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Mahua - Madhuca longifolia (Koenig) J.F. Macribide)

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 21
MAHUA
( Madhuca longifolia (Koenig) J.F. Macribide)
M.K. Dhakar, D.K. Sarolia, R.A. Kaushik, K.L. Kumawat,
Sanjay Singh and A.K. Singh

CONTENTS
1. INTRODUCTION
Mahua, the Indian Butter Tree (Madhuca 1. Introduction ...............................................305
longifolia (Koenig) J.F. Macribide) is an 2. Domestication ............................................307
important tree having vital socio­economic 3. Taxonomy ...................................................307
value and growing throughout the tropical 4. Centers of Origin/Centers of Diversity ......307
and subtropical region of the Indian 5. Objectives of Crop Improvement ...............308
subcontinent. It is a deciduous tree that 6. Cytogenetics...............................................308
grows widely under dry tropical and sub­ 7. Inheritance Pattern ....................................308
tropical climatic conditions. It is very hardy 8. Problem in Breeding ..................................309
and thrives well on rocky, gravely, saline 9. Floral Biology .............................................309
and sodic soils, even in pockets of soil 10. Different Species .......................................311
between crevices of barren rock (Singh,
11. Botanical Description .................................313
1998). It is one of those multipurpose forest
12. Crop Improvement Methods ......................313
tree species that provide an answer for
13. Future Thrust .............................................319
the three major Fs i.e food, fodder and
fuel (Patel et al., 2011).
Mahua is a tree valued for its fruit, seeds, which are the largest source of natural hard
fat commercially known as mahua butter or mowrah butter. Fruits are eaten as raw or
cooked. The fruit pulp may be utilized as source of sugar, whereas the dry husk makes
a good source of alcoholic fermentation. Seeds are good source of oil (Singh et al, 2005).
The oil obtained from kernel which is said to be useful for heart patients is used for edible
purpose and permitted for preparation of vegetable oil. Amount of oil obtained from seeds
of the fruit is higher than many oil seed crops and oil­bearing trees. In Mahua oil, linoleic
and unsaturated fatty acids are found, which are useful for heart patients, because it
reduces the cholesterol content in blood serum. Mahua oil is used in manufacture of soap,
306 Breeding of Underutilized Fruit Crops

lubricating grease, fatty alcohols and candles. Flowers of the plant are edible. The corolla
commonly called as mahua flowers is a rich source of sugar containing appreciable
amount of vitamins and minerals (Singh and Singh, 2005). The flowers are also used in
preparation of distilled liquor, portable spirits, vinegar and feed for livestock (Adhikary and
Adhikary, 1989). Midya and Brahmachary (1996) reported that the fresh flowers of
Mahua (B. latifolia Roxb.), that emit fragrance contain 2­acetyl­1­pyrroline (2AP), the
compound responsible for pleasant aroma in basmati and other scented rice. It was found
that 2AP gets synthesized only in fleshy corolla of mature flowers (Wakte et al., 2011).
The flowers show anti bacterial activity against E. coli and also are used against rice pest
diseases (Sujatha and Das, 1988). Next to sugarcane molasses, Mahua flowers contribute
as most important raw material for alcohol production (Singh and Agrawal, 1989).
Enzyme Invertase (â­D­fructofuranosidase, EC 3.2.1.26), present in the flowers of
Madhuca longifolia plays an important role during the preparation of fermented Ayurvedic
drugs known as ‘Arishta’ (Weerasooriya and Yatawara, 2003). Nutritive value of mahua
flower has been presented in the Table 1.

Table 1. Nutritional Properties of Mahua flower (Source: Kureel et.al., 2009)

Sr. No Constituents Value

1. Moisture (%) 19.8
2. Protein (%) 6.37
3. Fat (%) 0.50
4. Reducing Sugar (%) 50.62
5. Total Inverts (%) 54.24
6. Cane Sugar (%) 3.43
7. Total Sugar (%) 54.06
8. Ash (%) 4.36
9. Calcium (%) 8.00
10. Phosphorus (%) 2.00

The leaves are used for making plates for various purposes. Mahua seeds can also
be used for preparation of defatted flour, which has great potentiality in bakery products.
The saponin obtained after extraction has industrial and commercial application. Cake
obtained after extraction of oil is used as manure and has insecticidal properties. It
provides quality timber wood for various uses. Every part of Mahua yields an economic
product of great potential value; hence it is very useful tree for tribal and poor people of
India. People in small villages don’t have sufficient money for various resources to earn
their living that is why they are fully or partially dependent on the natural resources which
are easily available, cheap and renewable. Mahua flower is having potential to satisfy few
needs of rural people. High sugar content of Mahua flower allows it to be consumed as
jam, jelly or raw. The dried Mahua flower can be sold to the local distillery and hence
they can make good money from it. The fermentation waste can also be used as bio
 Mahua (Madhuca longifolia (Koenig) J.F. Macribide) 307

