Journal of Herbal Medicine 11 (2018) 1–11

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Journal of Herbal Medicine

journal homepage: www.elsevier.com/locate/hermed

Review article

Nutritional and medicinal applications of Moringa oleifera Lam.—Review of T

current status and future possibilities
⁎
Swati Guptaa, Rohit Jaina, Sumita Kachhwahab, S.L. Kotharic,
a
Department of Biosciences, Manipal University Jaipur, India
b
Department of Botany, and Bioinformatics Infrastructure Facility, University of Rajasthan, Jaipur, India
c
Amity Institute of Biotechnology, Amity University Rajasthan, Jaipur, India

A R T I C L E I N F O A B S T R A C T

Keywords: Moringa oleifera Lam. (vern. Drumstick) is a mineral rich, medicinally important tree species of family
Moringa oleifera Moringaceae. It has a wide range of culinary applications and has bioremediation, nutritional and medicinal
Alzheimer’s properties. Conventionally, seeds of the tree have been used as natural coagulants and flocculents in waste water
Cancer treatment. Its leaves and bark act as biosorbent for remediation of heavy metals and dyes. The tree provides
Moringa gum
remedies for a range of diseases and disorders by dint of its unique combination of numerous phytochemicals.
Antioxidant
Gum exudates of the tree are of immense medicinal importance due to their applications in biodegradable drug
delivery systems and in treatment of asthma, dysentery and intestinal cancer. The multidimensional utilities of
M. oleifera may cause overexploitation of this tree, posing danger to the existing natural variability in the near
future. Therefore, there is a need for conservation of the species for ethnobotanical, pharmacological, nu-
traceutical and biodiversity purposes. Development of tissue culture propagation methods will assist in pre-
serving some of the germplasm of the species. In the present review, consolidated analysis of the role of bio-
technology in conservation and genetic enhancement of nutritional, medicinal and commercial value of the tree
has been placed in perspective, together with an up to date review on phytochemical analysis of the plant, and its
utility has been discussed to invite the attention of the scientific community to further consider the study of this
miracle tree species and its nutritional and pharmaceutical properties.

1. Introduction Moringa oleifera Lam. (syn. M. pterygosperma Gaertn.) is a minerals

Plants provide the richest resource of natural compounds, which are
used directly or indirectly for a wide range of applications for the
wellbeing of human population and domestic animals (Khanuja, 2012).
These compounds are primarily phytomolecules synthesized through
primary and secondary metabolic pathways in plants (Arora et al.,
2013).
The dependence of humans on plants for their basic requirements
like food, medicine, clothes and shelter is as old as mankind itself
(Goyal et al., 2007), and still in the modern age the majority of com-
mercial products including pharmaceutical and healthcare, food and
beverages, textiles, cosmetics and aromas are obtained from plants
(Khanuja, 2012). Therefore, plants are and will remain economically,
industrially, environmentally, spiritually, historically, and aesthetically
important for survival, sustenance and prosperity of life on the Earth
(Arora et al., 2013).
and vitamin rich, nutritious and medicinally important tree species of
the Hindustan center of crop origin, belonging to the family
Moringaceae, and commonly called Drumstick, Ben Oil or Horseradish
tree (Arora et al., 2013; Ramachandran et al., 1980).
Moringaceae is a monogeneric family including the single genus
Moringa with 10–12 species. The plant M. oleifera is characterized by
tripinnate leaves, yellow or white petioles, hanging 3-sided pods and
whitish grey corky bark (Fig. 1a–c). Other characteristics include bi-
sexual, stalked, white or creamy axillary flowers, globular winged
seeds, pendulous ribbed capsules and soft, fissured tuberous taproots
(Bhandari, 1995; Ramachandran et al., 1980; Seshadri and Nambiar,
2003). The tree also exudes gum through lysigenous traumatic gum
ducts developed in the bark (Bhandari, 1995; Subrahmanyam and Shah,
1988). The shape and color of the gum exudates varies from stalactite
pieces to tears and yellow to reddish brown or black (Fig. 1d) in color
(Bhandari, 1995; Morton, 1991; Panda et al., 2006; Ramachandran

Abbreviations: Kn, Kinetin; BAP, 6-benzylaminopurine; IBA, Indole-3-butyric acid; IAA, Indole-3-acetic acid; NAA, α-naphthaleneacetic acid; TDZ, Thidiazuron; 2,4-D, 2,4-di-
chlorophenoxyacetic acid; TRIA, triacontanol; PGR(s), plant growth regulator(s); MS, Murashige and Skoog
⁎
Corresponding author.
E-mail address: slkothari28@gmail.com (S.L. Kothari).

