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Jackfruit (Artocarpus heterophyllus) and Breadfruit (A. altilis): Phytochemistry, Pharmacology, Commercial Uses and Perspectives for Human Nourishment

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The Artocarpus J. R. & G. Forster genus is comprised of about 50 species. Artocarpus is derived from the Greek word artos, meaning bread while karpos means fruit. There are two species that are widely distributed in tropical regions, Artocarpus heterophyllus Lam., known as jackfruit, and Artocarpus altilis (Parkinson) Fosberg, known as breadfruit, both in the Moraceae or mulberry family. Both of these Artocarpus species have medicinal properties and biological activities that are derived from almost every part of the tree, fruit, seed, wood, bark, leaves and sap. This review examines the limited work that has been conducted on the biology and biotechnology of these two Artocarpus species with the hope that this knowledge may spur further basic and applied research.
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Journal of Tropical Biology and Conservation 15: 6180, 2018 ISSN 1823-3902
E-ISSN 2550-1909
Received 30 May 2017
Reviewed 04 October 2017
Accepted 13 October 2017
Published 15 October 2018
Jackfruit (Artocarpus heterophyllus) and Breadfruit (A.
altilis): Phytochemistry, Pharmacology, Commercial Uses
and Perspectives for Human Nourishment
Reza Raihandhany1, Adhityo Wicaksono2, Jaime A. Teixeira da Silva3*
1Department of Forestry Engineering, School of Life Science and Technology,
Bandung Institute of Technology (Jatinangor campus), Sumedang, West Java, 45363
Indonesia
2Laboratory of Paper Coating and Converting, Centre for Functional Material, Åbo
Akademi University, Porthaninkatu 3, 20500 Turku, Finland
3P. O. Box 7, Miki-cho post office, Ikenobe 3011-2, Kagawa-ken, 761-0799, Japan
*Corresponding author: jaimetex@yahoo.com
Abstract
The Artocarpus J. R. & G. Forster genus is comprised of about 50 species.
Artocarpus is derived from the Greek word artos, meaning bread while karpos
means fruit. There are two species that are widely distributed in tropical regions,
Artocarpus heterophyllus Lam., known as jackfruit, and Artocarpus altilis
(Parkinson) Fosberg, known as breadfruit, both in the Moraceae or mulberry family.
Both of these Artocarpus species have medicinal properties and biological activities
that are derived from almost every part of the tree, fruit, seed, wood, bark,
leaves and sap. This review examines the limited work that has been conducted on
the biology and biotechnology of these two Artocarpus species with the hope that
this knowledge may spur further basic and applied research.
Keywords: fruit, medicine, Moraceae, secondary metabolites, tropical tree
Introduction
The genus Artocarpus (Moraceae), which contains food-producing plants that
are spread throughout tropical and subtropical regions of the world, consists of
about 50 species (Motley, 2014), but The Plant List (2018) lists 193 accepted
names for Artocarpus, although many of them are synonymous and unresolved
species. The word Artocarpus is a compilation of two Greek words, artos,
which means bread, and karpos, which means fruit (Jones et al., 2011).
Research Article
62 Raihandhany et al.
The species epithet of jackfruit, heterophyllus, is a compilation of two Greek
words, hetero, meaning different, and phyllus, which means leaf (Gupta,
2011). This implies existing variation in the shape and size of the leaves.
Jackfruit, which typically grows in the form of a tree, provides edible fruit and
medically potential secondary metabolites, is a source of timber, and has been
cultivated throughout China, Sri Lanka, India and Southeast Asia, but is also
found in Africa, the Caribbean islands, Brazil, Suriname and tropical parts of
Australia (Thaman & Ali, 1993). Jackfruit is known as nangka in Indonesia and
has various ethnobotanical properties that derive from its ripe fruit which
serve as ingredients for local sweets such as kolak and dodol in Java, young
fruit is consumed as a vegetable, and its leaves are used as cattle feed (Lim,
2012). Ash of leaves can be used to treat wounds and serve as medication to
treat ulcers (Gogte, 2000). Jackfruit timber is a good wood for furniture,
construction material, and musical instruments since it resists bacterial, fungal
and termite attacks (Orwa et al., 2009).