fertilizer which is eco­friendly and cheap in comparison to the expensive fertilisers (Jha
et al., 2013). Bulbul and Begum (2014) revealed that the methanol extract of M. indica
has got profound antioxidant activity. So the plants may be considered as good sources
of natural antioxidants for medicinal uses such as against aging and other diseases related
to free radical. Leaves and barks of Madhuca indica showed significant antioxidant
activity with IC50 value 61.832 ìg/ml and 66.342 ìg/ml respectively. The phenolic content
was found in leaf 62.43mg of GAE / gm of extractives and the amount of phenolic content
was 61.08mg of GAE / gm of extractives for bark which correlated with good antioxidant
potentiality. The ethanolic and aqueous leaf extracts of Madhuca indica has anti
hyperglycemic activity, is able to ameliorate the diabetic state and is probably a source
of hypoglycemic compound (Jebaseelan and Ramasubramanian, 2014). M. longifolia
leaves are effective biosorbent for the removal of Pb(II) ions from aqueous media. So,
these leaves can be used on industrial scale for removing Pb (II) ions effectively (Rehman
et al., 2013). The tree is very well known to rural folk since ages in India. The production
in India is mainly concentrated in the drier states and the produce is collected by the
villagers and sold in the local market. Its cultivation may be spread to arid and semi­arid
areas, resource­poor areas and wastelands where other crops can not be grown successfully.

2. DOMESTICATION
Mahua (Madhuca indica) is an oilseed tree which grows naturally on village common
lands in India. The local people, particularly the tribal, collect the fallen corolla and
consume it in the fresh or dehydrated form. Long gestation period and poor income are
major constraints for cultivation of mahua. Techniques have now been developed to
produce grafted plants of mahua which start bearing in the fourth year. Looking to the
economics and short gestation period, farmers are now keen to establish plantation of
mahua on their degraded lands as a commercial crop.

3. TAXONOMY
Mahua (Bassia latifolia Roxb) belongs to the family sapotaceae. The trees of Bassia
latifolia and B. longifolia grow up to an altitude of 1,200 m. The B. malabarica grows
in Western Ghats from Kanara to Travancore and also in the Himalayas. The trees of
Bassia butyracea grow in the Himalayan regions up to an altitude of 4,500 m.

4. CENTERS OF ORIGIN/CENTERS OF DIVERSITY

The mahua introduced from India to Australia and Polynesia (Troup, 1921; Anon. 1988).
The species Madhuca longifolia is distributed in northern, central and south­ern part of
peninsular India, and M. latifolia is found in some parts of central and north India.
Mahua, a characteristic tree of the dry region, is found in almost all parts of India.
It commonly grows in eastern Uttar Pradesh, Madhya Pradesh, Chattisgarh, Maharashtra,
Bihar, Jharkhand, Orissa, Andhra Pradesh and Gujarat. In Rajasthan, it is also found
growing on the wastelands particularly in southern part of the state. Dense population of
mahua trees can be seen in Dahod, Panchamahals and Vadodara districts of Gujarat.
 308 Breeding of Underutilized Fruit Crops

5. OBJECTIVES OF CROP IMPROVEMENT

Farmers’ preference and market demand should be kept in mind, while defining the
breeding objectives. Mahua is generally preferred by farmers as multipurpose tree for the
use of food, fodder, fruit, flower, timber and fuel wood. These intended uses help in
defining the ideotypes of this tree. Donald (1968) defined ideotype as “a biological model
which is expected to perform predictably, leading to greater quantities and qualities of crop
yield under defined environmental conditions”.
i. To develop promising genotypes having high yield potential along with early and
regularity in bearing.
ii. To develop short stature, precocious (rapid juvenile growth i.e. precocious in bearing)
and prolific bearer cultivars having high oil content in kernels.
iii. Breeding for resistance against biotic and a­biotic stresses may be the objective of
mahua improvement.
Ideotype specification for Maduca longifolia according to Chuntanaparb and
Ranganathan (1990) is given below:
Crown: Large with many branches for planting on common land, but narrow and
dance canopy for farm and home planting.
Stem: Single or multiple but with a long straight bole with branches high on the stem.
Root: Deep, strong taproot to withstand wind.

6. CYTOGENETICS
Mahua (Bassia latifolia Roxb) belongs to the family sapotaceae, indigenous to India,
having chromosome X = 7, 8.

7. INHERITANCE PATTERN
Correlation studies provide reliable information on nature and extent of relationship for
bringing out improvement in yield and other traits. There was significant positive association
of flowers and fruits per fascicle, flower weight and flower yield with fruit yield per plant.
These traits may be observed for their positive behaviour while selecting superior genotypes
Singh et al. (2005). Abraham et al. (2010) found significant positive correlation between
100 seed weight with seed length, breadth and thickness. Kernel oil content was negatively
correlated with all the above seed characters and found significant negative correlation
with hilum breadth, indicating that the seed size is not a selection criterion for identifying
a genotype with high oil content. Heteroblastic development associated with phyllotaxy of
leaves is generally noticed in Madhuca indica (cryptocotylar species) by Mundhra and
Paria (2009), where first two leaves are opposite and subsequent leaves are alternate.
This heteroblastic development of leaves serves as marker character and will help in
identification of the taxon from closely related species.
 Mahua (Madhuca longifolia (Koenig) J.F. Macribide) 309

8. PROBLEM IN BREEDING
i. It is cross pollinated species so leading to heterozygous nature of the crop.
Improvement of such crop require long period as compared to the annual crops.
ii. Mahua is generally associated with long gestation period of 8 to 10 years.
iii. Variability is declining in mahua at faster rate due to deforestation and human
interventions.
iv. Flowers and fruits drop is another major hindrance in breeding. Large number of
predators is associated with mahua due to its sweet corollas leading to more flowers
and fruit drop.
v. Self and hand pollination gives low fruit set as compared to natural pollinations.
vi. As other forest crop mahua seed collection is difficult. Birds and animals eat the
fleshy fruits.
vii. There is no commercial cultivar available in this crop but some of the promising
genotypes are available at NBPGR, New Delhi and NDUAT, Faizabad. Such genotypes
should be improved by using modern breeding techniques. Fruit or tree breeders
should lead the improvement work by collaboration with genetics person, biotechnologist,
botanist and taxonomist.

