameter. As they enlarge, they become raised, black and angular, are
limited by veins and surrounded by chlorotic haloes (Fig. 8.7). These lesions
are larger and more conspicuously raised than those caused by other xan-
thomonads that have been recovered from other species in the Anacardiaceae
(Ah-You et al., 2007a). Lesions can merge to form large necrotic patches, and
bacteria may ooze from lesions during wet conditions. Old lesions dry out,
turn white or grey, and crack. Defoliation occurs in severe cases. Anthracnose
R.C. Ploetz and S. Freeman 242
lesions are not raised or as black and angular as those caused by bacterial
black spot. On branches, bacterial black spot lesions are dark and cracked
along the long axis (Fig. 8.8). They develop only on highly susceptible culti-
vars and are often associated with wounds. Conspicuous, star-shaped lesions
are produced on fruit.
Aetiology
Diverse xanthomonads have been recovered from mango and other hosts in
the Anacardiaceae (Gagnevin and Pruvost, 2001; Ah-You et al., 2007a). Only
some of these cause symptoms of bacterial black spot. Early reports that this
disease is caused by Pseudomonas mangiferae-indicae (Patel et al., 1948) and
(a)
(b)
Fig. 8.7. (a) Symptoms of bacterial black spot on the undersurface of a mango leaf,
caused by Xanthomonas axonopodis pv. mangiferaeindicae (Photograph courtesy
of O. Pruvost). (b) Symptoms of bacterial spot on the undersurface of a mango leaf,
caused by a yellow-pigmented xanthomonad (Photograph courtesy of R.C. Ploetz).
Bacterial black spot lesions are larger and more raised than those of bacterial spot.
Foliar, Floral and Soilborne Diseases 243
Erwinia mangiferae (Steyn et al., 1974) are erroneous. The pathogen’s place-
ment in Pseudomonas was probably due to its production of non-pigmented
colonies in culture (P. syringae pv. syringae causes a different disease of mango,
apical necrosis – see above). Cook (1975) indicated that E. mangiferae is a

saprophyte that reached high populations in old lesions.
Pathological, cultural, biochemical, physiological, serological and genetic
data indicate that strains of the pathogen from different production areas are
diverse (Sanders et al., 1994; Gagnevin and Pruvost, 1995, 2001; Kishun, 1995;
Pruvost et al., 2005). Genetic diversity is greatest among strains from South-
east Asia, suggesting that this region of host diversity is also a centre of
pathogen diversification (Gagnevin and Pruvost, 1995).
The pathogen has a single flagellum, is Gram-negative, rod shaped and
0.4–0.5 × 1.0–1.5 m (Manicom and Wallis, 1984). It is oxidase negative and
does not reduce nitrates to nitrites. It cannot use asparagine as a sole carbon
and nitrogen source, but is able to hydrolyse starch, esculin, gelatin and
casein. On artificial media, colonies are cream coloured. (The latter trait is
atypical for xanthomonads, which are usually yellow in culture.) Yellow-
pigmented xanthomonads have been recovered from mango in Brazil,
Réunion, South Africa and Florida USA. These strains cause non-raised leaf
lesions (Fig. 8.7b), do not cause fruit or stem lesions, and should not be
Fig. 8.8. A twig canker caused by X. axonopodis pv. mangiferaeindicae that is
associated with a wound (Photograph courtesy of O. Pruvost).
R.C. Ploetz and S. Freeman 244
classified as pv. mangiferaeindicae (Ah-You et al., 2007a). Three genetically and
pathologically distinct groups were identified from different geographic
regions and hosts in the Anacardiaceae by Ah-You et al. (2007a). Group I
strains were from the Old World, multiplied in mango and cashew (Anacar-
dium occidentale L.), fell in amplified fragment length polymorphism (AFLP)
group 9.5 of Xanthomonas axonopodis, and contained strains that produced
typical bacterial black spot symptoms on mango. These strains of Xanthomo-
nas campestris pv. mangiferaeindicae sensu lato were redescribed as X. axonopo-
dis pv. mangiferaeindicae sensu novo (s.n.) (Ah-You et al., 2007a). In contrast,
Group II strains were from Brazil, multiplied in cashew, but not mango, and
fell in AFLP group 9.6 of X. axonopodis. They are associated with symptoms
on mango that differ from those of bacterial black spot, including brown, flat
leaf lesions, and black, depressed lesions on fruit of only a few cultivars that
is sometimes associated with pulp rot. These strains were responsible for
previous reports of bacterial black spot in Brazil (Gagnevin and Pruvost,
2001; Ah-You et al., 2007a). Group III strains are responsible for a unique syn-
drome on Spondias dulcis and Spondias mombin in the French West Indies; they
fell in AFLP group 9.4. Groups II and III were described, respectively, as X.
axonopodis pv. anacardii and X. axonopodis pv. spondiae (Ah-You et al., 2007a).
Epidemiology and management
The pathogen is disseminated by wind-driven rain (Gagnevin and Pruvost,
2001). Long-distance dissemination occurs via infected propagation material
and, less frequently, in infected seed. True seed are not infected, but surface
contamination is known. Insects may play a role in dissemination, but these
interactions are little studied. The pathogen is an epiphytic colonist of leaves
(Manicom, 1986; Pruvost et al., 1990), buds (Pruvost et al., 1993) and fruit
(Pruvost and Luisetti, 1991b). Infection occurs through wounds and, less
often, stomata on old leaves (young leaves are thought to be resistant due to
their non-functional stomata) (Gagnevin and Pruvost, 2001). High RH
(> 90%) and moderate temperatures (25–30°C) favour disease development
(Kishun and Sohi, 1983; Pruvost and Luisetti, 1991b). There is a direct rela-
tionship between the level of disease that develops on leaves and fruit (Man-
icom, 1986; Pruvost et al., 1990). Pruvost and Luisetti (1991b) considered that
leaf susceptibility was an important criterion when cultivars were selected
for lower fruit susceptibility.
Resistance to bacterial black spot varies among mango cultivars, and
resistant cultivars should be used where disease pressure is high (Manicom
and Pruvost, 1994). When new orchards are established, pathogen-free plant-
ing material should be utilized. Since the pathogen moves short distances in
wind-blown aerosols (usually within orchards), the long-distance spread of
the pathogen depends almost entirely upon the movement of infected plants
(Manicom and Pruvost, 1994). Windbreaks can be used to reduce wounding
and infected twigs (Fig. 8.8) should be pruned to reduce inoculum in the
canopy.
Bacterial black spot can be quite difficult to control, particularly on sus-
ceptible cultivars. Available chemicals may be only marginally effective
Foliar, Floral and Soilborne Diseases 245
under high disease pressure (Pruvost et al., 1989). During rainy weather,
applications of Cu-based bactericides are recommended. The application
schedules for these compounds focus on protecting fruit, and vary according
to the length of time fruit are exposed to wet conditions (Manicom and Pru-
vost, 1994). Although agricultural antibiotics (e.g. various formulations of
streptomycin sulfate or nitrate) have been reported to be effective (Misra and
Prakash, 1992; Viljoen and Kotzé, 1972), resistance that develops to these
products after continuous use limits their long-term effectiveness against
this disease. Biological control measures have not been widely researched.
Pruvost and Luisetti (1991a) reported little success. In India, Kishun (1994)
indicated that a strain of Bacillus coagulans from the phylloplane of mango
was effective against strains of the pathogen, although control of bacterial
black spot in the field was not reported.
Black-banded disease
This is a relatively unimportant disease that affects mango leaves and
branches (Reddy et al., 1961). The causal fungus, Rhinocladium corticola Mas-
see (described as ‘corticolum’) (teleomorph: Peziotrichum corticolum (Massee)
Subrumanian), was described on the bark of trees in Poona, India (Hughes,
1980). It produces repent, intricately branched, septate, olivaceous hyphae
5–7 m in diameter. Erect hyphae bear globose, olivaceous, densely and
minutely tuberculate, 15–18 m conidia. Hughes (1980) questioned whether
this was a species of Rhinocladium since it was quite different from other spe-
cies in the genus. It forms a black, velvety mass of hyphae on affected sur-
faces in conspicuous blotches or bands. The fungus is restricted to the outer
portions of bark. Bordeaux mixture controls the disease, but is not required
in most situations.
Black mildew, sooty blotch and sooty mould
Several ascomycetes produce dark-coloured, usually superficial growths
on the surfaces of stems, leaves and fruit (Lim and Khoo, 1985). These
range from thin, diffuse webs of dark hyphae to opaque, felty layers; in
extreme cases, a thick crust of hyphae forms. The variations in appearance
result, presumably, from the different species of fungi that are involved.
These are usually not important problems in well-maintained orchards.
However, layers of hyphae that the black mildew and sooty mould fungi
form may be thick enough to block sunlight and inhibit photosynthesis.
These blemishes also detract from the appearance and marketability of
fruit.
Sooty moulds develop in the presence of aphids, mealybugs, scales and
other sucking insects that produce honeydew (excreta) when feeding. Hon-
eydew is a required food source for these fungi, and the problems they cause
dissipate if the associated insects are controlled. In contrast, black mildews
R.C. Ploetz and S. Freeman 246
and sooty blotch are not dependent on honeydew and grow directly on host
surfaces.
Aetiology
Black mildew and sooty mould are similar in appearance, but their respec-
tive causal agents are distinct (Lim and Khoo, 1985). The black or dark mil-
dews are a group of mostly tropical obligate plant pathogens that produce
two types of hyphopodia (Alexopoulos et al., 1996). Capitate hyphopodia are
lobed appressoria from which infection haustoria are formed, whereas
mucronate hyphopodia function as conidiogenous cells. Black mildew of
mango is caused by Meliola mangiferae Earle (Sordariomycetes, Ascomycota)
(Fig. 8.9).
In contrast, the fungi that cause sooty moulds are diverse saprophytes
that require honeydew to colonize plant surfaces. In Malaysia, Lim and Khoo
(1985) listed coelomycetes (Polychaeton), hyphomycetes (Tripospermum) and
loculoascomycetes (Antennulariella, Chaetothyrium, Limacinula and Scorias),
whereas the reported agents in India were hyphomycetes (Leptoxyphium,
(a)
(c)
(b)
10 mm
10 µm
(d)
250 µm
1 mm
Fig. 8.9. (a–c) Signs of black mildew, and (d) microscopic features of the causal
agent, Meliola mangiferae (Source: from CMI description no. 1355).
Foliar, Floral and Soilborne Diseases 247
Microxyphium and Tripospermum) and loculoascomycetes (Capnodium) (Butler
and Bisby, 1931; Prakash, 1988). In Pakistan, 18 species in eight genera were
associated with sooty mould, including the foliar pathogens Aspergillus,
Alternaria, Botryodiplodia, Capnodium, Cladosporium, Curvularia, Fusarium and
Helminthosporium spp. (Hamid and Jalaluddin, 2006). Since some of the sooty
mould fungi may not sporulate on plants and because they are often found
in combination with one another, it is usually difficult to identify the specific
species that are involved.
Although ‘sooty blotch’ has been used as a synonym for ‘sooty mould’
(Singh, 1968), sooty blotch refers specifically to disease complexes on tem-
perate and tropical plants that are not associated with honeydew and are
caused by a diverse group of Dothidiomycetes (Ascomycota) (Johnson et al.,
1997; Ploetz et al., 2000; Batzer et al., 2005). The specific agents that are associ-
ated with sooty blotch on mango are not known, but resemble those from
apple, carambola and pear (Plate 44).
Epidemiology and management
Sooty moulds develop on honeydew that is produced on plant surfaces by
aphids, mealybugs, scales and other sucking insects. They are managed by
controlling the associated insects with oils and insecticides (Lim and Khoo,
1985). In Pakistan, spraying of fungicides (sulfur (S) and mancozeb) and
insecticides (malathion, diazinon and coal tar at 1 kg/tree) separately reduced
the incidence of sooty mould on foliage, whereas a mixture of fungicides and
insecticides further decreased sooty mould incidence (Hamid and Jalalud-
din, 2006). In India, sooty mould, caused by species of Microxyphium, Lep-
toxyphium and Tripospermum, was best controlled with a spray of S and
parathion-methyl (Prakash, 1991).
Sooty blotch management has not been investigated on mango;
however, signs of these fungi have been removed from apple with various
postharvest washes (Batzer et al., 2002), and managed in apple orchards
with diverse contact and systemic fungicides (Williamson and Sutton,
2000).
Blossom blight
Blossom blight can reduce fruit set and production considerably since flow-
ers and large areas of the panicle can be killed. When this disease was con-
trolled with fungicides in the Philippines, a 55–80% increase in fruit set
occurred (Pordesimo, 1982).