The methanolic extract of stem, root bark and heartwood, leaves, fruit, and
seed have multiple antibacterial compounds (Khan et al., 2003). One of those
compounds, artocarpin, is used as an antitermite agent (Shibutani et al.,
2006). The basal part of the fruit, which is fleshy, fibrous and rich in sugar,
provides a good natural source of carbohydrates and minerals such as calcium,
iron, magnesium, carboxylic acids, and vitamins A, C and E, primarily ascorbic
acid and thiamine (Rahman et al., 1999). Mature seed are edible when dried or
after cooking by boiling and roasting. Fresh mature seed contain 25 IU/100 g of
vitamin A, 4.3-6.6 g/100 g of protein, 23-25 g/100 g of calcium, 80-126 mg/100
g of phosphorus, and 10-17 mg/100 g of ascorbic acid (Acedo, 1992). In fresh
(raw) fruit, there are 23.25 g/100 g of carbohydrate, 24 mg/100 g of calcium,
0.23 mg/100 g of iron, 29 mg/100 g of magnesium, 110 IU/100 g of vitamin A,
13.7 mg/100 g of vitamin C (ascorbic acid), and 0.34 mg/100 g of vitamin E (α-
tocopherol) (USDA, 2016). The latex which also has anti-syphilitic and
vermifuge properties, contains 71.8% resin, 63.3% of which are yellow fluavilles
and 8.5% white albanes that are useful for varnishes (Rao et al., 2014). A study
conducted in New Delhi and Kerala, India by Suba Rao (1983) showed that
jackfruit is symbiotically asociated with Azotobacter and Beijerinckia, 35 and 4
× 104/g soil, respectively at pH 6.8-7.5, and 14 and 18 × 104/g soil, respectively
at pH 3.5-5.5. According to Prakash et al. (2009), a hot water extract of
jackfruit leaves when consumed orally by humans at 20 g/kg of the patient’s
weight, improves glucose tolerance for mature-onset diabetic patients, while
the crude methanolic extract of jackfruit parts (stem, root heartwood, bark,
leaves, fruits and seeds) and their subsequent partitioning with petrol,
Jackfruit and breadfruit: a review 63
dichloromethane, ethyl acetate and butanol gave fractions that exhibited
broad spectrum antibacterial activity, the most active fraction being the
butanolic extract of fruits and root bark. An extract from jackfruit shoots also
revealed nematicidal activity against Rotylenchulus reniformis,
Tylenchorynchus brassicae, Tylenchus filiformis and Meloidogyne incognita
(Sharma & Trivedi, 1995 cit. Prakash et al., 2009).
The species epithet of A. altilis, the word altilis itself is a Greek word that
means fat, refers to the fruit shape (Small, 2011). Breadfruit, also a source of
food, was first cultivated in the Western Pacific about 3,000 years ago and is
native to the eastern part of Indonesia, New Guinea, Malaysia and the
Philippines (Orwa et al., 2009). The migration of Polynesians to South and
South America, Africa (Senegal, Ghana, and Liberia), India, Maldives and Sri
Lanka contributed to the distribution of breadfruit (Deivanai & Bhore, 2010).
The breadfruit tree is often employed in a mixed cropping system with yams,
banana, black pepper and coffee, although details of these cropping systems
are lacking (Ragone, 1997). The fruit of ripe breadfruit can be eaten fresh or
cooked by steaming, roasting and frying (Ragone, 1997). Leaves and the non-
edible part of fruit can be used as cattle feed while tree bark can also serve as
feed for horses (Morton, 1987). In Samoa and several Pacific Islands, bark is
used to cure headaches, in Java and Malaya the toasted flower is used to treat
toothache, while in the Bahamas, leaves of A. altilis are used to relieve
headaches (Kuete et al., 2011). In Indonesia, the methanolic or
dichloromethane extracts of leaves have medicinal properties and are used to
cure liver cirrhosis, hypertension and diabetes (Kasahara & Hemmi, 1988;
Arung et al., 2009). Similar to jackfruit, breadfruit trunk wood is good for
construction and furniture, and its sap can be used to trap birds and houseflies
or to treat human skin and fungal diseases (Ragone, 1997).