9. FLORAL BIOLOGY
9.1. Flowering
M. latifolia is long­lived and starts bearing from about the 10th year. A full grown tree
can produce up to 90 kg of flowers in a year. Leaf fall occurs from February to April
and the trees are often leafless at the time of flowering which takes place, once in a year,
i.e. during March­April. The flow­ering time varies with the local conditions. Singh et al.
(2006) recorded that the peak period of flowering and fruit set commenced in the month
of March ­ April in different genotypes. Time taken for complete development of flower
bud from its visible initiation to anthesis varies from 20 to 30 days (Kureel et al., 2009).
Wakte et al. (2011) identified four different flower developmental stages viz.
Stage 1. Bud completely closed,
Stage 2. Bud closed with the style protruding,
Stage 3. Flower partly open with the style protruding and
Stage 4. Fully ripe flower with shedding corolla (Fig. 1).

9.2. Flower
Flowers are in dense clusters at the end of branches with long pedicels; calyx coriaceous
(i.e. leathery; stiff and tough, but somewhat flexible.); corolla tublar, fleshy, cream­
coloured, scented, caduceus (i.e. falling off early). The corolla falls off before or with the
appearance of new leaves. The copious fall of succulent corollas weave a cream coloured
 310 Breeding of Underutilized Fruit Crops

carpet on the ground. The stamens are very short and adhere to the inner surface of the
corolla. The pistil is a long, protruding green tongue. The pedicel length, pedicel thickness, bud
length and breadth at flower opening varied from 4.20­5.50 cm, 2.00­3.00 mm, 1.30­1.80 cm
and 1.00­1.20 cm, respectively in different genotypes. Total number of sepals was noted four.
The ovary and style length varied from 4.50­6.25 mm and 25.00­35.00 mm, respectively.

9.3. Anthesis, Stigma Receptivity and Pollen Viability

Variable percentage of anthesis was registered in different genotypes. Peak period of
anthesis was recorded from mid night till morning in all the genotypes. Anthers dehisced
2­4 days before the time of anthesis.
Bhattacharya and Mandal (2012) studied the floral biology of Madhuca indica and
reported that flower anthesis pattern and time (h) is Mid day (5.00­12.00), Pollen anthesis
time (h) is 6.00­15.00 and the type of pollen grain/aperture is 4­5­colporate (i.e. having
distinct furrow and pore). Maximum stigma receptive period is first day after anthesis.
Pollen viability and pollen germination ranged from 90.00­98.50 and 26.50­40.20 per cent
respectively, among the genotypes studied.

9.4. Mode of Pollination

Reddi (1976), Kuruvilla (1989) and Kundu et al.(2012) consider that Madhuca longifolia
(Syn. Madhuca indica), which flowers when tree about to leafless during March­April,
is wind­pollinated (anemophily), but other authors have suggested it is pollinated by fruit
bats which feed on the corollas (Rajan et al., 1999; Elangovan et al., 2000). During the
flowering period (March–April) bats visit Mahua trees during night in large numbers.
According to Nathan et al. (2009) fruit bats are the effective pollinators of M. latifolia
and Indian short­nosed fruit bat (Cynopterus sphinx) and Indian flying fox (Pteropus
giganteus) visit Madhuca latifolia. C. sphinx removed only the fleshy corolla and left
the pistil intact in the trees. While P. giganteus removes a bunch of corollas at a time
in the inflorescence along with a few flowers with pistils. So, it is possible that pollen
grains are transported and deposited on the stigma of intact, untouched and neighbouring
flowers during the detachment and affect pollination. Bhattacharya and Mandal, (2012)
were observed Thrips hawaiiensis and Formicidae to visit the flowers of mahua.

9.4.1. Pollen­pistil interaction

Madhuca indica shows strong self incompatibility reactions. It shows a few characters
of both sporophytic and gametophytic systems (Heslop­Harrison, 1978). It has trinucleate
pollen which needs controlled hydration for germination, characteristic of sporophytic
system. The pollen viability is retained for a longer period (about 5­6 days) which is
gametophytic in nature (Kuruvilla, 1985). The stigma is covered with copious exudates at
receptive stage as in a gametophytic system, but the pollen germination is inhibited on the
stigma surface even though the grains are hydrated as in a sporophytic system. Since the
genetics of the system is unknown, no conclusion can be drawn regarding the nature of
the self incompatibility system in Madhuca indica (Kuruvilla, 1989)
 Mahua (Madhuca longifolia (Koenig) J.F. Macribide) 311

9.5. Seed and Fruit Setting

The rate of seed setting in Mahua under normal environmental conditions is very low.
Kuruvilla (1989) found that usually only one ovule out of eight ovules develops into seed
per fruit and rarely two or upto seven ovules develop to seed. The seed setting rate is
low even in hand pollinations due to lack of distribution of the pollen tubes in all the stylar
canals and competition for stored resources between early fertilized ovules and late
fertilized ovules. The fertilized ovule started its development within three days after
pollination. As flowers are fully mature, the leaf initiation starts and the fruit development
occur simultaneously along with vegetative bud sprouting. The vegetative sink dominates
thereby resulting in the abscission of developing fruit. The matured fruits fall on the
ground in May and July in the North India and August and September in the South India.