Symptoms
Blossom blight starts as a wilt of the affected part of the inflorescence that is
often curved, the ‘shepherd’s crook symptom’ (Fig. 8.10). The peduncle
blackens and dies back from the tip. Internally, discoloration advances
beyond the surface lesion. Large black lesions can develop lower on the
peduncle, and once it is girdled the apex dies.
R.C. Ploetz and S. Freeman 248
Aetiology
The cause of blossom blight is confused. Colletotrichum gloeosporioides has
been reported most frequently as the responsible fungus (Fitzell et al., 1984;
Jeffries et al., 1990), and A. alternata has also been reported to attack panicles
and reduce fruit set (Cronje et al., 1990). Powdery mildew (see section below)
also damages panicles, but its symptoms are distinct from those of blossom
blight. In South Africa, symptoms caused by A. alternata and C. gloeosporioides
are small, mainly superficial black spots, 1–2 × 2–5 mm, on the peduncle
(Darvas, 1993; Lonsdale and Kotzé, 1993). Rather than blossom blight, Lonsdale
and Kotzé (1993) indicated that these pathogens caused blossom spot. In con-
trast, Lonsdale and Kotzé (1993) reported that Dothiorella mangiferae caused
extensive, systemic damage, and Darvas (1993) indicated that Dothiorella domini-
cana is the only fungus that caused typical symptoms of blossom blight.
Crous et al. (2006) placed these fungi in a new genus, Neofusicoccum (see
Decline disorders section below). Work is needed to determine the distribu-
tion of Neofusicoccum-incited panicle disease, and the identity of the most
important blossom blight pathogens worldwide.
Epidemiology and management
Little is known about the epidemiology of Neofusicoccum-incited panicle dis-
ease. Studies on the stem-end rot diseases have shown that the causal fungi
are endophytes. The roles of internal and external sources of inoculum in
the development of panicle disease are unknown. For optimal fruit set and
Fig. 8.10. Symptoms of blossom blight on panicles of mango. Note the blighted,
curved terminals and almost complete lack of fruit set (Photograph courtesy of
D. Benscher).
Foliar, Floral and Soilborne Diseases 249
development, blossom blight must be controlled. Once flowering begins,
early and frequent fungicide applications are necessary in many areas,
depending on rainfall. Previously published infection models can be used to
time applications appropriately (Fitzell et al., 1984; Dodd et al., 1991). Fitzell
et al. (1984) investigated environmental conditions that were conducive to
infection by C. gloeosporioides, and indicated that temperature and free mois-
ture were important determinants of infection. They developed a model for
scheduling fungicide application, which reduced the number of applications
that were needed to control blossom blight. Presumably, systemic fungicides
would be needed to control disease caused by endophytic agents.
Decline disorders
Several diseases of mango have been variously termed blight, canker, decline,
gummosis, twig blight, tip dieback and stem bleeding. They have similar
symptoms and aetiologies.
Symptoms
These widespread problems are not well understood. Symptoms include: (i)
marginal scorching of leaf lamina; (ii) foliar symptoms of nutritional defi-
ciencies, particularly of iron (Fe) and manganese (Mn); (iii) vascular discolor-
ation (Fig. 8.11a); (iv) dieback of small branches basipetally from the terminal
that may or may not progress to defoliation (Fig. 8.11b and c); (v) gummosis,
an oozing of a clear or cloudy exudate either from terminal buds or from
branches, scaffold limbs or trunks (Plate 45); and (vi) root degeneration (Lim
and Khoo, 1985; Ploetz et al., 1996a; Ploetz and Prakash, 1997).
Aetiology
Diverse biotic and abiotic factors may be primary causes of decline symp-
toms or predisposing agents (McSorley et al., 1980; Kadman and Gazit, 1984;
Schaffer et al., 1988; Ploetz and Prakash, 1997). Fungi are the most common
agents. They are endophytes that also cause stem-end rots on mango fruit,
and are usually secondary pathogens that cause disease on weakened, pre-
disposed hosts (Johnson et al., 1992; Ploetz and Prakash, 1997; Slippers et al.,
2005; Slippers and Wingfield, 2007). Several species cause all or some of the
above symptoms when used to individually inoculate plants (Ploetz et al.,
1996a). Their frequent association with one another in affected tissues may
indicate that these symptoms usually develop, or develop more severely,
after multiple infections.
Several of these pathogens are in the Botryosphaeriaceae (Dothidiomy-
cetes, Ascomycota). The taxonomy and nomenclature of these fungi has been
confused, and ‘phylogenetic understanding of major groups within Botry-
osphaeria remains poor’ (Crous et al., 2006). With 28S rDNA sequence data,
Crous et al. (2006) examined natural relationships among available members
of the family. Ten lineages were distinguished, most of which contained ana-
morphs with distinct morphological features. New relationships were revealed
R.C. Ploetz and S. Freeman 250
in some of the lineages that necessitated the renaming of several taxa (Crous
et al., 2006). These new names and holomorphs are used below when discuss-
ing the taxa that occur on mango.
Lasiodiplodia theobromae (Pat.) Griffon and Maubl.) (synonyms: Botryodi-
plodia theobromae Pat., Diplodia natalensis Pole-Evans, and Diplodia theobromae
(Pat.) W. Nowell) is the most common and widespread cause of decline dis-
eases of mango (Ploetz and Prakash, 1997), and affects many other host plants
in the tropics (Punithalingam, 1976). Crous and Palm (1999) declared B. theo-
bromae, a nomen dubium. Denman et al. (2000) reduced D. natalensis and L.
theobromae to synonymy with D. theobromae. However, adopting this change
was questioned by Burgess et al. (2006), who noted that five species of Lasio-
diplodia fell in a phylogenetic clade that had 100% bootstrap support; it was
distinct from a clade that included species of Diplodia and Dothiorella. The
teleomorph of L. theobromae, formerly Botryosphaeria rhodina (Cooke) Arx
(synonym: Physalospora rhodina Cooke), is usually not found in nature. In the
study of Crous et al. (2006), the genus Botryosphaeria was reserved for the type
species Botryosphaeria dothidea (Moug.:Fr.) (anamorph: Fusicoccum aesculi
Corda), which was in a different clade than L. theobromae. However, Crous et
al. (2006) refrained from renaming B. rhodina until the poorly resolved clade
in which it resided could be clarified with work with additional isolates and
analyses.
Lasiodiplodia theobromae attacks weakened trees that are predisposed by:
high and low temperatures; drought; high RH; hardpan soils; sun scorch;
and tar and tanglefoot (Muller, 1940; Das Gupta and Zacchariah, 1945;
Alvarez-García and López-Gracía, 1971; Acuña and Waite, 1977; Ploetz et al.,
1996a). It is often an endophyte, infects wounded plants, and is found in soil,
on dead twigs, mummified fruit and on organic debris beneath trees (Johnson
et al., 1992).
(a) (b) (c)
Fig. 8.11. Among the symptoms that are associated with mango decline are:
(a) internal/vascular discoloration and branch terminal death (tip dieback) that may not
(b), or may be associated with defoliation (c) (Photographs courtesy of D. Benscher).
(a) (b) (c)
Foliar, Floral and Soilborne Diseases 251
Dieback caused by L. theobromae has been recognized as a significant dis-
ease in India since the 1940s. It was the most serious disease of mango in the
Jaipur district (Verma and Singh, 1970), and affected 30–40% of the planta-
tions in the Moradabad region of Uttar Pradesh (Prakash and Srivastava,
1987). Das Gupta and Zacchariah (1945) indicated that only L. theobromae
caused dieback; Phoma sp. and two Fusariumspp. were not pathogenic. Lasio-
diplodia theobromae caused a canker on mango in Indonesia (Muller, 1940) and
Malaysia (Lim and Khoo, 1985), dieback in Egypt and the Sonsonate area of
El Salvador (Acuña and Waite, 1977), and gummosis and dieback in Puerto
Rico (Alvarez-García and López-Gracía, 1971).
Lasiodiplodia theobromae produces fluffy, grey-black mycelium on oatmeal
agar (OA) and PDA (Johnson, 1994b). Conidiomata may be simple or develop
into aggregated stromatic bodies (Burgess et al., 2006). Cirri of conidia may
ooze from ostioles. Conidia are initially hyaline, aseptate, granular, ovoid
to ellipsoid and thick-walled (Fig. 8.12). Mature conidia are two celled,
17–33 × 10–15 m, and brown walled with numerous longitudinal striations.
Paraphyses are usually present. The teleomorph was produced when indi-
vidual isolates were cultured on caimito fruits (Chrysophyllum cainito L.) or
papaya (Carica papaya L.) stems, suggesting that it was homothallic (Alvarez-
García and López-Gracía, 1971).
One of the most important mango pathogens causes stem-end rot on
fruit, dieback and blossom blight. Crous et al. (2006) refer to it as Neofusicoccum
parvum (Pennycook and Samuels) Crous, Slippers and A.J.L. Phillips (formerly
Fusicoccum parvum Pennycook and Samuels) (teleomorph: Botryosphaeria-like;
formerly Botryosphaeria parva Pennycook and Samuels). Slippers et al. (2005)
argued that D. dominicana Petro and Cif. may be synonymous with this fun-
gus. Johnson (1992), an author of the Slippers et al. (2005) paper, had placed
D. dominicana in synonymy with F. aesculi Corda (teleomorph B. dothidea
(Moug.:Fr.) Ces. and De Not.). They indicated that this fungus had been mis-
identified as Botryosphaeria ribis Gross. and Duggar (anamorph: Fusicoccum
sp.) and B. dothidea (anamorph: F. aesculi), due to overlapping host ranges
and spore dimensions. They felt that the tip dieback fungus reported by
Ramos et al. (1991) as B. ribis was probably N. parvum (= ‘B. parva’).
Neofusicoccum parvum produces cottony grey mycelium and discrete pyc-
nidia or stromatic multilocular fruiting bodies on, respectively, PDA and
OA (Johnson et al., 1991). Discrete, immersed pycnidia in a subcuticular
pseudostroma are produced on mango. Conidia are fusiform to navicular,
14.7–25.5 (19) × 4.5–7 (5.2) m, hyaline and unicellular (Slippers et al., 2005).
Sometimes, brown, biseptate conidia are observed. The teleomorph develops
infrequently on OA, and has been found on mango twigs in tree litter in
Australia (Johnson et al., 1991). On twigs, pseudothecia are subglobose to
pyriform, 210 × 120 m, and form beneath the epidermis. Ascostromata are
hemi-lenticular and up to 10 mm wide on OA. Asci are eight spored, bituni-
cate and irregularly biseriate. Ascospores are hyaline, single celled, fusiform
and 16–25 × 4.5–9.5 m.
Neofusicoccum mangiferum (Syd. and P. Syd.) Crous, Slippers and A.J.L.
Phillips (basionym: Dothiorella mangiferae Syd. and P. Syd.; synonyms:
R.C. Ploetz and S. Freeman 252
Nattrassia mangiferae (Syd. and P. Syd.) B. Sutton and Dyko; Fusicoccum
mangiferum(Syd. and P. Syd.) Johnson, Slippers and M.J. Wingf.) causes blos-
som blight and postharvest fruit rot in South Africa (Lonsdale and Kotzé,
1993; Saaiman, 1996). On PDA, N. mangiferum produces grey, felted myce-
lium with gregarious, partly immersed, discrete conidiomata, ‘pepper-spot’
patterns of pycnidial initials, and dark grey mycelium that lacks the white
tufts found in similar species (i.e. N. parvum) (Johnson, 1994b; Slippers et al.,
2005). On mango, the fungus produces unilocular conidiomata in subcuticu-
lar pseudostroma. The conidia of N. mangiferum differ from those produced
by other Neofusicoccum spp. by their shorter average length (13–14 m) and
smaller length/width ratio (2–2.5) (Slippers et al., 2005). They are usually
unicellular, ellipsoid to ovoid, 13.6 × 5.4 m and hyaline, although conidia
often become one or two septate, light brown with distinctly darker middle
cells. The teleomorph (not identified) resembles ‘B. parva’, and develops
occasionally on OA.
(b)
(d)
(c)
(a)
500 µ
10 µ
Fig. 8.12. Pycnidia (a and b), conidiogenous cells and immature conidia (c) and
mature and immature conidia (d) of Lasiodiplodia theobromae (Source: from CMI
description no. 519).
Foliar, Floral and Soilborne Diseases 253
A dieback disease of mango has been recognized in Niger (Reckhaus
and Adamou, 1987). Neoscytalidium dimidiatum (Penz.) Crous and Slippers
(basionym: Torula dimidiata Penz.; synonyms: Scytalidium dimidiatum (Penz.)
B. Sutton and Dyko (Fig. 8.13); Fusicoccum dimidiatum (Penz.) D.F. Farr;
Hendersonula toruloidea Natrass) causes sudden wilting of shoots to large
branches, firing of leaves and trunk cankers from which a clear exudate orig-
inates. Reckhaus and Adamou (1987) believed that water stress was a pri-
mary, predisposing factor in the development of this disease.