Jackfruit and breadfruit are tropical fruits with potential beneficial uses as
food, timber and ethnomedicines, but this requires scientific testing. This
paper, in a bid to expand research of these trees, and expand their sustainable
use and production through biotechnological interventions, highlights their
basic biology such as morphology, medicinal properties and propagation (both
in classical and biotechnological approaches). In this paper, we highlight
research that has been conducted on two species, A. heterophyllus Lam. (syn:
A. integrifolia Linn.) or jackfruit, and A. altilis (Parkinson) Fosberg (syn: A.
communis J.R. Forst & G. Forst; A. incisus (Thunb.) L.f.), or breadfruit.
64 Raihandhany et al.
Morphology
Jackfruit is an evergreen tree 8-25 m in height and with a trunk diameter of
30-80 cm that can live up to 100 years. Young trees grow with a conical or
pyramidal canopy shape that turns into a dome-shaped canopy as the plant
grows older. Canopy diameter which can reach 10 m, is close to the ground
and provides dense shade (Elevitch & Manner, 2006). Wood of jackfruit is
categorized as medium hardwood with a specific gravity of 0.6-0.7 (Orwa et al.,
2009). When the tree ages, wood turns from yellow to red or brown. Breadfruit
is also an evergreen tree 15-20 m in height and with a 1-2 m diameter trunk
whose bark is smooth, thick and light-grey while wood is golden although, after
exposure to air, it darkens (Ragone, 1997).
Jackfruit inflorescences sprout from a short, thick stalk and emerge from the
lateral side of the main stem and thick branches (Backer & Bakhuizen, 1965).
The male inflorescence forms in the axil of the apical branch with a cylindrical
to conic-ellipsoid shape 2-7 cm in diameter and a 1-5 cm long peduncle with a
tubular calyx that has a two-lobed apex 1-1.5 mm in diameter, pubescent
texture, straight filament and ellipsoid anther while the female inflorescence
has a globose fleshy rachis with a tubular calyx, lobed apex and a one-celled
ovary (Zhou & Gabriel, 2006). Some parts of the male inflorescence are sterile.
As in jackfruit, the breadfruit inflorescence emerges from the apical trunk
(Figure 1).
Figure 1. Jackfruit young fruit (left) and mature fruit (right). White scale bar = 10 cm.
Unpublished figure.
Jackfruit and breadfruit: a review 65
The breadfruit inflorescence has a cylindric-clavate shaped flower with a 3-6
cm long peduncle and globose or ellipsoid inflorescence shape with a diameter
up to 20-30 cm. It has a tubular calyx that is pubescent, has two lobes on its
apical surface and has a lanceolate-shaped lobe while the anthers are elliptic.
Female breadfruit flowers have a tubular calyx, an ovoid ovary with a long
style and two branches on the apex. Each flower consists of a reduced tubular
perianth that covers a single stamen with a two-lobed anther on a thick
filament (Sharma, 1962).