http://dx.doi.org/10.1016/j.hermed.2017.07.003
Received 1 November 2016; Received in revised form 17 March 2017; Accepted 31 July 2017
Available online 02 August 2017
2210-8033/ © 2017 Elsevier GmbH. All rights reserved.
S. Gupta et al.
Fig 1. Moringa oleifera Lam. a. Tree growing in natural habitat. b. Tree in flowering stage. c. Tree in fruiting stage. d. Gum secretion from the cut marks on the stem. (For interpretation of
the references to color in the text, the reader is referred to the web version of this article.)
et al., 1980). These gum exudates are water insoluble due to their
mucilaginous texture and belong to the tragacanth or hog gum series
(Ramachandran et al., 1980).
It is a fast growing, medium sized deciduous tree, propagated as a
perennial plant from cuttings and seeds (Ramachandran et al., 1980;
Seshadri and Nambiar, 2003). It mainly grows in semi-arid, tropical and
subtropical regions, although the drought resistance properties of the
plant makes it more suitable for drier regions (Farooq et al., 2012;
Leone et al., 2015; Seshadri and Nambiar, 2003; Thurber and Fahey,
2009). The roots are susceptible to water-logging and tend to rot in such
conditions. It is also able to tolerate a wide range of soil types, the
optimum being a well-drained sandy or loamy soil (pH 5–9) (Patel
et al., 2010a,b; Ramachandran et al., 1980).
M. oleifera is valued for its multiple economic, medicinal and neu-
traceutical properties worldwide. This plant has been honored as
“Botanical of the Year – 2007” by the National Institute of Health (NIH).
The Africans used to call it “Never Die” or “Miracle Tree” for its ability
to treat more than 300 diseases.
All the plant parts including immature pods, leaves, mature seeds,
flowers and roots have long been consumed by humans for a variety of
applications. The immature pods, leaves, flowers and mature seeds are
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Journal of Herbal Medicine 11 (2018) 1–11
used as vegetables (Farooq et al., 2012; Ramachandran et al., 1980;
Stevens et al., 2013). Culinary practices are affected by varying tradi-
tions and taste preferences (Seshadri and Nambiar, 2003). Traditional
dishes across the world include the use of tender young plants, young
and mature leaflets and flowers in soup and sauce preparations
(Morton, 1991; Stevens et al., 2013). Leaves are also used in salad
preparation with groundnut (Fahey, 2005; Morton, 1991; Stevens et al.,
2013), herbal tea preparations, porridge, complementary baby foods,
spice and as garnish. The fresh leaves serve as a good snack when
chewed raw (Ramachandran et al., 1980; Stevens et al., 2013).
In addition, seeds obtained from tender and mature pods are po-
pularly used in pickling (Ramachandran et al., 1980). Seeds obtained
from young pods are eaten green, and mature seeds are either roasted
or fried before consumption and taste like peanuts (Fahey, 2005;
Morton, 1991; Ramachandran et al., 1980). The extremely pungent
roots are mixed with vinegar and salt and serve as a substitute to the
popular condiment “horseradish” (Morton, 1991).
This review focuses the important medicinal and neutraceutical
properties and also the biotechnology of this high value plant M. olei-
fera.
S. Gupta et al.
2. Methodology
A thorough and critical survey of the literature related to Moringa
research was conducted up to March 2017. Various online and offline
resources were taken into consideration. The primary source of data
collection for the review included research papers and review articles
published by reputed publishers such as Springer, Elsevier, Routledge
and Taylor & Francis imprints, BMC and PLOS One. Online databases
including NCBI, Scopus and Science Direct were also referred to collect
the data on M. oleifera. The paid articles were accessed through the
Bioinformatics Infrastructure Facility Center sponsored by DBT, New
Delhi located at the University of Rajasthan campus. Some other ‘grey
literature’ sources such as webpages, conference proceedings, book
chapters and theses were also reviewed to access the maximum possible
information about the potential utilities, extent of research carried out
and current bottlenecks in the M. oleifera research. The Flora of the
Indian Desert (Bhandari, 1995), a comprehensive book based on prac-
tical survey of the Desert zone of India, was referred to for collection of
information about the geographical distribution, taxonomy and mor-
phology of the tree. Herbarium “RUBL”, located at University of Ra-
jasthan Campus was also visited to collect data regarding the mor-
phological and identification characteristics of the tree. The importance
of this plant in Ayurvedic, Unani, Siddha and folk medicinal formula-