10. DIFFERENT SPECIES

The family Sapotaceae consist of 58 genera and about 1,250 species in subtropical and
tropical regions of the world (Swenson et al., 2007a, b). Madhuca is an Indo­malayan
genus represented by about 100 species especially from Western Malaysia to Australia
(Mabberley 2008). The genus name Madhuca is derived from the Sanskrit word Madhu,
meaning honey. Mahua’s basic chromosome number (X=12) is agreed with Mimusops and
other genera of the same family and having diploid chromosome number 2n=24. In India
the genus is represented by 5 species viz. Madhuca longifolia, M. latofolia, M. butyracea,
M. neriifolia and M. bourdillonii (Awasthi et al., 1975; Govaerts et al., 2001) and the
present nomenclature of these species according to The Plant List Version 1 is as follows:
1. Madhuca longifolia (Koenig) J. F. Macb. (Syn. Bassia longifolia J.Konig ex L.):
The two varieties
i. Madhuca longifolia (Koenig) J.F. Macb. var. longi­folia [Syn. Bassia villosa
Wall. ex G.Don, Illipe malabarorum Gras, Illipe malabrorum subsp. alphonsae
Dubard, Vidoricum longifolium (J.König ex L.) Kuntze]
ii. Madhuca longifolia (Koenig) J.F. Macb. var. lati­folia (Roxb.) Cheval. (Syn.
Madhuca indica J.F.Gmel, Madhuca latifolia (Roxb.) J.F.Macbr.
2. Madhuca butyracea (Roxb.) J.F. Macbr. is a synonym of Diploknema butyracea.
(Roxb.) H.J.Lam.: It is locally known as Phulwara and grows chiefly in the sub­
Himalayan region of India and Nepal.
3. Madhuca neriifolia (Moon) H.J.Lam. [Syn. Bassia malabarica Bedd., Bassia
neriifolia Moon, Madhuca malabarica (Bedd.) R.Parker]: It is grows sparsely in
Bombay, south Kanara, Madras and Mysore and often not differentiated from Madhuca
longifolia.
4. Madhuca bourdillonii (Gamble) H.J.Lam. (Syn. Bassia bourdillonii Gamble):
Grows mainly in Mysore and western ghats along with Madhuca longifolia.
312 Breeding of Underutilized Fruit Crops

Prasad and Raveendran (2013) collected one interesting riparian Sapotaceae member
Madhuca insignis (Radlk.) H.J. Lam. (Syn. Bassia insignis Radlk.), a very narrow
endemic and critically endangered species from Kerala. Earlier it is known only from
Dakshina Kannada and Udupi Districts of Karnataka. Four new species i.e. M. chai­
ananii, M. chiangmaiensis, M. klackenbergii and M. smitinandii have also been
isolated from Thailand (Chantaranotha, 1998).

Description of Some Species

10.1. Bassia latifolia
It is a deciduous tree having 12­15 meter height. Bark thick, dark­coloured, cracked, the
inner bark red, milky, trunk short, branches numerous, spreading, forming a thick shady
head. Leaves clustered near the ends of the branches, coriaceous, hard and firm, elliptic
or elliptic­oblong, shortly acuminate, pubescent or tomentose when young, at length glabrous,
base rounded or acute. Petioles long at first pubescent, ultimately glabrous or nearly so.
Stipules long, subulate, densely pubescent, vary caducous. Flowers in dense fascicles near
the ends of the branches, below the terminal leaf­bud. Pedicels long, drooping, fulvous or
rusty­pubescent or tomentose. Calyx long, divided nearly to the base, segments ovate,
subacute, rusty­tomentose, usually 4 (rarely 5), the two outer subvalvate and enclosing the
others. Corolla cream coloured, tube long fleshy, rugose, when dry, lobes7­14 (usually 7­
9), ovate­lanceolate, acute, erect. Stamens 20­30 (usually 24­26), anthers in 3 series,
acuminate, hairy at the back. Ovary hirsute, style long and hairy at the base. Berry long,
fleshy, ovoid, greenish. It flowers in January­ April under different climatic conditions.

10.2. Bassia longifolia

It is commonly found in south India. A large tree, branches numerous, bark thick, dark
brown, scaly, the inner dark red, milky, young parts pinkish­white silky­pubescent. Leaves
thin, clustered near the ends of the branches, linear­lanceolate, acute, glabrous when
mature, much tapered towards the base, petioles long slender, stipules long, linear­subulate,
hairy and caducous. Flowers appearing with the young leaves, in dense clusters near the
ends of the branches below the leaves, pedicels long, glabrous, erect at first, afterwards
more or less drooping. Calyx long divided nearly to the base, densely rusty­pubescent,
segments usually 4, ovate­oblong, acuminate. Corolla long, tube fleshy, in­ flatted, glabrous,
rugose when dried, lobes 6­12, lanceolate, subobtuse, glabrous. Stamens 16­20 in 2 rows
one above the other, subsessile, anthers hairy, cordate at the base, the connective produced
to a point. Ovary densely hairy. Berry oblong, the size of a plum, hirsute while young
ultimately nearly glabrous, yellowish when ripe. Seeds 1­2, compressed, straight on one
side, curved on the other. It flowers November­ January.