Both jackfruit and breadfruit exude a sticky white latex from the injured parts
of the plant (Rahman & Khanom, 2013), and forms part of the plants’ defense
against herbivory (Agrawal & Konno, 2009). The phyllotaxis (i.e., leaf
arrangement) of jackfruit and breadfruit is distichous or spiral with simple,
leathery leaf blades with a full margin and plants are monoecious (i.e., male
and female flowers on the same tree) with inflorescences growing from the
main branch or trunk (cauliflory) for jackfruit but sprouting from the apex of
the main branch, also where new leaves emerge, and arising from simple,
pseudomonomerous ovaries as in other Moraceae species (Singh, 2016). Both
jackfruit and breadfruit form a single leaf blade that is lobed, but mature
jackfruit leaves become entire and lose their lobes, hence the species epithet,
heterophyllus. The leaves of jackfruit and breadfruit have stipulate leaf types,
with an ovate form for jackfruit and a lanceolate to broadly lanceolate form
for breadfruit. Jackfruit leaves are spirally arranged with an elliptic to obovate
leaf blade, leathery, leaf margins are lobed in seedlings but entire in mature
trees, with pale green on the lower leaf surfaces displaying scattered globose
to ellipsoid resin cells while the axial surface is dark green, smooth and glossy
(Zhou & Gilbert, 2003) with a cuneate, subdecurrent base, firmly coriaceous,
leaf size is 10-20 × 5-10 cm (l × b), the stipule is 1.5-5 cm, and the petiole is 2-
4 cm long (Backer & Bakhuizen, 1965). Breadfruit leaves are also spirally
arranged, elliptic in shape with a broadly cuneate or obtuse base, up to 3-7
lobed along each margin, lobes are oblong, long-acuminate acute, the stipule
is 16-20 cm long, the petiole is 2-4 cm ling, and leaves are 30-100 cm × 25-65
cm (Backer & Bakhuizen, 1965).
Jackfruit and breadfruit have a compound fruit or syncarp that is classified as
a compound false fruit or pseudofruit that forms from the enlargement of the
stigma, and the inflorescence is composed of 1,500-2,000 flowers attached to
the fruit’s axis (Jarret, 1976). The fruit of jackfruit can weigh 4.5-30 Kg and
can reach 30-40 cm in length, with an oblong-cylindrical shape and dark green
coloration when young that turns greenish-yellow or brownish when mature.
66 Raihandhany et al.
The fruit grows and matures on the trunk for 90-180 days (Elevitch & Manner,
2006). Some jackfruit achenes contain multiple fruits, each with a bulk
composed of seed and with a waxy and soft texture, golden-yellow with a
sweet and aromatic aril (Orwa et al., 2009). The fruit of breadfruit is formed
from the fused flower perianth, except for the base (Reeve, 1974), young fruit
is light-green but turns yellowish-green when mature, and as the fruit
develops, perianths fuse, becoming the fleshy edible portion of the fruit
(Ragone, 1977). When sliced, breadfruit has a white flesh composed of dense
perianths (Figure 2).
Jackfruit seed are semi-round, light brown to brown, 2-3 cm in length and 1-
1.5 cm in diameter, wrapped in a whitish seed coat/testa, and a yellow aril
(Figure 3). The seed is recalcitrant and can be stored for up to a month in
humid conditions (Elevitch & Manner, 2006). Adelina et al. (2014) air-dried
seeds for 0 h (control) to 5 h (treatments separated by 1 h) at 28°C and 70%
humidity, noticing that water content was reduced from 75.03% to 22.95%,
seed respiration rate declined from 7.189 mg CO2/kg h to 5.32 mg CO2/kg h,
and seed viability dropped after 14 days of germination from 97.33% to 24.67%.
The seed of breadfruit is brown, round or obovoid in shape with a thin wall 1-2
cm thick with reduced or no endosperm, hence its recalcitrance to storage or
desiccation (Ragone, 1997). Some modern bread breadfruit cultivars are
seedless (Devanai & Bhore, 2010). The male inflorescence of seedless cultivars
produces less viable pollen than fertile, less-seeded cultivars and only few
flowers in the male inflorescence produce and release pollen (Devanai &
Bhore, 2010). In seedless breadfruit cultivars, nectar is only produced in male
flowers but not in female flowers (Ragone, 1997). In general, the loss of
Figure 2. Breadfruit: whole (left) and sliced (right). Scale bar = 5 cm. Unpublished figure.