Botryosphaeria dothidea (anamorph: F. aesculi) is an uncommon mango
pathogen (Ploetz and Prakash, 1997; Slippers et al., 2005). It produces a fluffy
grey mycelium with discrete pycnidia on PDA or stromatic multilocular
fruiting bodies on OA. Discrete, immersed pycnidia are produced on mango.
(e)
(d)
(c)
(a)
(b)
10 µ
50 µ
Fig. 8.13. (a) Vertical section of stroma, (b) part of pycnidial wall and conidiophores,
(c) conidiophores, (d) conidia, and (e) the Scytalidium-like synanamorph of Neoscyta-
lidium dimidiatum (Source: from CMI description no. 274).
R.C. Ploetz and S. Freeman 254
Conidia are 18.8–30.4 × 4.5–7 m, hyaline, and single celled (Fig. 8.14). The
teleomorph is occasionally produced on OA and has been found in litter
beneath avocado and mango trees (Johnson, 1994b; Michailides et al., 1999).
On twigs, pseudothecia are subglobose to pyriform, 210 × 120 m, and im-
mersed beneath the epidermis. On OA, ascostromata are hemi-lenticular and
up to 10 mm wide. Asci are eight spored, bitunicate and irregularly biseriate.
Ascospores are hyaline, single celled, fusiform and 16–25 × 4.5–9.5 m.
Mango decline is an important disorder in Florida USA, Israel and
other areas that have calcareous soils (Schaffer, 1994; Ploetz et al., 1996a).
Symptoms include interveinal chlorosis and marginal necrosis of leaves,
(a)
(c)
(b)
10 µ
500 µ
Fig. 8.14. (a) Conidia, (b) conidiophores and (c) a vertical section of a conidioma of
Fusicoccum aesculi, anamorph of Botryosphaeria dothidea (Source: Sutton, 1980).
Foliar, Floral and Soilborne Diseases 255
dieback of young twigs that progresses to larger branches, reduced growth of
secondary roots, gummosis and vascular discoloration. Several different
factors have been associated with mango decline in Florida. Schaffer et al.
(1988) used the Diagnosis and Recommendation Integrated System (DRIS) to
assess the nutritional status of declining and healthy ‘Tommy Atkins’ trees.
The nutrient imbalance index, an indication of the overall nutrient balance in
trees, was greatest for declining trees. DRIS identified Mn, Fe or both ele-
ments as the most deficient in declining trees, and in two of the three declin-
ing orchards that they investigated, concentrations of these elements were
below the critical range. Mineral deficiencies may be predisposing factors in
the development of mango decline, since pathogenic fungi are recovered
from symptomatic trees (see below).
McSorley et al. (1980) detected the nematode Hemicriconemoides mangiferae
Siddiqi at low, but consistent levels on declining mango trees. They sug-
gested that it may have been responsible for the reduced root growth in
affected trees, and could play a role in the development of the disorder.
Smith and Scudder (1951) reported that Diplodia sp. caused a dieback
of mango. Additional species of fungi were examined by Ploetz et al.
(1996a). Alternaria alternata, C. gloeosporioides, N. parvum (D. dominicana), L.
theobromae and two Phomopsis spp. were recovered from trees with diverse
decline symptoms, and caused one or more of these symptoms on ‘Keitt’ and
‘Tommy Atkins’. Colletotrichum gloeosporioides, N. parvum and L. theobromae
were most damaging, and caused significant bud necrosis, tip dieback,
gummosis and vascular discoloration (Fig. 8.15); these symptoms were
distinguishable only when C. gloeosporioides sporulated on inoculated
branches.
In summary, several different fungi cause, or are associated with, decline
symptoms worldwide; most are endophytes that have Botryosphaeria or
Botryosphaeria-like teleomorphs (Botryosphaeriaceae). Stress and wound pre-
disposition are usually associated with symptom development.
Management
Controlling decline disorders of mango is difficult. Techniques that could
detect these pathogens in plants would be useful to identify pathogen-free
propagation materials. The internal location and the diversity of fungi that
are involved decrease the opportunities for controlling these disorders with
fungicides (Peterson et al., 1991). Because significant movement of some of
these pathogens may occur via wind and rainsplash, strategic applications of
broad-spectrum protectant fungicides may be effective at certain times of the
year (Lonsdale and Kotzé, 1993), but have not been tested. In India, dieback
was managed by pruning affected portions of the canopy and treating the
wounded areas with a 5:5:50 Bordeaux mix (Prakash and Raoof, 1989). Man-
agement of the controllable predisposing factors, such as drought stress, may
also be beneficial. A better understanding of the epidemiology of these dis-
eases would assist these efforts. Pruning to force synchronous flushes of
foliar growth might enable the avoidance of windows of infection for certain
pathogens (Johnson, 1994b).
R.C. Ploetz and S. Freeman 256
Galls and scaly bark
Gall and scaly bark disorders of mango are known in several producing
regions. These diseases are usually minor problems.
Symptoms
In India, bark scaling develops as deep cracks along the entire rootstock por-
tion of the plant, and cracks may penetrate the phloem and become necrotic
(Prakash and Srivastava, 1987). These symptoms resemble those of a scaly
bark disorder, ‘cuarteado’, in Colombia (Cook, 1975). In Hawaii, similar
symptoms developed on mango seedlings (Cook et al., 1971). The bark from
the soil line to the first branches was rough and scaly, and xylem pegs, 5 mm
long, were evident when the bark was removed around leaf scars and
secondary branches.
In Mexico, a disorder known as ‘nanahuate’, ‘bolas’ or ‘buba of mango’,
causes galls, 5–10 cm in diameter, which resemble a cauliflower, are initially
light green, but become dark brown as they die (Fig. 8.16) (Angulo and
Villapudua, 1982). The galls remain attached to trees for many years, and
severely affected branches die. Similar symptoms are found in Florida USA,
and are associated with pruning injuries. Larger galls have also been noted
in Puerto Rico, as well as the US Department of Agriculture (USDA)
Agriculture Research Service (ARS) in Miami and University of Florida
in Homestead (Fig. 8.17) (Ploetz et al., 1996b; R. Rodriguez, personal
communication). Some of the latter galls are large, have rough, scaly exteri-
ors, and are usually found on the main trunk or scaffold limbs of affected
trees. Cracks may penetrate the phloem and become necrotic, but the branch
death that is associated with galls in Mexico and India has not been
observed.
(a) (b) (c)
Fig. 8.15. Decline symptoms induced on ‘Tommy Atkins’ plants artificially inoculated
with isolates of: (a) C. gloeosporioides; (b) Neofusicoccum parvum; and (c) L. theobro-
mae (Photographs courtesy of D. Benscher).
(a) (b) (c)
Foliar, Floral and Soilborne Diseases 257
Aetiology
Fusarium decemcellulare C. Brick (synonym: Fusarium rigidiuscula (Brick) Snyd.
and Hans.) causes these diseases in Florida USA, Mexico and Venezuela
(Malaguti and de Reyes, 1964; Angulo and Villapudua, 1982; Ploetz et al.,
1996b). Colonies on PDA are dark carmine-red on the underside. The fungus
produces microconidia in false heads or chains on branched and non-
branched monophialides (Fig. 8.18). Large macroconidia, 92–55 × 7–5.5 m,
are produced in slimy yellow sporodochia, c.1 mm in diameter. The fungus’s
Fig. 8.16. Galls of the ‘buba’ type in Haiti (Photograph courtesy of Carolyn Cohen,
USDA, Animal and Plant Health Inspection Service (APHIS)).
Fig. 8.17. Large, 30-year-old gall on ‘Langra’ in the USDA-ARS mango germplasm
repository in Miami (Photograph courtesy of R.C. Ploetz).
R.C. Ploetz and S. Freeman 258
teleomorph, Albonectria rigidiuscula (synonyms: Nectria rigidiuscula Berk. and
Broome, and Calonectria rigidiuscula) (Rossman et al., 1998), has not been
observed on mango.
Fusarium decemcellulare causes corky bark, gall, canker and dieback dis-
eases on diverse woody hosts in the subtropics and tropics (Holliday, 1980;
Farr et al., 1989; Alfieri et al., 1994). It causes an important disease of cacao
(Theobroma cacao L.), cushion gall, as well as a stem gall on loquat (Eriobotrya
japonica (Thunb.) Lindl.) and scaly bark of pongam (Pongamia pinnata (L.)
Pierre). Host specialization in the fungus has not been reported. Fusarium
decemcellulare has not been reported to cause gall and scaly bark disorders of
mango in other areas. The possible role of Agrobacterium tumefaciens was
examined in Miami; however, the bacterium could not be recovered from
affected tissues (R. McGuire, Miami, 1993, personal communication). In
Hawaii, microorganisms were not recovered from affected plants (Cook,
1975).
Epidemiology
Isolates of F. decemcellulare from mango are only mildly aggressive (Ploetz
et al., 1996b), and require wounding in order to infect. A recent outbreak of
scaly bark in a commercial mango orchard in Florida USA was associated
with pruning wounds. In the cushion gall disease on cacao, F. decemcellulare
interacts with several different insect pests and pathogenic agents (Holliday,
(b)
(c)
(a)
20 µ
Fig. 8.18. (a) Ascus and ascospores of Albonectria rigidiuscula, and (b) micro-
conidia and conidiophores, and (c) macroconidia and conidiophores of its anamorph,
Fusarium decemcellulare (Source: from CMI description no. 21).
Foliar, Floral and Soilborne Diseases 259
1980; Ploetz, 2007). These insects may facilitate infection and disseminate the
pathogen. Insect-F. decemcellulare interactions have not been investigated on
mango.
Management
No pesticides have been identified to control this problem. Measures that
should be helpful include the removal and destruction of affected branches
and trees in the orchard, disinfestation of pruning equipment to ensure that
the pathogen is not spread during pruning operations, and the use of healthy
planting material in new orchards.
Grey leafspot
Pestalotiopsis mangiferae (Henn.) Steyaert (synonym: Pestalolia mangiferae
Henn.; no teleomorph of the fungus is known) causes grey leafspot and stem-
end rot of mango fruit (Lim and Khoo, 1985; Johnson, 1994b). It is a weak
pathogen that usually requires wounding in order to infect mango. Grey
leafspot is usually unimportant and occurs mainly on unhealthy or poorly
maintained trees.
Pestalotiopsis mangiferae produces abundant conidia in acervuli that
develop in grey leafspot lesions and necrotic areas on fruit (Lim and Khoo,
1985). As lesions age, black columns of spores emanate through ruptures in
the host epidermis. Conidia are produced that have three thick-walled,
brownish, concolorous median cells and thin-walled, hyaline apical and
basal cells; the apical cells bear three characteristic appendages (Fig. 8.19).
Conidia are 20 × 5 m, fusiform and straight to slightly curved. Two other
species of Pestalotiopsis that occur on mango produce larger conidia, Pestalo-
tiopsis mangifolia Guba and Pestalotiopsis versicolor Speg. (synonyms: Pestaloti-
opsis cliftoniae Tracy and Earle and Pestalotiopsis coccolobae Ellis and Everh.).
On leaves, symptoms are light grey spots, usually 2–20 mm in diameter
(Lim and Khoo, 1985). These may coalesce to form large patches of necrotic
tissue on leaves. Lesions are surrounded by dark, raised margins, and as they
mature, raised black dots (which are columns of the pathogen’s conidia) are
evident in lesion centres. Although most cultivars are susceptible, specific
control measures are usually not required. Dithiocarbamate fungicides con-
trol this disease.
Leaf blight
This disease has been reported in India and Nigeria (Hingorani et al., 1960;
Cook, 1975; Okigbo, 2001; Okigbo and Osuinde, 2003), and the causal fun-
gus, Macrophoma mangiferae Hingorani and Sharma (Ascomycota), has also
been intercepted in shipments to the USA from Mexico (Systematic Mycol-
ogy and Microbiology Laboratory, USDA-ARS, Beltsville). This is not a
serious problem.
R.C. Ploetz and S. Freeman 260
Macrophoma mangiferae produces subepidermal, globose pycnidia, 77–231
m in diameter. Hyaline conidiophores, 1.5–2 × 8–11 m, produce unicellular
conidia, 5.3–7 × 10.5–24.5 m. No teleomorph is known. Since the genus Mac-
rophoma has been placed in synonymy with Sphaeropsis, this species should
be redescribed.
Leaves, twigs and fruit are affected, especially when the latter are stored.