10.3. Bassia malabarica

It is a medium sized tree. Bark dark coloured, scaly. Leaves very coriaceous, oblong to
lanceolate, sub­obtuse, glabrous, dark green and shining, base tapering, main nerves 15­
25 pairs, very slender with closely reticulate veins between, petioles long. Flowers crowded
 Mahua (Madhuca longifolia (Koenig) J.F. Macribide) 313

towards the ends of the branches in dense fascicles. Calyx long, divided nearly to the
base, segments 4, ovate, subacute, the 2 outer glabrous or nearly so, the 2 inner hairy.
Corolla long, tube densely rufous­hairy, both inside and outside, slightly shorter than the
lobes. Lobes 6 (rarely 7), oblong, obtuse, fulvous­hairy outside. Stamens 16­18 in 2 rows,
filaments densely rufous­hairy, anther glabrous or with a very small tuft of hairs between
the basal lobes, cordate at the base. Ovary glabrous. Berry oblong­lanceolate, glabrous
while young. It flowers November­ January. It grows in Western Ghats from Kanara to
Travancore and also in the Himalayas.

11. BOTANICAL DESCRIPTION

Mahua is a medium sized to large deciduous tree, usually with a large rounded crown
found up to an altitude of 1,200 meter and of 12 to 15 meter height. Bark grey to black
with cracks, inner bark dark red, milk, trunk short, branches numerous (Behl and Sriwasrawa,
2002). The tree matures from 8 to 15 years and fruits up to 60 years. Leaves are thick,
leathery, pointed at the tip and clustered at the end of branches. It exudes a milky sap
when broken. Young leaves pinkish and wooly underneath. Flowers are small and fleshy,
dull or pale white in colour and in define fascicles near end of branches. Corolla tubular,
freshly, pale yellow aromatic and caduceus (Variers, 1995). Fruits are green at maturity
and turn pinkish yellow when ripe. Botanically fruit is fleshy berry. Fruits are 2­6 cm long,
ovoid, fleshy and having 1­4 seeds coloured brown to black. Seed contains two kernels.
Seed of Mahua is highly sensitive to desiccation and freezing, indicating recalcitrant nature
of seed. Edible part is Mesocarp (flesh) of fruit

12. CROP IMPROVEMENT METHODS

Its breeding work is carried out by National Bureau of Plant Genetic Resources (NBPGR), New
Delhi; Central Institute for Subtropical Horticulture (CISH), Lucknow; Central Arid Zone Research
Institute (CAZRI), Jodhpur; The National Oilseeds and Vegetable Oils Development (NOVOD)
Board, Gurgaon; Central Institute for Arid Horticulture (CIAH), Bikaner; Tropical Forest Research
Institute (TFRI), Jabalpur; Narendra Deva University of Agriculture & Technology, Kumarganj,
Faizabad and Chaudhary Charan Singh Haryana Agricultural University, Hisar etc.

12.1. Introduction
Collection, introduction and evaluation of mahua variability was undertaken by CIAH, Bikaner
after making survey of the Panchmahals district and adjoining areas and thirty­five genotypes
were selected and evaluated for flowering, fruiting and fruit quality attributes. Highest
number of fruits per cluster (14.00) was found in collection No.2. Corolla (Mahua flowers)
of collection No.5 had maximum weight (2.5g) and Juice (68.00%). TSS (26.00%) and total
sugar (22.10%) was also found to be highest in the same genotype. Ripening took place in
the month of May and June in different genotypes. The weight of mahua fruits varied
from12.50 to 40.00 g and seed weight from 5.67 to 22.50 g. Kernel weight ranged from
4.20 to 16.00 g. TSS per cent of fruits varied 12.00­16.30, total sugar 10.50­14.00 per cent
and vitamin C content 42.00­63.50 mg/100g. On the basis of overall performance, collection
no 2, 5, 8, 10 and 14 were found to be promising (Anonymous, 2004).
 314 Breeding of Underutilized Fruit Crops

Germplasm collection and evaluation is carried out by many institutes which are
presented in Table 2 and 3.
Abraham et al. (2010) carried out exploration and collected mahua variability from
Tamil Nadu and the Union Territory of Pondicherry. This resulted in the collection of 54
accessions. Four unique accessions IC554527, IC554535, IC554553 and IC554554 were
identified. Chemical evaluation of 54 accessions for kernel oil content revealed a range
of variation from 44.43 to 61.50%. Three accessions viz., IC556617 with 61.50, IC556632
with 60.80 and IC556632 with 60.55% of kernel oil are superior to the rest and may be
utilized for selection of promising material for extraction of oil.
Variability in oil content and seed weight of 37 accessions of mahua (Madhuca longifolia
Koenig) J.F. Macribide collected from different part of Tamil Nadu, India were assessed
by Yadav et al. (2011b) and found the highest oil content in accessions IC556617 (61.5%)
and was followed by IC554529 (58.7%). Out of the total 37 accessions, 14 accessions were
having more than 55 per cent kernel oil content, which is a desirable trait for industrial use
of this tree. These accessions can be used as a rich source of fat for industrial purposes.
Variability in oil content, fatty acid profile and biodiesel traits of mahua presented in the
Table 4. These thirty seven accessions were grouped in three clusters; cluster I with 28
accessions, cluster II with three accessions, namely, IC555369, IC555370 and IC555371 and
Cluster III with six accessions, namely, IC554546, IC554553, IC554547, IC554526, 554535
and 554551. The accessions in cluster II had low palmitic acid and high oleic acid, while
cluster III had high palmitic acid and low oleic acid content. So they suggested that hybridization
between accessions of cluster II and III will result in wide spectrum of variability in
subsequent generations for medicinal, edible applications and biodiesel production.