Jackfruit and breadfruit: a review 67
fertility in breadfruit is caused by triploidy (2n = 3x = ~84) or by sterile diploids
(2n = 2x = 56) that result from hybridization (Ragone, 2001).
Medicinal properties
Artocarpus produces various secondary metabolites and biologically active
compounds, particularly phenolic compounds such as flavonoids (Table 1),
stilbenoids, and arylbenzofurans (Hakim et al., 2006), extracted from leaves,
the stem, fruit and bark, which have ethnomedicinal uses and antibacterial
(Khan et al., 2003), antiviral (Likhitwitayawuid et al., 2005; 2006), antifungal
towards Herpes Simplex Virus (HSV) and Human Immunodeficiency Virus (HIV)
(Jayasinghe et al., 2004; Trindade et al., 2006), antiplatelet (inhibitory of
thromboxane formation) (Weng et al., 2006), antiarthritic (Ngoc et al., 2005),
tryrosinase inhibitory (Arung et al., 2006; Likhitwitayawuid & Sritularak, 2001)
and cytotoxicity properties (Hakim et al., 2006) (reviewed in greater detail by
Figure 3. Mature fruit of jackfruit (A), sliced (B), part of the fruit with arils and the seed
covered with testa (C), and jackfruit seeds with testa (left) and still wrapped with aril
(right). Blue lines indicate the direction of cuts. Scale bar = 5 cm. Unpublished figure.
68 Raihandhany et al.
Jagtap & Bapat, 2010). Jacalin, which is a tetrameric two-chain lectin
extracted from A. heterophyllus, has strong mitogenic activity against human
CD4+ T lymphocytes, serving as an immunobiological diagnosis agent for HIV-1
patients (Kabir, 1998).
Jackfruit contains various components used for medical benefits. Some
flavonoids (Table 2) are used as antinflammatory agents (Wei et al., 2005).
Fang et al. (2008) extracted three phenolic compounds from the ethyl acetate
fraction of jackfruit fruit: artocarpesin (5,7,2’,4’–tetrahydroxy6–β–methylbut
3–enyl flavone), norartocarpetin (5,7,2’4’-tetrahydroxyflavone), and
oxyresveratrol (trans-2,4,3’,5’tetrahydroxystilbene). All three compounds
showed a potent anti-inflammatory property after inhibiting
lipopolysaccharide-activated RAW 264.7 murine macrophage cells. Other
Table 1. Typical flavonoids, modified flavonoids, and flavonoid-derived xanthones found
in Artocarpus (Hakim et al., 2006)
Compound class
Flavonoids
Modified flavonoids
Flavonoid-derived xanthones
Table 2. Flavonoids with anti-inflammatory properties (Wei et al., 2005)
Flavonoid Compounds
Cycloartomunin
Cyclomorusin
Dihydrocycloartomunin
Dihydroisocycloartomunin
Cudraflavone A
Cyclocommunin
Artomunoxanthone
Cycloheterohyllin
Artonin A and B
Artocarpanone A
Heteroflavone A, B, and C
Jackfruit and breadfruit: a review 69
compounds, cycloheterophyllin and artonins A and B, showed antioxidant
properties as they inhibited iron-induced lipid peroxidation after exposure to
oxygen radicals in more than 60% of a rat brain homogenate after the addition
of 1 μM of each of the three compounds and in more than 80% when 3 μM was
used (Ko et al., 1998). A chitin-binding lectin, jackin, which was purified from
a saline crude extract of jackfruit seed, displayed anti-fungal properties,
inhibiting the growth of Fusarium moniliforme and Aspergillus niger cultures
(2.25 mg/ml, but no effect for A. niger at 4.5 mg/ml) and induced
hemagglutination against human and rabbit erythrocytes (with at least 0.15
mg/ml) (Trindade et al., 2006). Jacalin, a 65 kDA two-chain lectin, has
potential as an immunomodulatory agent, having shown mitogenicity against
human CD4+ T lymphocytes when added at 100 μg/ml (Blasco et al., 1995). The
addition of 10, 20, 30, and 40 μg/ml of jackfruit lectin displayed in vitro
inhibitory activity against herpes simplex virus type HSV-2, varicellazoster virus
(VSZ), and cytomegalovirus (CMV) via a cytopathic effect, and inhibited HIV-1
infection in vitro by preventing the binding of the virus to host cells (Wetprasit
et al., 2000; Swami et al., 2012).