Small, yellow spots gradually enlarge to become brown with wide purple
margins. The lesions are initially circular, but develop an irregular appear-
ance and may encompass large portions of the leaf surface. Pycnidia form
most frequently on the underside of leaves. Elliptical stem lesions are infre-
quent but can girdle stems. In India, the disease is most serious during the
rainy season (Verma and Singh, 1996b). Macrophoma mangiferae survives in
pycnidia that develop on bark of twigs of young mango plants, blighted
leaves and as dormant mycelium in wood (Verma and Singh, 1996a). Okigbo
(2001) reported that the fungus survived adverse conditions best in stems
and branches.
Four applications of captan, Bordeaux mixture, captafol, carbendazim
and thiophanate-methyl were effective on young plants (Verma and Singh,
1996a). The bacterium Bacillus subtilis NCIB 3610, isolated from soil under a
mango tree, inhibited M. mangiferae on agar plates, and symptoms were
reduced in the field by the application of the antagonist (Okigbo and Osuinde,
2003).
(a)
(b)
50 µm
(c)
10 µm
Fig. 8.19. (a) Vertical section of an acervulus, (b) mature conidia, and (c) conidiog-
enous cells and developing conidia of Pestalotiopsis mangiferae (Source: from CMI
description no. 676).
Foliar, Floral and Soilborne Diseases 261
Malformation
Malformation is one of the most destructive mango diseases (Ploetz, 2001).
Although trees are not killed, the vegetative phase of the disease impedes
canopy development and the floral phase dramatically reduces fruit yield.
Based on the incidence and severity of malformation in Egypt, an estimated
US$15 million of fruit were lost in 1998 (Ploetz et al., 2002). Losses in more
important producing countries (i.e. India) are undoubtedly much greater.
Malformation was first reported in India in 1891 (Kumar and Beniwal,
1991), and has subsequently been observed in Brazil, Myanmar, Egypt, El
Salvador, India, Israel, Malaysia, Mexico, Nicaragua, Oman, Pakistan, South
Africa, Sudan, Spain, Swaziland, Uganda and the USA (Flechtmann et al.,
1973; Crookes and Rijkenberg, 1985; Lim and Khoo, 1985; Kumar and Beni-
wal, 1991; Ploetz, 2001; Kvas et al., 2007; S. Freeman, Bet Dagan, 2007, unpub-
lished data; G.I. Johnson, Jamison, Australia, 2007, personal communication;
J.F. Leslie, Manhattan, Kansas, 2007, personal communication). Since the
pathogen is easily disseminated in infected germplasm and there are con-
spicuous gaps in the disjunct distribution (note African and American records),
malformation is probably more widely spread.
Symptoms
Malformation affects vegetative and floral meristematic tissues (Fig. 8.20)
(Ploetz, 2001). Vegetative malformation is most serious on seedlings and
small plants in nurseries, especially where seedlings are grown beneath
affected trees, a common practice in the Middle East (Ploetz et al., 2002;
Youssef et al., 2007). Vegetative malformation also occurs on mature trees.
Apical and axillary buds produce misshapen shoots with shortened inter-
nodes and dwarfed leaves that are brittle and recurve towards the sup-
porting stem (Fig. 8.20). Shoots may not expand fully, resulting in a bunched
appearance (i.e. the ‘bunchy-top’ symptom of the disease).
Floral malformation is most important. Affected inflorescences usually
do not set fruit or fruit are aborted. Primary or secondary axes on affected
panicles are shortened, thickened and highly branched (Fig. 8.20). Malformed
panicles produce up to three times the normal number of flowers, range from
one-half to two times the normal size, and have an increased proportion of
male to perfect flowers. Malformed panicles may also produce dwarfed and
distorted leaves (exhibit phyllody).
Aetiology
The aetiology of malformation has been controversial for almost as long as
the disease has been recognized (Ploetz, 2001). Suggested causes include
mites (Narasimhan, 1954), nutritional problems (Prasad et al., 1965), physio-
logical or hormonal imbalances (Dang and Daulta, 1982; Singh and Dhillon,
1989), viruses (Kauser, 1959) and unknown causes (Kumar and Beniwal,
1991). Summanwar et al. (1966) demonstrated that a fungus, identified then
as Fusarium moniliforme Sheld., was responsible for the floral phase of this
disease. Varma et al. (1974) later showed that F. moniliforme also caused
R.C. Ploetz and S. Freeman 262
(a)
(b)
Fig. 8.20. Among the symptoms caused by malformation are: (a) in panicles, an in-
crease in the size and number of flowers and interspersed floral and vegetative organs
(phylody); and (b) in vegetative shoots, compact or retarded growth of buds and brittle,
dwarfed and recurved leaves. Symptoms in (a) are on ‘Haden’ in Michoacan, Mexico
and are associated with an undescribed species in the Gibberella fujikuroi species
complex, whereas (b) is on a ‘Van Dyke’ plant artificially inoculated with an isolate of
Fusarium mangiferae (Photographs courtesy of R.C. Ploetz).
(a)
(b)
Foliar, Floral and Soilborne Diseases 263
vegetative malformation. This pathogen has had several synonyms in
the literature, including: Fusarium subglutinans (Wollenweb. and Reinking)
Nelson, Toussoun and Marasas; F. moniliforme Sheldon var. subglutinans
Wollenweb. and Reinking; and F. moniliforme Sheldon emend. Snyd and
Hans. ‘Subglutinans’ sensu Snyd., Hans. and Oswald.
In 2002, 29 strains of the pathogen from Egypt, Florida USA, Israel,
Malaysia and South Africa were redescribed as a new species in the Gibberella
fujikuroi species complex (GFSC), Fusarium mangiferae Britz, Wingfield and
Marasas (Steenkamp et al., 2000; Britz et al., 2002). Fusarium mangiferae resem-
bles morphologically other members of the GFSC. It was established based
on -tubulin and histone H3 DNA sequences, subtle morphological differ-
ences, and because most of the examined strains had been shown in previous
studies to cause malformation on artificially inoculated mango. The presence
of F. mangiferae has been verified in India (O’Donnell et al., 1998; Zheng and
Ploetz, 2002), Oman (Kvas et al., 2007) and Spain (S. Freeman, Bet Dagan,
2007, unpublished results). Although a recent report from Pakistan mentions
F. mangiferae, the identity of the pathogen there is not clear since the authors
only discussed morphological characteristics of the pathogen (Iqbal et al.,
2006).
Based on DNA sequence data (O’Donnell et al., 1998, 2000; Steenkamp
et al., 1999, 2000), F. mangiferae is related to a lineage that includes Fusarium
fujikuroi Nirenberg, Fusarium proliferatum (Matsushima) Nirenberg, and
Fusarium sacchari (E. J. Butler) W. Gams (Marasas et al., 2006), and corresponds
to the ‘Asian Clade’ of O’Donnell et al. (1998). Based on combined sequence
data for five genes, the closest known relative of F. mangiferae is an isolate
from tropical rainforest soil in Papua New Guinea (Marasas et al., 2006).
Fusarium mangiferae produces white, floccose mycelium on PDA with
light- to dark-purple pigments in the agar (Leslie and Summerell, 2006).
Cream-coloured sporodochia on carnation leaf agar (CLA) produce abun-
dant thin-walled, long, slender and straight to slightly curved, three- to five-
septate macroconidia with curved apical cells and foot-shaped basal cells
(Fig. 8.21). Single celled, rarely single septate, obovoid microconidia are
produced in false heads on polyphialides with two to five conidiogenous
openings and on monophialides. Microconidial chains and chlamydospores
are absent.
A second species, Fusarium sterilihyphosum Britz, Wingfield and Marasas,
was described originally for isolates from a small area in South Africa (Britz
et al., 2002). In subsequent work, it was detected in Brazil (Ploetz, 2003; K.
O’Donnell, unpublished results), and was recently shown to cause malfor-
mation in Brazil after artificial inoculation (Lima et al., 2006b). On PDA, colo-
nies of F. sterilihyphosum produce white, floccose mycelium with rose to light
purple pigmentation in the agar (Leslie and Summerell, 2006). Uncommon,
cream- to orange-coloured sporodochia are produced on CLA that produce
rare, long, slender, three to five-septate macroconidia (Fig. 8.22). On mono-
and polyphialides, obovoid, oval to allantoid microconidia that are usually
single celled are produced on false heads. Distinctive sterile coiled hyphae
are produced by some isolates of this species.
R.C. Ploetz and S. Freeman 264
Fusarium sterilihyphosum is relatively uncommon in South Africa and
Brazil where, respectively, F. mangiferae and a third, unnamed species that
resembles F. sterilihyphosum morphologically, predominate. The latter taxon
is phylogenetically distinct from F. mangiferae and F. sterilihyphosum, produces
a unique teleomorph in the GFSC, and has been shown to cause malforma-
tion (Lima et al., 2006a, b).
Fusarium mangiferae has not been found in either Brazil (Lima et al., 2006a,
b) or Mexico (Rodríguez-Alvarado et al., 2006, 2008). In the latter studies,
isolates that were recovered from malformed trees resembled F. sterilihypho-
sum in that they induced malformation symptoms, formed sterile coiled
hyphae and produced a PCR fragment that is also produced by isolates of F.
sterilihyphosum (see below). Translation elongation factor-1 DNA sequences
for isolates from several areas in Mexico are identical, but differ signifi-
cantly from other taxa in the GFSC; they probably represent a new species
(Rodríguez-Alvarado et al., 2006, 2008; K. O’Donnell, Peoria, 2007, personal
communication). Additional work is needed to clarify relationships among the
strains in Brazil and Mexico, and whether they are found elsewhere in the Amer-
icas. Likewise, whether F. mangiferae is found outside Florida USA in the western
hemisphere should be determined; it predominates in the eastern hemisphere.
PCR primer pairs have been used to diagnose some of the above taxa.
Zheng and Ploetz (2002) developed a pair, 1-3F/R, that amplifies a 608 bp
fragment for F. mangiferae. It has been used extensively for diagnostic pur-
poses (Youssef et al., 2007). Another pair, 61-2F/R, developed to diagnose
Fusarium verticilloides (published as F. moniliforme in Müller et al., 1999), did
not amplify F. mangiferae DNA, but when amplification protocols were mod-
ified, amplified a 445 bp-fragment for strains of F. sterilihyphosum and the
new species from Mexico (Zheng and Ploetz, 2002; Rodríguez-Alvarado et al.,
2008). It has not been tested with the unnamed pathogen from Brazil.
(a)
(b)
(c)
(d)
(e)
(f )
Fig. 8.21. Microscopic features of Fusarium mangiferae: (a) and (b), macroconidia;
(c) and (d), microconidia, and (e) and (f), microconidia in situ on carnation leaf agar.
Scale bars for (a)–(d) = 25 m, and (e)–(f) = 50 m (Photographs courtesy of Suzanne
Bullock).
Foliar, Floral and Soilborne Diseases 265
Three other taxa have been associated with mango malformation. Fusar-
ium oxysporum Schlecht emend. Snyder and Hansen (Fig. 8.23) was reported
in Egypt and Mexico (Bhatnagar and Beniwal, 1977; Kumar and Beniwal,
1991), but these reports appear to be erroneous since bona fide, vouchered
specimens have neither been described nor shown to cause the disease (Plo-
etz, 2001; Rodríguez-Alvarado et al., 2008). Fusariumsp. nov. (Britz et al., 2002)
and F. proliferatum (Gibberella intermedia (Kuhlman) Samuels, Nirenberg and
Seifert) were recovered from malformed mango trees in Malaysia (Leslie,
1995), but their pathogenicity has not been determined.
Epidemiology
Although malformation has been reproduced with F. sterilihyphosum and the
unnamed taxa from Brazil and Mexico, no work has been conducted on the
epidemiology of disease that is caused by these pathogens. Thus, results
below are from work on F. mangiferae or what is presumed to be this species.
Fusarium mangiferae is spread by grafting and in infected nursery stock
(Prakash and Srivastava, 1987). Since seed do not appear to harbour the
fungus (Saeed and Schlosser, 1972; Youssef et al., 2007), seedlings should be
disease free. Microconidia of F. mangiferae are probable infective propagules
since they are the primary spores that are produced by the fungus and form
profusely on dead malformed tissues. The disease spreads slowly in orchards,
perhaps because conidia of the pathogen die quickly when exposed to sun-
light (Manicom, 1989). Experimentally, populations of F. mangiferae in infected
panicles in Egypt and Israel declined rapidly during the summer (Youssef
et al., 2007). Wounding enhances infection and subsequent disease develop-
ment (Ploetz, 2001).
(a) (c) (e)
(b) (d) (f )
(g)
(h)
Fig. 8.22. Microscopic features of Fusarium sterilihyphosum: (a) and (b), macroconidia; (c) and
(d), microconidia; (e) and (f), coiled hyphae; and (g) and (h), microconidia in situ on carnation
leaf agar. Scale bars for (a)–(d) = 25 m and (e)–(f) = 50 m (Photographs courtesy of Suzanne
Bullock).