Table 2. Variability in oil content, fatty acid profile and biodiesel traits of mahua.

Parameter Range Mean

Kernel oil (%) 44.43­61.50 54.37
Palmitic acid (%) 11.65­25.86 19.56
Stearic acid (%) 19.11­32.16 24.78
Oleic acid (%) 32.91­48.65 42.87
Linoleic acid (%) 9.36­15.42 12.64
O/L ratio 2.39­4.48 3.43
Total saturated fatty acid (%) 39.33­52.75 44.34
Total unsaturated fatty acid (%) 46.66­60.65 55.52
Saturation ratio 0.64­1.13 0.80
Saponification value (SV) 198.34­202.78 201.76
Iodine Value (IV) 52.09­68.59 61.46
Cetane number (CN) 58.0­61.58 59.52

Source: Yadav et al. (2011a)

 Mahua (Madhuca longifolia (Koenig) J.F. Macribide) 315

Table 3. Mahua germplasm collected at various Institutes/University in India.

Sr.No Institutes/ Germplasm Source

University collected
1 CCS HAU, Hisar 86 (http://www.hau.ernet.in/coa/forestry.htm)
2. TFRI, Jabalpur 53 (http://tfri.icfre.gov.in/
Research%20highlights.htm)
3. NBPGR, Regional 53 Anonymous (2008)
Station, Thrissur
4. CISH, Lucknow 30 Anonymous (2011)

Table 4. Some of the interesting collections made by NBPGR regional station Thrissur
(Annonymous, 2008)

Botanical name Accs. No. Speciality District State

Madhuca longifolia IC554527 More than one seed, Krishnagiri Tamil Nadu
extending up to 5
seeds per fruit.
IC554535 Profusely bearing with Vilupuram Tamil Nadu
bold seeds
IC554553 Curved seeds and sharp
Curved seeds and sharp Krishnagiri Tamil Nadu
IC554554 Sharp, long pointed beak Salem Tamil Nadu
IC555358 Profuse bearing and more Salem Tamil Nadu
than 10 fruits per bunch
Madhuca indica IC555367 Short, thick, bitter and Vilupuram Tamil Nadu
hairy fruit; the beak of
which is protruding and
the seed is straight
IC555372 ­ do ­ Dindugal Tamil Nadu

According to Jha et al. (2006) alteration of the fatty acid composition was directed
predominantly towards palmitate and to a lesser extent myristate and oleate due to acyl chain
termination activity of plant thioesterase in bacteria. Thus, this new MbFatB gene (gene
isolated from Madhuca butyracea cloned a cDNA of fatty acyl­acyl carrier protein thioesterase)
can be used in future for transgenic development of oil­seed Brassica, that expresses
predominantly oleoyl­ACP thioesterase (FatA) in its seed tissue and has high amount of
unwanted erucic acid in edible oil in order to alter the fatty acid profile in a desirable way.
Mahua germplasm (TNML 1 to TNML 33) were collected from various places of
Tamilnadu and maintained at the Forest College and Research Institute, Mettupalayam,
Tamilnadu, India.
Twenty four promising accessions have been identified, collected, multiplied and
established in field gene bank for evaluation of growth and fruiting pattern at CISH,
316 Breeding of Underutilized Fruit Crops

Lucknow. The flowers from these selected accessions were analyzed for flower weight
(1.10 to 2.63g), flower length (1.53 to 2.60cm), diameter (1.46 to 1.90cm) and TSS (14.53
to 26.0°B). The juice content in the accessions varied was from 44.25 to 64.90 per cent.
On the basis of these characters CISH M­8 was found to be superior type having
maximum flower weight (2.63g) and TSS 23.26°B with juice content 64.90 per cent
(Anonymous, 2011). Further, nutraceutical value with respect to total antioxidant (44.48
mg AEAC/g) and phenol (9.53 mg/g) content maximum recorded in accession CISH M­
4 while total caroteniod (18.69 ug/g) in CISH M­1 (Singh et al., 2012).
ND University of Agriculture and Technology, Faizabad has raised about 500 Mahua
seedling plants for transplanting. The sucrose content of 3.7% (NDMC­9) to 4.8% (NDMC­
7) and range of TSS 17% (NDMC ­10) to 24% (NDMC ­3) was recorded in mahua
germplasm flowers (Anonymous, 2012).
Wani and Ahmad (2013) estimated the pattern of field environment genetic variation
for twenty genotypes of Madhuca indica Gmel distributed over different sites/locations
of district Allahabad and adjoining areas of Uttar Pradesh, India. They found genotypes
S20, S9, S12 and S19 showed better germination and seedling growth performance for
majority of the characters as compared to the rest of the genotypes and are recommended
for further genetic improvement programme in this species.