The methanolic and ethyl acetate extracts from breadfruit fruit contain
steroids, phenolics and flavonoids that can inhibit the growth of human
pathogenic bacteria like Enterococcus faecalis, Staphylococcus aureus,
Streptococcus mutans and Pseudomonas aeruginosa by establishing a defense
mechanism (Pradhan et al., 2013). During a test on mice, the methanolic
extract of breadfruit fruit and leaves (500 μg/ml each) was used to treat
inflammation by lowering the intensity of leukocyte infiltration by preventing
skin tumor growth and angiogenesis induced by carcinogenic chemicals 30
minutes after treatment (Lin et al., 2014). Fruitackin, a lectin isolated from
the saline crude extract of breadfruit seed, induced hemagglutination against
human and rabbit erythrocytes when added at 0.15 mg/ml and exhibits
antifungal activity against Fusarium moniliforme and Aspergilus niger at the
same concentration as used for jackin (2.25 mg/ml, but no effect on A. niger
at 4.5 mg/ml) (Trindade et al., 2006).
Propagation (classical and biotechnological)
Conventional vegetative propagation using cuttings, grafting and rootstocks
have unsuccessfully been used to propagate A. heterophyllus and A. altilis,
thus seed serve as an effective choice to propagate A. heterophyllus (Roy et
al., 1993). In vitro culture is an effective solution to cultivate and mass-
produce both species. Roy et al. (1993) first washed adventitious shoot buds in
100 ml of 0.7% polyvinylpyrrolidone (PVP) with 2% sucrose, shook them at 100
70 Raihandhany et al.
rpm for 3 minutes then washed buds with tap water to remove PVP. Buds were
disinfected in 0.2% HgCl2 for 5 minutes then rinsed with sterile double-distilled
water (SDW) for 3 minutes and this procedure was repeated 3-5 times. Buds
cultured on Difco bacto-agar-solidified Murashige & Skoog (1962) (MS) basal
medium supplemented with 8.88 μM 6-benzyladenine (BA) and 2.68 μM α-
naphthaleneacetic acid (NAA) induced 10 shoots/explant after the 7th
subculture. Shoots were elongated on MS medium with 4.44 μM BA, 0.54 μM
NAA and 10% (v/v) coconut milk. Shoots were rooted in vitro on half-strength
MS medium with 5.37 μM NAA and 4.92 μM indole-3-butyric acid (IBA), 80% of
shoots being able to root. Plantlets were transplanted into earthen pots
containing sterile sand, soil and humus (1:2:1, v/v/v), and 75% survived after
30 days.
Amin & Jaiswal (1993) used 10-20 days old terminal buds from an A.
heterophyllus trunk from a 30-50 year-old tree grown from seeds. Stems were
washed in running tap water, treated with 1% (v/v) Cevalon® (an antiseptic and
detergent), disinfected in 0.1% HgCl2 for 5 minutes, then rinsed with SDW 4-5
times. Explants (5-10 mm denuded buds) were prepared by removing the outer
cover of green stipules and excising inner buds encased by creamy-white
stipules before implanting them vertically on growth medium, and placing
cultures at 26±1°C, a 16-h photoperiod (50-70 μmol m-2 s-1), and subculturing
them every 4-5 weeks. MS basal medium with four concentrations (4.5, 9.0,
18.0, and 36.0 μM) of BA and kinetin (Kin) and a combination of BA and Kin
(4.5 μM each) were used to induce shoots while MS with two concentrations of
BA (4.5 μM and 9.0 μM) and BA with Kin (4.5 μM each) were used to multiply
shoots. Roots were successfully induced from shoots with four combinations
(0.5, 5.0, 10.0, and 25.0 μM) each of NAA and IBA, or two combinations (5.0 +
5.0 and 10.0 + 10.0 μM of NAA and IBA). The highest percentage of bud break
resulted from 9.0 μM BA (82 ± 6%) while BA + Kin (4.5 μM each) resulted in 90 ±
7%. The highest number of shoots/explants formed with 4.5 μM BA (3.5±0.6),
or 38±1.1 for BA + Kin (4.5 μM each). Under ex vitro conditions, the survival
percentage of regenerated plantlets was 50%.