R.C. Ploetz and S. Freeman 266
The mango bud mite, Aceria (Eriophyes) mangiferae Sayed, is often
observed in high numbers on malformed trees and has been indicted as the
cause, or a factor in the development, of this disease (Narasimhan, 1954,
1959; Nariani and Seth, 1962). Circumstantial evidence indicates that the mite
does not cause malformation (Ploetz and Prakash, 1997); for example, A.
mangiferae is present in Australia, but the disease is not (Ridgeway, 1989).
However, A. mangiferae probably vectors the pathogen. It has been recovered
from the mite’s body on culture media (Crookes and Rijkenberg, 1985; Sattar,
Ismailia, 2006, personal communication), and was recently shown to adhere
to its body (Gamliel-Atinsky, Freeman and Palevsky, unpublished data) (Fig.
8.24). The mite cannot ingest the pathogen, due to its small mouthparts.
However, it was able to move spores of F. mangiferae to infection courts in
mango buds via external contamination of its body, and increased infection
Fig. 8.23. Microscopic features of Fusarium oxysporum: (a) sporodochia, (b) macro-
conidia, (c) microconidia in false head on monophialide, (d) terminal and intercalary
chlamydospores, and (e) macroconidia and microconidia (Photographs courtesy of
K. O’Donnell).
(a) (b)
(c)
(d) (e)
Foliar, Floral and Soilborne Diseases 267
of buds by the pathogen (Gamliel-Atinsky et al., 2007). Presumably, the mite’s
feeding on buds facilitated infection. Aceria mangiferae does not appear to
play a significant role in disseminating the pathogen among trees. In Israel,
mites were not found in traps that were designed to monitor their movement
in a malformed mango orchard, although high numbers of F. mangiferae
conidia were recorded in the air in this orchard (Gamliel-Atinsky et al.,
2007).
The distribution of F. mangiferae in affected trees suggests that vegetative
and floral buds are the primary sites of infection and that systemic coloniza-
tion of older, subtending tissues does not occur. Freeman et al. (1999) trans-
formed isolates of F. mangiferae from mango with the uidA reporter gene
(-glucuronidase), and used them to artificially inoculate mango. Their results
verified that bud and flower tissues of the host are primary infection sites,
and that wounds provide points of entry for the pathogen. In Florida USA, F.
mangiferae was restricted almost entirely to malformed floral and vegetative
tissues (Ploetz, 1994). Levels of infection were highest in malformed flowers
and vegetative shoots (c.65–85%), were much lower or non-existent in asymp-
tomatic tissues (0–11%), and were rare in branches (0–4%) even when they
supported malformed flowers or shoots. Remnant infections of F. mangiferae
in scaffold branches and trunks were restricted almost exclusively to branch
scars or dormant apices (Freeman, unpublished data). Reports of root
infection by F. oxysporum (Kumar and Beniwal, 1991) or F. mangiferae (Abdel-
Sattar, 1973; Kumar and Beniwal, 1991) causing malformation in seedling
plants have not been corroborated. Although roots can be infected by
GFP-labelled microconida of
Fusarium mangiferae
20.0 µm
Aceria mangiferae
Fig. 8.24. Body of the mango bud mite, Aceria mangiferae, to which green
fluorescent protein (gfp)-labelled microconidia of Fusarium mangiferae have adhered
(Photograph courtesy of E. Gamiel-Atinsky).
R.C. Ploetz and S. Freeman 268
F. mangiferae, these infections are not systemic and do not appear to result in
symptom development (Youssef et al., 2007).
The localized and variable levels of infection by F. mangiferae that have
been noted in diseased and non-symptomatic tissue (Ploetz, 1994; Youssef
et al., 2007), suggest that there are thresholds of infection, whereby malfor-
mation develops only after a sufficient proportion of a host meristem is colo-
nized by the pathogen. This hypothesis is supported by the long latent period
that exists before symptoms develop in artificially inoculated plants and the
hormonal perturbations that probably occur when meristematic tissues are
infected with this pathogen (van Staden et al., 1989; van Staden and Nichol-
son, 1989; Ploetz, 2001).
Management
Management of malformation can be difficult. New plantings should be
established with pathogen-free nursery stock. Scion material should never be
taken from an affected orchard, and any affected plants that are observed in
the nursery should be removed and burned immediately. Nurseries should
not be established in orchards that are affected by malformation. Once the
disease is found in an orchard, control is possible, but time consuming. In
these cases, cultural management has been most effective (Narisimhan, 1959;
Singh et al., 1974; Manicom, 1989). All affected terminals and the subtending
three nodes are cut from trees, removed from the field and burned. Unfortu-
nately, producers may be unwilling to devote the effort that is required to
ensure that this approach succeeds. In addition, it may be difficult to impose
this treatment on large trees.
A diverse array of pesticides, hormones and growth regulators has been
tested against malformation, but these measures have been marginally effec-
tive. Singh et al. (1994) obtained moderate control with sulfates of cobalt (Co),
cadmium (Cd) and nickel (Ni) in India, but it is unlikely that these toxic com-
pounds could be used safely. Darvas (1987) reduced the percentage of mal-
formed inflorescences from 96% to 48% by injecting ‘Keitt’ trees with the
fungicide fosetyl-Al. This reduction was significant (P < 0.05), but the increase
in fruit yield, 46–95 kg of fruit/tree, was not. Other fungicidal compounds
have been generally less effective (Diekman et al., 1982; Chakrabarti and
Ghosal, 1989). In general, the protected, internal location of the pathogen in
affected trees makes it difficult to control this disease. When applied as foliar
sprays or via continuous drip irrigation, prochloraz reduced the severity of
malformation significantly in Israel, but this was dependent on the timing of
application (Freeman et al., unpublished data). Although disease was not
completely controlled, this and other systemic fungicides might be useful in
future integrated management programmes that would incorporate other
measures such as removal of symptomatic terminals and use of tolerant
cultivars.
Prakash and Srivistava (1987) indicated that ‘There is great variation in
the susceptibility of existing varieties.’ Unfortunately, controlled inoculations
have not been used to determine cultivar resistance, and these reports have
come from non-replicated tests; cultivars listed as ‘resistant’ may have come
Foliar, Floral and Soilborne Diseases 269
from healthy nursery stock or may have escaped infection once planted in
the field (Ploetz, 2001). For example, Bastawros (1996) reported that two
newly introduced cultivars in Egypt, ‘Kent’ and ‘Keitt’, were immune (0%
disease); however, these cultivars are susceptible in Florida USA (Ploetz,
unpublished data). Controlled inoculations with grafted plants of different
genotypes and quantified levels of virulent isolates of F. mangiferae have not
been reported.
Recently, Ploetz (unpublished data) utilized previously described proto-
cols (Freeman et al., 1999) to assess malformation development on grafted
plants of diverse genotypes. Disease development was affected by: the length
of time after inoculation and inoculated apical buds remained dormant after
inoculation; the isolate of F. mangiferae that was used for inoculation; and
mango genotype. Virulent isolates, patience (latent period ranged from 40 to
210 days), and sufficient replication were needed to successfully conduct
screenings for response to malformation. Future work is warranted to investi-
gate attributes that are related, and might predict resistance, to this disease.
The symptoms of malformation suggest that a hormone imbalance
occurs in affected tissues. Singh and Dhillon (1989) assayed levels of indole
acetic acid (IAA), gibberellic acid (GA
3
) and zeatin in malformed and healthy
mango seedlings. Whereas IAA and GA
3
levels were, respectively, about ten
and five times lower in malformed plants, levels of zeatin were about five
times higher. Van Staden and colleagues (Nicholson and van Staden, 1988;
van Staden and Nicholson, 1989; van Staden et al., 1989) examined specific
cytokinins produced by mango and ‘F. moniliforme’ (presumably F. mangiferae).
They determined that the cytokinin complements in healthy and malformed
panicles differed qualitatively and quantitatively, and that the pathogen pro-
duced some of the hormones and metabolites that were implicated in dis-
ease development. However, it was impossible to assign unequivocal roles
for production of hormones by the host and pathogen and the subsequent
development of symptoms. For example, whether production of hormones
by the pathogen directly caused the noted changes or whether hormone pro-
duction by the host was somehow altered in the presence of the pathogen
was not clear. Additional work would be needed to clarify these interactions,
and whether F. sterilihyphosum and the unnamed pathogens in Brazil and
Mexico also produce cytokinins or other hormones in affected plants.
Parasitic plants
The family Loranthaceae contains several parasitic plant species that affect
mango. In Malaysia, Dendrophthoe (fomerly Loranthus) pentandra Linn. is
the most important species (Lim and Khoo, 1985). Other Dendrophthoe spp.,
Elytranthe spp. and Viscum spp. are also known in Malaysia, but are less
important. In India, Dendrophthoe falcata (formerly Loranthus longiflorus) is
common, and other, less frequently encountered, species include Macrosolen
cochinchinensis, Helicanthes elasticus and Elytranthe capitellata (Majumder
and Sharma, 1990). These parasites are usually only important in neglected
R.C. Ploetz and S. Freeman 270
orchards. Although they are photosynthetically self-sufficient, the plants
obtain water and minerals from the host plant via haustoria that penetrate
and colonize the host vascular system. In severe cases, the removal of water
and minerals from parasitized branches is sufficient to reduce the vitality and
yields of trees.
Since the appearance of these plants is distinct from the mango host, they are
easily distinguished in infected trees (Lim and Khoo, 1985). The points at which
the mango host is penetrated are usually characterized by swollen growths
called burrs. Lim and Khoo (1985) and Majumder and Sharma (1990) indicated
that affected portions of trees should be removed far enough below burrs to
remove haustoria of the parasite. After affected tissues are removed, cut sur-
faces can be treated with creosote or other wound dressings. These plants can
also be treated with herbicides, such as 2,4-dichlorophenoxyacetic acid (2,4-D),
but these are dosage sensitive treatments and pose a risk to the host plant.
Phoma blight
Phoma blight is widespread in India (Prakash and Singh, 1977). It occurs only on
old leaves. Initially, lesions are minute and yellow-brown (Prakash and Singh,
1977). As they expand they darken to brown or cinnamon, become irregular, and
may ultimately develop dark margins and dull-grey centres. In severe cases,
necrotic patches as large as 13 cm in diameter may form that cause defoliation.
The disease is caused by Phoma glomerata (Corda) Wollenw. and Hochapf
(Prakash and Singh, 1977). It forms globose to obpyriform, light-coloured to car-
bonaceous pycnidia that average 30–400 m in diameter. Pycnidia have one to
three ostioles, form singly or in clusters, and have short necks. On PDA, pyc-
nidia and conidia are abundant. Conidia are hyaline to dark coloured, ovoid to
ellipsoid, unicellular or occasionally bicellular, and average 8.3 × 3.2 m.
Phoma leafspot
Another Phoma sp. causes a leafspot in India (Prakash and Singh, 1976b), and
is referred to as phoma leafspot. On young leaves, Phoma sorghina (Sacc.)
Boerema. Doren. and Vankest causes irregular to roughly circular, water-
soaked spots, up to 2.5 mm in diameter. Lesions are brown with a yellow to
brown margin. Lesions on midribs are elongated and more conspicuous, and
may coalesce to up to 14 cm in diameter. They can be confused with those
caused by anthracnose.
Pink disease
A basidiomycete, Erythricium salmonicolor (Berk. and Broome) Burdsall (syn-
onyms: Corticium salmonicolor Berk. and Broome, and Phanerocbaete salmoni-
color (Berk. and Broome) Jülich; anamorph: Necator decretus Massee) causes
Foliar, Floral and Soilborne Diseases 271
pink disease. Pink disease affects many economically important woody
plants in the humid tropics, where it is one of the most destructive diseases
of mango (Holliday, 1980). The disease is also known as cobweb, rubellosis
and thread blight (Prakash and Srivistava, 1987). It has been most thor-
oughly studied on rubber, Hevea brasiliensis, an important host in South-east
Asia (Rao, 1975). On mango, pink disease can significantly reduce tree vigour
and yield, especially in 6- to 15-year-old trees (Lim and Khoo, 1985).
Symptoms first appear as white, felty mycelial threads on twigs and
branch crotches (Lim and Khoo, 1985). If favourable conditions persist, the
mycelial threads coalesce to form a rough, pink crust on the bark surface. This
stage usually takes 1 to several months to develop and coincides with the
penetration of bark and internal colonization of the tree. Affected bark often
cracks. As the fungus kills the vascular and cambial areas beneath the bark,
branches above the colonized areas die, resulting in a sparse, patchy canopy.
Two types of sporulation occur (Holliday, 1980; Lim and Khoo, 1985).
Erythricium salmonicolor produces a smooth, clammy, pinkish white hyme-
nium over the pink crust it forms on bark. Basidiospores form on the hyme-
nium and are borne on sterigmata on narrowly clavate to cylindrical basidia
(Fig. 8.25). Basidiospores are hyaline, broadly ellipsoidal and 8–10 × 5–7 m.