12.2. Selection
It is highly heterozygous, cross­pollinated fruit crop and as such seedlings exhibit a wide
range of variations, which aids in the selection of the superior desirable genotypes. Due
to cross pollination and predomination of seed propagation over a long period of time, it
gives immense opportunity to locate elite trees having positive horticultural traits. Wide
variations were observed in sweetness, acidity, size, shape and bearing habits in Mahua
under Uttar Pradesh and Gujarat conditions (Singh et al., 1999 and Anonymous, 2004).
It is important to study the variability existing in the species and to select superior
genotypes for adoptability, fast growth, precociousness, oil content, sugar content, flower
size and disease resistance. It is desirable to screen the naturally available genetic variation
so as to ensure that only the best material is utilized for maximum productivity and for
further breeding work. Superior genotypes are then propagated clonally or by seed into
special seed production areas or orchards where open or controlled pollination provide
seed for planting and if needed, for further selection, testing and breeding (Burley, 1980).
Improvement was made through selection. Elite genotypes were earmarked among
the existing population of mahua based on the horticultural traits and evaluated under field
conditions. Generally, the trees of B. latifolia, B. longifolia are found in the North and
South parts of the country, which bear flowers containing a large amount of sugar and
seeds rich in quality oil.
 Mahua (Madhuca longifolia (Koenig) J.F. Macribide) 317

Promising Genotypes Developed Through Selection

Generally, the trees of B. latifolia, B. longifolia are found in the North and South parts
of the country, which bear flowers containing a large amount of sugar and seeds rich in
quality oil. Though a large variability occurs in its fruits but still there is no improved variety
of mahua for an organized orcharding. Recently some selections have been made at
N.D. University of Agriculture and Technology, Faizabad in Uttar Pradesh. These are
NM 2, NM 4, NM 7, and NM 8 (Singh, 1998).

12.2.1. NM–2
Flowering is taken place in the first week of April. Average number of flowers / fascicles
weight of fresh flower, yield of flower (dry) were recorded 66, 2.8g, and 9.9 kg, respectively
from 7­year­old grafted plants. Fresh flower had 25.50 % total sugar, 19.60% reducing
sugar, 5.6 % non­ reducing sugar, 60.90 mg/ 100 g Vitamin C, 650.70 IU Vitamin A and
0.60 % mineral content. Total sugar content of dry flower was 58.70 % with reducing
sugar 55.60 and non­reducing sugar 3.0 %. Mineral content of dry flower was found to
be 2.20 per cent. Ripening of fruit took place by second week of June. Average fruit
weight 16.80g, husk content 58.80 %, total sugar13.60 %, reducing sugar 7.8 %, non­
reducing sugar 5.80 %, vitamin C 51.30 mg/ 100 g, vitamin A 586.70 IU and mineral
content 1.90 % was recorded. Average seed weight and kernel weight was found to be
7.2 g and 5.50 g respectively. Oil content, 47.40 %, free fatty acid 0.53 %, protein content
23.60 % and mineral content 5.70 % was recorded. Unsaturated fatty acid 53.80 % with
40.30 oleic and 13.50 % linoleic and where as saturated fatty acid was recorded 45.40
% (palmitic 24.30, stearic 20.20 and archidic 0.90 %).

12.2.2. NM–4
Flowering is taken place in the fourth week of April. Average number of flowers /
fascicles 54.6, weight of fresh flower 2.6g and yield of flower (dry) of 7­year­old grafted
plant is 8.5 kg. Fresh flower recorded 24.50 % total sugar, 19.60% reducing sugar, 4.80
% non­ reducing sugar, 49.20 mg/ 100 g Vitamin C, 528.20 IU Vitamin A and 0.67 %
mineral content. Total sugar content of dry flower was 57.30 % with reducing sugar 53.80
and non­reducing sugar 3.30 %. Mineral content of dry flower was found to be 2.30 per
cent. Ripening of fruit started by third week of June. Average fruit weight 19.50 g, husk
content 65.30 %, total sugar 11.20 %, reducing sugar 8.20 %, non­ reducing sugar 3.20
%, vitamin C 55.80 mg/ 100 g, vitamin A 860 IU and mineral content 2.1 % was recorded.
Average seed weight and kernel weight were found to be 6.00 g and 4.70 g, respectively.
Oil content 45.20 per cent, free fatty acid 0.63 per cent, protein content 25.70 per cent
and mineral content 4.40 per cent were recorded. Unsaturated fatty acid 50.60 % with
oleic 38.00 and linoleic 12.60 % and saturated fatty acid 48.50 % (palmitic 19.90, stearic
27.00 and archidic 1.60 %).
318 Breeding of Underutilized Fruit Crops