A. altilis can be propagated vegetatively in vivo and in vitro. In vivo
vegetative propagation can be achieved by cuttings and air layering of
branches by removing the ring bark, covering the wound with peat moss and
then encapsulating in plastic to induce rooting before being cut and placed on
soil (Deivanai & Bhore, 2010), although details about how long it takes to
achieve each step was not explained. In vitro propagation of A. altilis can be
achieved using shoot tips (Rouse-Miller & Duncan, 2000; Murch et al., 2008).
Jackfruit and breadfruit: a review 71
Rouse-Miller & Duncan (2000) collected shoot tips from a 6-7 year-old tree
during the dry season (December to April in Trinidad-Tobago). Explants with
one or two expanded leaves and 3-6 cm of associated stem were collected and
placed in water (period of time not specified). Expanded leaves and bracts
surrounding the shoot tip were removed and shoots were rinsed in tap water
before cleansing in 70% ethanol for 1 minute. Shoots were reduced to 1 cm,
dipped in 70% ethanol for 30 seconds, 10% household bleach (5.25% available
chlorine) for 10 minutes and rinsed three times in sterile distilled water. The
Rouse-Miller & Duncan (2000) study used Margara (1978) nutrients (Table 3).
For shoot induction, N5K and N15K macronutrients (Margara, 1978), MS
micronutrients and vitamins with 3% sucrose, 0.8% agar and 4.4 μM BA were
necessary. Shoot proliferation required Margara (1978) N30NH4 macronutrients,
MS micronutrients, vitamins, 3% sucrose and 2.2 μM zeatin. Rooting required
N30NH4 macronutrients, vitamins, 2% sucrose, with 0.5, 1.0, 1.5, 2.0, and 2.5
μM IBA. However, IBA alone could not induce roots, and 60% of shoots formed
roots in auxin-free medium (N3ONH4 in Table 3; Margara, 1978). Murch et al.
(2008) used MS or B5 (Gamborg et al., 1968) media with 2.5 g/L gelrite and 3%
sucrose, 2 μM BA and 3 μM Kin to induce shoots in A. altilis within one week
and 1 μM IAA to induce roots.
72 Raihandhany et al.
Table 3. Margara (1978) nutrient lists according to Karla da Silva (2010).