Conidia of N. decretus, which are produced in reddish-orange sporodochia,
are hyaline, ellipsoidal, unicellular and 10–18 × 6–12 m. Although the infec-
tion process has apparently not been studied in mango, basidiospores can
infect rubber trees (Holliday, 1980). The anamorph and teleomorph are
(c)
(a)
(b)
(f )
(e)
(d)
100 µ
20 µ
Fig. 8.25. (a) Conidium-bearing pustule, and (f) conidiogenous cells and conidia
of Necator decretus, and (b) hymenium, (c) basal hyphae, (d) immature and mature
basidia, and (e) basidiospores of its teleomorph, Erythricium salmonicolor (Source:
from CMI description no. 511).
R.C. Ploetz and S. Freeman 272
formed during wet conditions, and conidia and basidiospores are dispersed
by rainsplash and wind.
Pink disease management relies on early detection and removal of
affected tissues from orchards. When removal of syptomatic tissues is imprac-
tical, control depends upon treatment with fungicides. Several protectant
and systemic fungicides are effective (Lim, 1994). They should be used as
soon as symptoms are evident, and as long as the disease is present. All cul-
tivars of mango that have been tested in Malaysia are susceptible (Lim and
Khoo, 1985).
Powdery mildew
Powdery mildew is a widespread and important disease of leaves, panicles
and fruit. The disease can result in yield reductions as high as 90%, due
mainly to its effect on fruit set and development (Schoeman et al., 1995).
Symptoms
Mango cultivars vary in their response to powdery mildew (Palti et al., 1974).
On the most susceptible cultivars, virtually all foliar, floral and fruit parts of
the plant are affected (Plate 46). Powdery growth can cover all tissues on
panicles, resulting in a brown, shrivelled necrosis. Since fruit set and reten-
tion can be affected, the disease can have a profound impact on yield. Foliage
can also be damaged significantly, and young leaves are most susceptible.
White, powdery coatings of conidia develop on either side or both sides of
leaves, depending on the cultivar. When damage occurs on the undersides of
leaves it is often restricted to the mid-rib. In all cases, leaves become dis-
torted, and affected areas turn purple and ultimately necrotic.
Aetiology
Powdery mildew is caused by Oidium mangiferae Berthet, a host-specific fun-
gus (Prakash and Srivistava, 1987; Ploetz and Prakash, 1997). It was first
described in Brazil (Berthet, 1914), and is now recognized in most mango-
producing regions (Palti et al., 1974). Conidium and haustorium traits indi-
cate that O. mangiferae belongs to the Erysiphe polygoni group (Johnson, 1994a).
Although the pathogen was originally classified as Erysiphe cichoracearum by
Wagle (1928), Uppal et al. (1941) noted that it produced saccate and lobed
appressoria, which are not characteristic of E. cichoracearum. The pathogen
has no known teleomorph, a common trait for powdery mildew fungi in the
tropics (Holliday, 1980). Conidia of O. mangiferae are unicellular, hyaline,
elliptical to barrel-shaped and measure 33–43 × 18–28 m (Uppal et al., 1941;
Palti et al., 1974). They are produced in large numbers on host surfaces, and
impart a powdery appearance to affected tissues (Plate 46). The lengths of
germ tubes vary depending upon RH, and they terminate in appressoria. Glob-
ular haustoria form in host epidermal cells. Conidiophores are of the pseudoid-
ium type, with two to four septa and a straight basal cell (Boesewinkel,
1980).
Foliar, Floral and Soilborne Diseases 273
Epidemiology
Powdery mildew is most severe during cool, dry weather. Conidia are dis-
seminated by wind and are released on a diurnal basis (Schoeman et al.,
1995). Peak spore release, between 11:00 to 16:00 h, was positively correlated
with hourly temperature and negatively correlated with RH, vapour pres-
sure deficit and leaf wetness (all P < 0.01). Conidia germinate at temperatures
ranging from 9 to 32°C (23°C is optimal), and at RH as low as 20% (Palti et al.,
1974). Since germination occurs in such a wide range of RH, disease develop-
ment is usually independent of this parameter. Infection can occur within
5–7 h, and conidia are produced within 5 days of infection. Disease develop-
ment occurs within a very broad range of temperatures, 10–31°C. Gupta
(1989) reported that dry weather encouraged disease development.
Management
Mango cultivars vary in their resistance to powdery mildew (Palti et al.,
1974). ‘Zill’, ‘Kent’, ‘Alphonso’, ‘Seddek’ and ‘Nam Doc Mai’ are very sus-
ceptible; ‘Haden’, ‘Glenn’, ‘Carrie’, ‘Zebda’, ‘Hindi be Sennara’, ‘Ewaise’ and
‘Keitt’ are moderately susceptible; and ‘Sensation’, ‘Tommy Atkins’ and
‘Kensington’ are slightly susceptible (Ploetz et al., 1994; Nofal and Haggag,
2006). In India, Tiwari et al. (2006) reported that ‘Baigan Phalli’, ‘Barbalia’,
‘Dabari’, ‘Dilpasand’, ‘Khirama’, ‘Nagarideeh’, ‘Oloor’ and ‘Totapari’ were
highly resistant and ‘Amrapali’ was most susceptible.
Schoeman et al. (1995) recommended that the first fungicide application
to control this disease should occur when panicles begin to change colour.
Assuming an effective period of 3 weeks for a given application, they con-
cluded that applications should continue every third week until panicle sus-
ceptibility decreased at the end of fruit set. Powdery mildew is easily
controlled with S (Johnson, 1994a). Other fungicides are effective, but many
have negative environmental impacts (Ray, 2003; Tavares et al., 2004). Foliar
sprays of di-potassium hydrogen orthophosphate (K
2
HPO
4
) and potassium
di-hydrogen orthophosphate (KH
2
PO
4
), systemic fungicides, and an alterna-
tion of fertilizer and systemic fungicides inhibited powdery mildew on pan-
icles (Reuveni et al., 1998; Nofal and Haggag, 2006). Treatments of the
fertilizers with half or a quarter of the recommended rate of sterol-inhibitor
fungicides and Kresoxym-methyl provided protection comparable with or
superior to that of standard fungicides alone (Oosthuyse, 1998; Reuveni et al.,
1998). Sulfur can burn flowers and young fruit during warm, sunny condi-
tions (Johnson, 1994a), and three fungicides used during bloom, dinocap,
fenbuconazole and hexaconazole, can reduce pollen germination (Dag et al.,
2001).
Scab
Elsinoë mangiferae Bitancourt and Jenkins (anamorph: Sphaceloma mangiferae
Bitancourt and Jenkins) causes scab on mango (Bitancourt and Jenkins, 1943;
Cook, 1975). The disease was first recognized in Cuba and Florida USA in the
R.C. Ploetz and S. Freeman 274
1940s and is now widespread in the western hemisphere. Scab is important
in nurseries since young host tissues are most susceptible, and because moist
environments aid infection (Ruehle and Ledin, 1955). Lesions, usually first
observed on the underside of leaves, are dark brown to black, and 1–2 mm in
diameter. They may enlarge to 5 mm in diameter and become light grey with
narrow, dark borders. Affected foliage develops a distorted appearance, and
greyish blotches are produced on stems.
Elsinoë mangiferae produces brownish ascocarps, 30–48 × 80–160 m, in
the host epidermis. Globular asci, 10–15 m in diameter, are dispersed in
ascocarps, and contain one to eight hyaline ascospores. Ascospores are
4–6 × 10–13 m, three septate and constricted at the median septum; the sub-
apical cell is longitudinally septate. Conidiophores of S. mangiferae are erect,
sinuous, 2.5–3.5 × 12–35 m, and wider at the base. Conidia are brown, one
or two celled, and 2–4 × 6–29 m.
Young host tissues are most susceptible. Rainy weather promotes sporu-
lation of the fungus, but specific information is not available on the epidemi-
ology of scab. Whether conidia and ascospores are infectious is not known.
Seca and sudden decline
This is a disease that is known by several different names in Brazil and the
Middle East and is the only one that routinely kills mango trees. ‘Seca’ (dry-
ing), ‘murcha’ (withering), branch blight and Recife sickness in Brazil, was
first recognized in Pernambuco in 1938, and is now also found in Bahia,
Goias, the Federal District, Rio de Janiero and São Paulo (Ribeiro, 1997; Colo-
simo et al., 2000; Silveira et al., 2006). It threatens neighbouring states due to
its efficient movement in infected propagation materials, on pruning equip-
ment, and via a mobile beetle vector.
In 1998, a disease termed ‘sudden decline’ began to kill trees in Oman
(Al Adawi et al., 2003, 2006), about the same time a similar problem (i.e. quick
decline or sudden death) was observed in Pakistan (Malik et al., 2005). In
many ways these diseases resembled seca. Circumstantial evidence sug-
gested that the disease was introduced from Brazil by a producer with
orchards in Oman and Pakistan (M. Deadman, Muscat, 2005, personal com-
munication). By 2007, many mango-producing areas in Oman and Pakistan
were affected and uncontrolled dissemination of infected germplasm may
have spread the disease elsewhere in the region. Its spread into India should
be investigated (A.W. Cooke, Indooropilly, 2007, personal communication).
Symptoms
Symptoms include: discoloration of the vascular cambium; exudation of an
amber-coloured gum from the trunk and branches, particularly from galler-
ies of the putative beetle vector of the pathogen; wilting; rapid death of
branches and entire trees without defoliation; and a scorched appearance of
dead trees (Plate 47) (Junqueira et al., 2002; Al Adawi et al., 2006). On grafted
trees, scions, rootstocks or both may be susceptible and exhibit vascular
Foliar, Floral and Soilborne Diseases 275
symptoms. In Oman, where susceptible Omani seedlings are used as root-
stocks, the disease is frequently a problem of rootstocks (Al Adawi et al.,
2006), whereas in Brazil, the disease is associated with the scion (P < 0.01)
(Colosimo et al., 2000); rootstock cultivar had an insignificant impact on dis-
ease development in the latter work (P > 0.05). When disease development
begins in the canopy, symptoms may initiate in a branch or portion of a tree,
but death of the entire plant usually follows. Where root/rootstock infection
is involved, sudden death of the entire tree usually occurs.
Aetiology
Ceratocystis fimbriata Ellis and Halst. sensu lato (s.l.) (anamorph: Thielaviopsis
sp.) was reported in Brazil in the 1930s (Viegas, 1960; Ribiero, 1980; Silveira
et al., 2006), and is recognized as the primary cause of seca. Diplodia recifiensis
Batista (= Lasiodiplodia theobromae?) was indicted as the cause of Recife sick-
ness in Brazil (Batista, 1947), but it probably plays no role or a secondary role
in the development of this disease (see below). In Oman, C. fimbriata s.l.
causes sudden decline, but L. theobromae and Ceratocystis omanensis Al Subhi,
M.J. Wingf., M. van Wyk and Deadman are also associated with the disease
(Al Adawi et al., 2006; van Wyk et al., 2007). The ease with which L. theobromae
and the difficulty with which C. fimbriata s.l. are recovered from affected trees
may have been responsible for previous assumptions that ‘D. recifiensis’
caused Recife sickness in Brazil and L. theobromae caused sudden decline in
Oman (Batista, 1947; Ploetz and Prakash, 1997; Al Adawi et al., 2003, 2006).
Ceratocystis contains many pathogens, particularly of trees (Kile, 1993).
The wide host range of C. fimbriata s.l. led Webster and Butler (1967) to
hypothesize that it was a species complex, and DNA sequences have begun
to delineate some of the host-specific, often morphologically indistinct, taxa
in the species (van Wyk et al., 2007). A contemporary view is that C. fimbriata
sensu stricto (s.s.) specifically refers to the cause of black rot of sweet potato
(Ipomoea batatas L.) on which it was first described (Halsted and Fairchild,
1891). Other cryptic, monophyletic lineages of C. fimbriata s.l. have been
described as distinct species (Engelbrecht and Harrington, 2005; Johnson
et al., 2005; van Wyk et al., 2005, 2007), and more will likely follow.
Two new Ceratocystis spp. have been described on mango in the Oman
Gulf region. Ceratocystis omanensis belongs to the Ceratocystis moniliformis
Hedgc. s.l. species complex (Al Subhi et al., 2006), which are typically not
primary pathogens. Ceratocystis omanensis is a minor pathogen on mango.
The primary sudden decline agent in Oman and Pakistan, C. fimbriata s.l.,
represents a monophyletic lineage based on ITS, -tubulin and translation
elongation factor (TEF) 1- DNA sequence comparisons, and it has unique
morphological characteristics; it was renamed Ceratocystis manginecans M.
van Wyk, A Al Adawi and M.J. Wingf. sp. nov. (van Wyk et al., 2007).