12.2.3. NM–7
Flowering is taken place in the second week of April. Average number of flowers /
fascicles 64.30, weight of fresh flower 2.50g and yield of flower (dry) of 7­year­old
grafted plant is 10.00 kg. Fresh flower recorded 22.70 % total sugar, 19.50% reducing
sugar, 3.00 % non­ reducing sugar, 48.50 mg/ 100 g Vitamin C, 564.50 IU Vitamin A and
0.82 % mineral content. Total sugar content of dry flower was 58.40 % with reducing
sugar 56.60 and non­reducing sugar 1.50 % while mineral content of dry flower was
found to be 3.00 per cent. Ripening of fruit took place by third week of June. Average
fruit weight 21.8 g, husk content 55.00 %, total sugar 10.10 %, reducing sugar 8.00 %,
non­ reducing sugar 2.30 %, vitamin C 59.20 mg/ 100 g, vitamin A 620.70 IU and mineral
content 2.4 % was recorded. Average seed weight and kernel weight was found to be
9.50 g and 7.20 g, respectively. Oil content 46.70 per cent, free fatty acid 0.55 %, protein
content 22.40 % and mineral content 4.80 % was recorded. Unsaturated fatty acid
49.20% with oleic 34.40 % and linoleic 14.90 % and saturated fatty acid 47.90 % with
palmitic 20.80, stearic 26.90 and archidic 1.20 % was recorded.
Singh and Singh (2005) studied the genetic diversity in mahua and identify twenty
promising germplasm and grouped them early, mid and late based on flowering time. Early
flowering was observed in the first week of March in MH 1, MH 4, MH 5 while late
in the second week of April in MH 6. They found MH 2 and MH 4 as promising
genotype.
The genotype MH­2 having highest flowers (45) and fruits (8) per fascicle. Dry
flower yield and fruit yield varied from 30 to 46 kg/plant and 35 kg to 82 kg/plant,
respectively being both found highest in MH 2. Maximum fruit weight (34.50g), seed
weight (14.50g), kernel oil (46.50%), minerals (4.90%) and protein content (24%) were
also found in MH 2. MH­ 4 having the highest total soluble solids (27.80%), total sugar
(24.24%) and vitamin C content (64.00 mg/100g). These genotypes would be exploited
as potential parents to develop high yielding stable genotypes having positive horticultural
traits.
Description of some promising genotypes evaluated at CHES, Godhra (Singh
et al., 2008a)
The study revealed that there was a wide variation among the genotypes. With respect
to all traits studied genotypes, MH 32, MH 34, MH 35, MH 26, MH 27, MH 23 and MH
33 were found to be promising and would be exploited as potential parents to develop high
yielding stable genotypes.

12.2.4. MH–10
Peak period of flowering is the second week of March. Flowers recorded 2.29 g weight,
65.00 % juice and 26.37 % TSS. It ripens in second week of May and recorded 30.50g
fruit, 13.50g seed and 11.00g kernel weight.
 Mahua (Madhuca longifolia (Koenig) J.F. Macribide) 319

12.2.5. MH–14
Peak period of flowering is the second week of March. Flowers recorded 2.24 g weight,
66.00 % juice and 25.00 % TSS. It ripens in the second week of May and recorded 29.00
g fruit, 12.70g seed and 9.53g kernel weight.

12.2.6. MH–32
Singh et al. (2005) found promising this genotype in respect to maximum number of
flowers and fruits per fascicle, highest dry flower yield (27–48 kg/plant), fruit yield (98.00
kg/plant), total soluble solids (26.20%), total sugar (23.78%) and vitamin C content (62.60
mg/100g) of flowers and fruits.

12.3. Hybridization
12.3.1. Crossing Technique
In this method, flower buds of the cultivars should be emasculated a day prior to anthesis
and covered with butter paper bags. Next morning, emasculated flowers should be pollinated
with the pollen of desired cultivars, after pollination, it should be covered again. These
bags should be removed after ensuring that the fruit set is taken place.

13. FUTURE THRUST

Breeding work with respect to horticultural traits is limited due to limited interest in this
crop by horticulturists. There is vast scope for the improvement of scion and rootstock
in this crop. The dwarf scion with precocious bearing may helpful for closer planting
leading to more number of plants per unit area and efficient cultural practices. Another
area in scion improvement is through increasing the shelf life of flowers (Corolla) which
is economic part of plant and deteriorate very quickly. Under high temperature (<400C)
during summer months (April to June) mahula flowers suffer post­harvest losses. Spoilage
is the most significant form of wastage that accounts for 20–25 per cent of post­harvest
losses in mahua flowers, which become unsuitable for the liquor distillation units and as
cattle feed (Behera et al., 2011; Patel et al., 2011).
Biotechnological tools must be utilized for fasten the breeding programme. Seedlings
must be evaluated as rootstock for better encore, dwarfness and adaptability to different
climate.
Studies on floral biology, inter and intra specific/generic hybridization, documentation
of floral biology and crossing techniques, plantation of selected plants for ex situ
conservation and screening trees for higher oil content, sugar content, high flower and
fruit yield are some of the other breeding objectives which should be carried out for the
improvement of mahua.
320 Breeding of Underutilized Fruit Crops

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 MAHUA (Madhuca longifolia (Koenig) J.F. Macribide)

Fig. 1: Flower developmental stages in Bassia latifolia.

(a) Inflorescence representing different stages of flower development, (b) Detached

fleshy corollas, (c) Growth stages of flower

Flowering
MAHUA (Madhuca longifolia (Koenig) J.F. Macribide)

Fruiting

Fruits of MH­10

Fruits of MH­14
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