Medium
Macronutrients (mg/L)
KNO3
NaNO3
NH4NO3
Ca(NO3)2•4H2O
CaCl2•2H2O
MgSO4•7H2O
KCl KH2PO4
N5Ca
80
354
292
246
149
136
N30Ca
808
480
1180
246
74.5
136
N30K
1313
480
590
246
74.5
136
N15K*
606
240
354
246
149
136
N15Ca
101
240
944
246
149
136
N45K
1818
85
720
944
246
372.5
136
N5K*
75.8
80
265.5
246
372.5
136
N3ONH4*
606
800
472
246
372.5
136
Medium
Micronutrients (μg/L)
MnCl2
ZnSO4•H2O
H3BO3
KI CuSO4•5H2O
NaMoO4•H2O
FeSO4•7H2O
NaEDTA•2H2O
All
157
500
500
10
100
59
35000
30000
* only the macronutrients were used in the Rouse-Miller and Duncan (2000) study
Jackfruit and breadfruit: a review 73
Molecular advances and future perspectives
Molecular studies of both jackfruit and breadfruit offer promising prospects for
exploiting biotechnology- and industry-derived benefits. Breadfruit molecular
genetics has been studied more than in jackfruit. Studies on the genetic
identification and profiling of breadfruit used microsatellite or short sequence
repeats, identifying around 65 loci for nuclear genomic DNA (Witherup et al.,
2013; De Bellis et al., 2016) or 15 loci for chloroplast genomic DNA (Elliot et al.,
2015). Multi-access identification key software to identify breadfruit cultivars
has been developed from a prototype version on a Lucid 3.3 platform based on
quantitative and qualitative traits (Jones et al., 2013). Amplified fragment
length polymorphism (AFLP) has been used to identify and track the origin of
breadfruit cultivars as linked to the routes of human migration in Oceania
(Zerega et al., 2004), or to assess genetic diversity (Shyamalamma et al.,
2008). Random amplified polymorphic DNA (RAPD) was also used to assess
genetic diversity (Prasad et al., 2014) and fruit cracking in jackfruit (Singh et
al., 2011). Chloroplast and nuclear DNA were used to assess the phylogeny of
60 Moraceae taxa, including the Artocarpus genus (Zerega et al., 2010).
Gibberellin 20-oxidase genes isolated from breadfruit allowed for the
detection of sequence variants, their role in stem elongation after cuttings
were treated with paclobutrazol (a GA inhibitor), and their regulation of
abiotic stress, namely salinity and drought (Zhou & Underhill, 2015, 2016).
Future research needs to identify breadfruit and jackfruit genetic diversity
more precisely while studies on molecular genetics related to metabolic
biosynthetic pathways, for example the elucidation of genes coding for
artocarpatin synthesis, would allow for applications in the pharmaceutical
industry.
Jackfruit and breadfruit are still known locally and may be good sources of
nutrients ranging from carbohydrates to secondary metabolites. These fruits
could be useful germplasm in future plant breeding projects for improving fruit,
such as fortifying stress tolerance. Roy et al. (1993) bred flood-resistance
jackfruit plants in vitro as a way to solve the problem of annual flooding in
Bangladesh. A breeding programme conducted in South Florida aimed to
improve jackfruit aroma, edible percentage, flesh firmness, colour and flavour
(Campbell et al., 2004). A red-fleshed variant of jackfruit exists in India
(International Tropical Fruits Network, 2011). These colour variants can be
used to attract more consumers and thus achieve the maximum benefits of
jackfruit, thus breeding for more colourful fruit flesh could be important. For
the nutraceutical and pharmaceutical industries, future jackfruit breeding for
higher content of specific metabolites can be achieved in a similar way as
“Gama Melon Parfum,a melon cultivar that was developed in Indonesia to
74 Raihandhany et al.
obtain higher yield of sesquiterpenes aimed for perfume production (Maryanto
et al., 2014). Breadfruit colouration is mostly only white, but it has some
shape variants ranging from oval to long fruits (McCormack, 2007). As
breadfruit appears to have potential as a better source of starch used in drug
tablets than cornstarch (Adebayo et al., 2006), a breeding programme to
produce a higher yield of starch in breadfruit could be a good prospect. Similar
prospects for jackfruit could also be applied to breadfruit in future by creating
colour variants for increased appeal or to improve metabolite content for the
food, pharmaceutical and nutraceutical industries. As one example, breadfruit
flour was found to be a good substitute for wheat flour when used as a
composite breadfruit-wheat flour mix for donuts, with a larger ratio of
breadfruit flour resulting in lighter donuts, apparently as a result if its lower
gluten content, although panelists preferred the color, aroma, taste, and
texture of donuts with more wheat flour in the dough (Oke et al., 2018).
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... Jackfruit is an evergreen tropical tree which lives up to 100 years (Raihandhany et al., 2018) and growing up to 8-25 m in height. The male and female inflorescence are present on the same tree due to monoecious pattern. ...
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