On 2% malt extract agar (MEA), colonies of C. manginecans are greyish
olive and have a banana odour (van Wyk et al., 2007). Hyphae are smooth
and segmented (Fig. 8.26). Ascomatal bases are globose, black and (153–)192–
254(–281) m in diameter; ascomatal necks are dark brown, lighter towards
the apices (514–)557–635(–673) m long, (25–)32–42(–48) m, wide at the
R.C. Ploetz and S. Freeman 276
base, and (14–)16–22(–26) m wide at the tip; and ostiolar hyphae are hya-
line, divergent and (42–)45–59(–69) m long. Asci are evanescent, and
ascospores are hyaline, hat-shaped, 3–4 m long, and 4–5 m wide without,
and 7–8 m wide within the sheath. Primary conidiophores are phialidic,
lageniform, hyaline, (72–)81–109(–144) m long, 5–7(–9) m wide at the base,
6–8(–9) m wide at the broadest point, and 3–6 m wide at the tip. Secondary
conidiophores are tube like, flared at the mouth, short, hyaline, (59–)65–
77(–84) m long, 5–8 m wide at the base and (5–)6–8 m wide at the tip.
Primary conidia are hyaline, cylindrical, (15–)23–29(–33) m long, and 3–6
m wide; and secondary conidia are hyaline, barrel shaped, (8–)9–11(–12) m
long, and 5–7(–8) m wide. Chlamydospores are brown, thick-walled, glo-
bose to subglobose, (11–)12–14 m long and 9–11(–12) m wide.
Two isolates of C. fimbriata s.l. from mango in Brazil (CBS 114721 and CBS
600.70) have been compared to isolates of C. manginecans (van Wyk et al.,
2005, 2007). They are similar to, but differ significantly from, C. manginecans.
They reside with C. manginecans in a clade that contains other New World
(a)
(d) (g)
(c)
(e) (b) (f)
Fig. 8.26. Microscopic features of Ceratocystis manginecans: (a) globose ascoma,
(b) divergent ostiolar hyphae, (c) hat-shaped ascospore, (d) segmented hypha,
(e) primary phialidic conidiophore with emerging cylindrical conidia, (f) secondary
conidiophore with emerging chain of barrel-shaped conidia, and (g) dematiaceous
chlamydospores and cylindrical- and barrel-shaped conidia. Scale bars: (a) = 100 m,
(b) = 20 m, (c) = 5 m, (d) = 20 m, (e) = 20 m, (f) = 20 m, (g) = 20 m.
(Source: van Wyk et al., 2007).
Foliar, Floral and Soilborne Diseases 277
species, Ceratocystis cacaofunesta and Ceratocystis platani. Research is needed
to: (i) examine additional isolates of C. fimbriata s.l. from mango in Brazil; (ii)
describe the putative, new species; (iii) determine whether C. manginecans is
present in Brazil; and (iv) clarify pathogenic variation in the agent(s) in Oman
and Brazil. At least two pathotypes of C. fimbriata s.l. are evident in Brazil
(Rossetto et al., 1996; Junqueira et al., 2002; Silveira et al., 2006).
Rossetto et al. (1996) evaluated 15 cultivars against two isolates of the
pathogen in Brazil. Eight-year-old trees were inoculated c.40 cm beneath
branch apices with IAC FITO 4905, which is pathogenic to ‘Jasmim’, and IAC
FITO 334-1, which is not. ‘São Quirino’, ‘Irwin’, ‘Edwards’ and ‘Van Dyke’
were resistant, and ‘IAC 100 Bourbon’ was moderately resistant. ‘Glenn’, ‘Joe
Welch’, ‘Zill’ and ‘Haden’ were susceptible, and ‘Kent’ responded as ‘Jas-
mim’, resisting IAC FITO 334-1 and succumbing to IAC FITO 4905.
Epidemiology
Genotype has a profound impact on disease development, and severe epi-
demics occur wherever susceptible rootstocks and/or scions are used. Greater
disease develops when trees are stressed, although it is not clear whether this
results from an increased attraction of the vector to stressed trees or reduced
disease resistance in the host. The associated pathogens are moved easily in
infected germplasm, the route by which the diseases have spread in Brazil
and probably to Oman. Pruning implements also move the pathogen, and soil,
once infested with chlamydospores of the pathogen, can be a long-term reser-
voir of inoculum. Insect dissemination plays a particularly insidious role.
Beetles (Coleoptera: Scolytidae) are closely associated with seca in Brazil
(Batista, 1947; Viegas, 1960; Piza, 1966; Ribiero, 1980). Batista (1947) indicated
that Xyleborus affinis was the sole vector of D. recifiensis. In contrast, Ribiero
(1980) reported that Hypocryphalus mangiferae Stebbing was the primary vec-
tor of C. fimbriata s.l. (Fig. 8.27). It produced galleries in the cambium of
affected trees (Plate 47a), and was the only scolytid found on healthy, as well
as diseased, trees. Hypocryphalus mangiferae is also associated with the dis-
eases in Oman and Pakistan, where C. manginecans is recovered from the
insect and trees are commonly found with insect probing damage before dis-
ease develops (Al Adawi et al., 2006; van Wyk et al., 2007).
The interactions between H. mangiferae and the Ceratocystis pathogens
of mango are incompletely understood. In olfactometer tests in Brazil,
H. mangiferae was attracted to cultures of C. fimbriata s.l., and larvae of the
insect were raised to adulthood on the fungus (Ribiero, 1980). Several other
species, many of which are in the genus Xyleborus, were also associated with
seca, but because they were found only in diseased trees they appeared to be
opportunistic feeders on C. fimbriata s.l. rather than primary vectors. Although
the sequence of events in Brazil and the Oman Gulf has not been studied, it
is probable that H. mangiferae contaminates its body with these pathogens
while feeding in diseased trees and subsequently disseminates the pathogen
to healthy trees.
Hypocryphalus mangiferae is thought to be native to some of the same
areas in southern Asia where mango evolved (Wood, 1982; Butani, 1993;
R.C. Ploetz and S. Freeman 278
Atkinson and Peck, 1994; Mukherjee, 1997). Thus, the insect would have
been introduced into Brazil and would have been a new encounter, rather
than coevolved, relationship with C. fimbriata s.l. (van Wyk et al., 2007). In
contrast, if C. manginecans were introduced into Oman and Pakistan from
Brazil, it may have then established a relationship with a native insect.
Although the available information suggests that the H. mangiferae–
Ceratocystis interactions on mango were recent, opportunistic encounters in
the New World, additional work is needed.
Management
Given the ease with which these pathogens are moved and their destructive
impact, preventing their dissemination to new areas must be a high priority.
Pathogen-free propagation material should be used whenever new plantings
are established and germplasm is moved. Clean pruning implements should
be used in affected areas, and should be frequently disinfested with bleach,
formalin or other disinfestants (Junqueira et al., 2002). Trees that have been
killed by the disease must be removed and destroyed as they are significant
reservoirs of inoculum and infested vectors. Where partially resistant culti-
vars are grown, the removal and burning of affected branches and treatment
of the exposed branch stubs with Cu fungicides is recommended (Ribeiro
et al., 1995; Ribeiro, 1997).
Managing these diseases with fungicides on susceptible cultivars would
be a challenge. External applications of protectant or systemic fungicides
would probably be ineffective, given the internal, protected location of the
pathogen. Injecting fungicides, as is done to control Dutch elm disease, might
be effective. However, this would probably not be allowed where concerns
Fig. 8.27. Hypocryphalus mangiferae, vector of the seca and sudden decline
pathogens (Photograph courtesy of R.C. Ploetz).
Foliar, Floral and Soilborne Diseases 279
with pesticide contamination of fruit exist. Treatment of germplasm collec-
tions and young, non-bearing trees might be the only situations in which
fungicide injection would be possible.
Genetic resistance offers the best hope for managing these diseases. Var-
ious levels of tolerance have been observed in Brazil and resistant clones
have been developed. However, pathogenic variation in the causal fungus in
Brazil has hindered progress (Rossetto et al., 1996; Junqueira et al., 2002; Sil-
veira et al., 2006). Although disease responses of some genotypes vary in dif-
ferent production areas in the country, ‘Manga D’agua’, ‘Pico’, ‘IAC 101’,
‘IAC 102’, ‘Edwards’, ‘Van Dyke’ and ‘Carabao’ resist two races of the patho-
gen, and ‘Rosa’, ‘Sabina’, ‘Sao Quirino’, ‘Oliveira Neto’, ‘Jasmim’, ‘Sensation’,
‘Irwin’ and ‘Tommy Atkins’ are generally tolerant (Ribiero, 1997; Junqueira
et al., 2002). ‘Kent’ and ‘Jasmim’ respond differentially (see above), and
‘Coquinho’, ‘Glenn’, ‘Joe Welch’, ‘Zill’ and ‘Haden’ are susceptible. Although
‘Espada’ is also reported to be tolerant, old trees are frequently attacked. In
commercial orchards, the disease on ‘Espada’ is managed by grafting onto
resistant rootstocks and pruning diseased branches. Colosimo et al. (2000)
worked with other scion cutivars, although in a single location (and against
a single pathotype?). They reported that ‘Oliveira’ was most resistant, ‘Car-
lota’, ‘Imperial’, ‘Extrema’ and ‘Pahiri’ had intermediate resistance, and
‘Bourbon’ was most susceptible.
One must also recognize the impact of other diseases on different
cultivars. Carvalho et al. (2004) described two new cultivars, ‘IAC 103 Espada
Vermelha’ and ‘IAC 109 Votupa’, which had moderate resistance to seca.
‘IAC 103 Espada Vermelha’ also had moderate resistance to powdery mil-
dew but was susceptible to anthracnose. Both cultivars were susceptible to
malformation.
Stigmina leafspot
Stigmina leafspot is caused by Stigmina mangiferae (Koorders) Ellis (synonym:
Cercospora mangiferae Koorders; a teleomorph for the fungus is not known).
Lim and Khoo (1985) indicated that the disease occurs on a range of cultivars,
and is most severe during rainy weather. Both young and old leaves are
affected. Dark-brown spots, 1–2 mm in diameter, are formed initially by the
fungus. These can enlarge and coalesce to 1 cm or larger, and are surrounded
by conspicuous chlorotic haloes that aid diagnosis of this disease. The fungus
produces large, olive-grey conidia, 30–60 × 3.5–5.0 m, usually on the lower
leaf surface (Fig. 8.28). Conidia are wider at the base than the apex, are
straight to curved, have three to seven septa, and are borne in subglobular,
dark stromata that are 20–60 m in diameter.
Although the fungus sporulates sparsely on artificial media, it produces
copious conidia on necrotic host tissue, especially in leaf litter. Thus, Lim and
Khoo (1985) suggested that removing such debris from orchards and burning
it would assist control efforts. Several different fungicides are effective (Lim
and Khoo, 1985).
R.C. Ploetz and S. Freeman 280
8.3 Soilborne Diseases
Although soilborne diseases of mango are relatively less important than
foliar and floral diseases, they can cause significant damage to seedlings,
nursery stock and mature trees. In general, the pathogens that are involved
are different from those that cause problems above ground. Chemical
Fig. 8.28. Conidia of Stigmina mangiferae, cause of stigmina leaf spot (Source: from
CMI description no. 097).
Foliar, Floral and Soilborne Diseases 281
management is indicated rarely for these diseases; sanitation and other
cultural measures are used most often.
Black root rot
Black root rot is reported to be an uncommon problem on young mango trees
(Lim and Khoo, 1985). Canopies of affected plants wilt suddenly and subse-
quently defoliate. Roots exhibit a water-soaked, blackened decay, and have
an unpleasant, putrid odour. Although black root rot is associated with pro-
longed flooding, its precise aetiology is not known. Several species of fungi
have been recovered from affected plants, including Fusarium solani, F. oxyspo-
rum and L. theobromae, but these were thought to be secondary colonizers of
roots (Lim and Khoo, 1985). Although mango is generally considered to be
flood intolerant, its flood tolerance is actually variable (Larson, 1991). Varia-
tion in the responses of individual trees in orchards is evident after flooding,
and when potted plants are flooded experimentally, they usually adapt by
forming hypertrophied lenticels (intumescence) (Larson et al., 1993). Plants
that do not adapt in this manner succumb fairly rapidly. Roots of the latter
plants have symptoms of black root rot (R.C. Ploetz, Homestead, Florida,
1988, personal observations). Although flood tolerance is environmentally
and biochemically complex (Larson et al., 1993), some of the fungi reported
by Lim and Khoo (1985) may interact with flood-induced stress in this host
to cause black root rot.
Nematode damage
Decline of mango trees due to nematodes has been reported in various
regions (Khan et al., 1971, 2005; McSorley et al., 1980; Anita and Chaubey,
2003). Infestations occur in areas where warm temperatures and sandy,
moist, well-drained soils predominate (Ploetz et al., 1994). Many nematode
species have been recovered from mango roots, including: Helicotylenchus
dihystera (Cobb) Sher, Quinisulcius acutus (Allen) Siddiqi, Rotylenchulus reniformis
Linford and Oliveira, Criconemella sp., Pratylenchus brachyurus (Godfrey) Fil-
ipjev and Schuurmans Stekhoven, Xiphinema sp., Meloidogyne sp., Praty-
lenchus sp. and Hoplolaimus sp. However, only Hemicriconemoides mangiferae
Siddiqi is pathogenic (Powers and McSorley, 1994). Although high popula-
tions of R. reniformis are often found on mango trees, no correlation has
been shown between their density and tree health.
Populations of H. mangiferae vary according to soil moisture and tem-
perature (Khan et al., 1971). Soil moisture < 10% and > 30%, as well as tem-
peratures < 15°C and > 35°C are detrimental to nematode survival and are
likely to reduce populations. In addition, tree age appears to be a significant
factor, since H. mangiferae is found more frequently on old (> 10 years) than
young trees (< 3 years). Management is difficult and may depend on pre-
plant chemical applications plus cultural control measures (McSorley et al.,
R.C. Ploetz and S. Freeman 282
1981). Phenamiphos and 1,2-dibromo-3-chloropropane reduce levels of
H. mangiferae after planting; however, neither are registered for use in the
USA. Anita and Chaubey (2004) reported that high organic matter content in
the rhizosphere had a detrimental effect on nematode populations in the
field.
During orchard establishment, nematode-free nursery stock should be
used. Since H. mangiferae is partially endoparasitic, it is moved easily to clean
field sites. The use of clean planting material in infested fields should also be
avoided. If such land must be used, soil fumigation prior to planting should
be conducted. Fruit yields may still be maintained if infected trees are well
irrigated and fertilized.
Phytophthora diseases
Phytophthora palmivora (E.E. Butler) (Oomycota) causes diseases of mango in
several areas. It caused wilt, crown rot, root rot and the death of nursery trees
in Arizona USA, the Philippines and Thailand (Kueprakone et al., 1986;
Matheron and Matejka, 1988; Tsao et al., 1994). Gumming and conspicuous
bark lesions develop above ground on these plants, whereas root and crown
rots are evident at or below the ground level. Crowded conditions and exces-
sive irrigation and rainfall exacerbate these diseases. Sanitation, the use of
less-crowded conditions and reduced irrigation would be beneficial.
Damage has also been recorded on the trunks of field-grown, mature
trees in the Ivory Coast (Lourd and Keuli, 1975), and on fruit in Australia,
Malaysia and West Africa (Turner, 1960 cited in Chee, 1969; Cooke, 2007).
Mortality of trees is not observed, but substantial stem cracking and bleed-
ing does occur. The symptoms that occur on fruit have not been recorded in
Malaysia and West Africa, but on ‘Calypso’ fruit in Australia, a firm chocolate-
brown decay is produced that has a sweet odour. Fruit isolates in Australia
caused leaf blight and crown canker on mango seedlings (Fig. 8.29).
Phytophthora palmivora has coenocytic hyphae, up to 7 m in diameter, papil-
late sporangia, 31–56.4 × 20.7–36.7 m, which germinate either directly with
germ tubes or indirectly with motile zoospores (Fig. 8.30) (Waterhouse, 1970;
Erwin and Ribiero, 1996). Zoospores are the primary infective propagule,
and require free water for movement. Phytophthora palmivora is heterothallic.
Antheridia are amphigynous and oogonia are spherical. Chlamydospores,
c.35 m in diameter, are also formed.
Recently, a Phytophthora sp. was isolated in Spain from mango trees that
were wilted, chlorotic and had sparse canopies and cracked bark (Zea-
Bonilla et al., 2007). On V8 agar, sporangia were semi-papillate, obovoid
and 51 (28–52) × 36 (22–37) m. Paragynous antheridia, spherical oogonia
and oospores of 28 (19–32) m in diameter were produced homothalli-
cally. Ribosomal DNA sequences (ITS1, 5.8S rDNA and ITS2) (GenBank
Accession No. AM235209) were compared with those in GenBank; the
closest match, 99% identity, was with Phytophthora citricola, corroborating
the above morphological identification (Fig. 8.31). An isolate deposited in
Foliar, Floral and Soilborne Diseases 283
Fig. 8.29. Symptoms induced by Phytophthora palmivora after artificial inoculation
of (a) stems and (b) foliage of mango seedlings (Photographs courtesy of A.W. Cooke).
Fig. 8.30. (a) Sporangia, (b) oogonia with amphygynous antheridia and oospores,
and (c) chlamydospore of Phytophthora palmivora (Source: from CMI description
no. 831).
(a) (b)
R.C. Ploetz and S. Freeman 284
the Spanish Type Culture Collection, CECT 20567, caused root rot on
‘Florida’ and lesions on leaves and stems of seedlings of ‘Gomera 3’.
Root rot and damping off
The oomycete, Pythium vexans de Bary, can cause root rot and wilt of seed-
lings (Lim and Khoo, 1985). In Malaysia, this pathogen caused seedling losses
of up to 30% in nurseries. Symptoms included wilting of foliage, which
initially becomes pale green, but later develops necrotic patches. Roots
develop a wet, blackened necrosis that begins in fine roots and progresses to
larger roots and the root collar. Death of seedlings often occurs. Lim and
Khoo (1985) indicated that overcrowding, excessive moisture and the use of
polybags favoured this disease.
Prakash and Singh (1980) reported that the basidiomycete Rhizoctonia
solani Kuhn (teleomorph: Thanatephorus cucumeris (Frank) Donk) caused root
and damping off of seedlings in India (Fig. 8.32). Affected tissues become
soft, dark brown or black, and seedlings may ultimately become completely
girdled and collapse. Mycelia and sclerotia of the pathogen form conspicu-
ously on affected tissues.
Sclerotium rot
This disease has been reported in Brazil (Almeida et al., 1979), India (Prakash
and Singh, 1976a) and the Philippines (Palo, 1933). The causal fungus, Sclero-
tium rolfsii Sacc. (teleomorph: Athelia rolfsii (Curzi) Tu and Kimbrough;
Fig. 8.31. (a) Semipapillate sporangia and (b) oogonia of Phytophthora citricola
(Source: from CMI description no. 114).
Foliar, Floral and Soilborne Diseases 285
synonyms: Corticium rolfsii Curzi and Pellicularia rolftii E. West), produces
globular, brown sclerotia, 1.0–2.6 mm in diameter. Sclerotia are resilient struc-
tures that enable the pathogen to survive adverse environmental conditions.
Symptoms begin with the formation of felty white tufts of mycelium of the
pathogen around the base of seedlings. The fungus can girdle the entire stem
to a height of 5 cm or more above the soil line. It eventually forms conspicu-
ous sclerotia in high numbers. Ultimately, seedlings wilt and die. Seed may
also rot prior to germination. The disease is controlled via sanitation and the
disinfestation of seedbeds.
Verticillium wilt
Verticillium wilt of mango was first reported in Florida USA (Marlatt et al.,
1970). The disease was originally attributed to Verticillium albo-atrum Reinke
and Berth., but this was before Verticillium dahliae Kleb. was recognized as a
distinct species. Since the causal fungus described by Marlatt et al. (1970)
formed microsclerotia, it is clear that V. dahliae was involved (Fig. 8.33).
Symptoms of the disease include a ‘firing’ and necrosis of leaves, usually
in a portion of the canopy. Sectoral development of the disease often does not
progress to other portions of the trees, which may recover. Killed leaves usu-
ally remain attached to the tree, and the xylem of affected branches is dis-
coloured brown (Fig. 8.34). Verticillium wilt is relatively uncommon, and is
(a)
(b)
(c)
20 µ
(d)
Fig. 8.32. (a) Sclerotial cells, (b) mycelium and (c) monilioid cells of Rhizoctonia
solani, and (d) basidia and basidiospores of its teleomorph, Thanatephorus cucumeris
(Source: from CMI description no. 406).
R.C. Ploetz and S. Freeman 286
found on land where susceptible vegetable crops (i.e. potato, tomato and
aubergine) were recently grown (Pohronezny and Marlatt, 1982). New mango
orchards should not be planted on such sites.
White root disease
Rigidoporus lignosus (Klotzsch) Imazeki, the basidiomycete that causes white
root disease, is a common soil inhabitant in the humid tropics of Africa and
Asia (Holliday, 1980). It has also been reported in the western hemisphere,
but the identity of the fungus there is unclear. Rigidoporus lignosus has a wide
host range on woody perennials, including rubber, the host on which the
pathogen was first reported (1904 in Malaysia). The significant losses in rub-
ber plantations in the eastern hemisphere have resulted in considerable
research on this pathogen (Nandris et al., 1987).
Rigidoporus lignosus produces white rhizomorphs on the surfaces of roots
and root crowns that later darken to a yellowish and then reddish colour
(Lim and Khoo, 1985; Nandris et al., 1987). The leading edge of the rhizo-
morph is well defined and seldom appears above ground. It undergoes a
morphogenic change to produce infectious hyphae that penetrate the host
epidermis and subsequently degrade wood. Rigidoporus lignosus produces a
non-differentiated white rot that affects lignin in host cell walls.
(a)
(b)
10 µ
(c) (d)
Fig. 8.33. (a) Verticilliate conidiophore, (b) phialospores and (c) immature and (d) mature
microsclerotia of Verticillium dahliae (Source: from CMI description no. 256).
Foliar, Floral and Soilborne Diseases 287
The fungus is most damaging on mango if orchards are established in
old rubber plantations or newly cleared jungle sites (Lim and Khoo, 1985).
Previously colonized stumps and debris from rubber and other hosts are pri-
mary sources of inoculum. Orange-yellow, bracket-like sporophores are pro-
duced during the rainy season on the root collar, trunk or exposed roots
(Fig. 8.35). Basidiospores are viable, but may play a secondary role in dis-
seminating the disease. Rhizomorphs are more significant epidemiologically,
since they grow rapidly and can advance great distances in soil in the absence
of woody substrates.
Fig. 8.34. Vascular discoloration caused by V. dahliae (Photograph courtesy of
R.C. Ploetz).
R.C. Ploetz and S. Freeman 288
White root disease is managed by eliminating or avoiding colonized
woody debris when new orchards are established (Lim and Khoo, 1985).
Unfortunately, this is difficult and often impractical. Alternative measures
include: (i) treating planting holes with S to promote the growth of antago-
nistic microorganisms; (ii) treating stumps after clearing operations with
chemicals that discourage their colonization by basidiospores; and (iii) estab-
lishment of leguminous cover crops that promote growth of the pathogen
and the eventual exhaustion of its energy reserves.
8.4 Conclusions
As mango production continues to increase in different regions, so will the
scope and types of disease problems. Although new fungicides and bacteri-
cides will be developed in the future, it is probable that reliance on these
tools will diminish. In recent years, the regulation of pesticides has increased
while the number of new compounds that have been registered for use has
decreased. Since it appears certain that this trend will continue, the problems
posed by diseases must be solved increasingly with alternative disease
control strategies.
10 µ (c)
(d)
(a) (b)
20 µ
Fig. 8.35. (a) Upper and (b) lower surface of sporophore, (c) basidiospores, and (d) hyphae of
Rigidoporus lignosus (Source: from CMI description no. 198).
Foliar, Floral and Soilborne Diseases 289
A willingness among producers to utilize new technologies will play a
role in this process. New methods to detect important pathogens have been
developed (Zheng and Ploetz, 2002), but more work is needed. Effective
detection protocols, especially for those pathogens that can colonize host tis-
sues without causing symptoms, could be used to interdict important, exotic
pathogens and identify pathogen-free propagation materials. Detection pro-
tocols could also indirectly assist disease control efforts through their use in
epidemiological studies, research on disease resistance, or to clarify portions
of disease cycles.
Acknowledgements
The authors thank Francisco Cazorla for comments on apical necrosis; Ber-
nard Slippers and Mike Wingfield for comments on the decline section; Syl-
via Fernandez, John Leslie, Christiano Lima, Kerry O’Donnell and Gerardo
Rodriquez for information on malformation; Ali Obaid Al-Adawi and Mike
Wingfield for comments on seca and sudden decline; and Robert Knight for
translating Portuguese articles on seca. The following individuals are thanked
for pictures: Francisco Cazorla, T.-K. Lim, Oliver Pruvost, Carolyn Cohen,
Suzanne Bullock, Efrat Gamliel-Atinsky, Eric Palevsky and Tony Cooke.
Kevin Hyde, editor of Fungal Diversity, is thanked for permission to use
micrographs of Ceratocystis manginecans.
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