tions was further validated through the Indian Medicinal Plant Data-
base, NMPB (http://www.medicinalplants.in).
Literature search was conducted using Google Scholar and PubMed
using the following keywords: Moringa oleifera; drumstick; sajina;
Moringa pterygosperma; metabolite profiling of Moringa; metabolite
profiling of drumstick; phytochemical analysis of Moringa; in vitro
propagation; micropropagation; uses of drumstick tree; and importance
of Moringa oleifera. The commercial importance of the tree was re-
viewed by surveying different market places and online stores for the
availability of Moringa oleifera products.
In this review the challenges associated with commercialization of
the products of drumstick tree, along with the scope of biotechnological
interventions as a remedy, are discussed. The review thus highlights the
need for future research pertaining to the application of biotechnology
for enhancement of commercial value of the tree.
3. Advances in M. oleifera research/multidimensional potential
and utilities of M. oleifera
3.1. Bioremediation potential
Use of M. oleifera as a water purifier has been a common practice for
decades, and research in this direction revealed the significant bior-
emediation potential of different components of the tree.
During the early 1990s, the efficiency and properties of the drum-
stick tree as a natural coagulant were assessed and it was found that M.
oleifera seeds, while being biodegradable and non-toxic, can act as a
viable substitute to industrial coagulants such as alum (Sutherland
et al., 1994). This coagulating potential is attributed to dimeric cationic
proteins (13 kDa) and flocculating proteins (about 6.5 kDa) as isolated
and purified from aqueous extracts of M. oleifera plant parts
(Gassenschmidt et al., 1995; Ndabigengesere et al., 1995). In fact, the
purified proteins were reported to be more effective as the coagulation
process is mediated through adsorption and neutralization of colloidal
charges (Ndabigengesere et al., 1995). These promising results led to
extensive research on the isolation and purification of efficient coagu-
lants from M. oleifera extracts, and as a result a variety of coagulating
and flocculating components were isolated from different plant parts.
Okuda et al. (2001) successfully purified an active coagulant from the
salt solution of M. oleifera seeds crude extract, while hemaggulatinating
proteins and a lectin coagulant were isolated and purified from M.
oleifera tissue extracts and seeds, respectively by Santos et al. (2009).
The coagulant activity of different extracts and purified proteins was
3
Journal of Herbal Medicine 11 (2018) 1–11
reported to be comparable to that of the most commonly used coagu-
lant, aluminium sulphate and as a consequence use of M. oleifera as a
substitute to these chemical coagulants is encouraged for turbid water
treatments (Pritchard et al., 2010), along with the use of its flower
extracts for disinfection of the waste water (Moura et al., 2011). Fur-
ther, Broin et al. (2002) cloned cDNA encoding one of the flocculating
proteins present in M. oleifera seeds and the resulting recombinant
proteins were used against clay particles, as well as Gram positive and
Gram negative bacteria. Besides being used as a natural coagulant,
flocculent and microbicidal, the tree has also been explored for its
potential to remediate toxic compounds usually found in industrial soil
and waste water.
The presence of toxins in effluents, due to improper disposal prac-
tices, lead to soil and water pollution and disturbance in the food chain
associated with a particular ecosystem. Hence, strong emphasis has
been made over the past few years for the development of efficient
methods for removal of toxicants. Use of M. oleifera as a natural bio-
sorbent for remediation of toxicants is one novel approach, in which
researchers have explored different parts of the tree for their remedial
efficiency using different parameters. Biosorbents developed by che-
mical modification of leaves serve as an excellent substitute to the
conventionally used adsorbents for removal of heavy metals, particu-
larly Cd(II), Cu(III), Ni(II) and Pb(II) from the aqueous solution (Reddy
et al., 2010a, 2012). In a similar study, Reddy et al. (2010b) demon-
strated that M. oleifera bark can serve as an efficient biosorbent for
biosorption of Pb2+, and concluded that such biosorbents can serve as
efficient substitutes to the existing synthetic adsorbents such as acti-
vated carbon and ion exchange resins. Efficient removal of azo and
anthroquinonic dyes and organic compounds such as benzene, toluene,
ethylbenzene and cumene using seed extracts and pods, respectively
has also been illustrated (Akhtar et al., 2007; Beltrán-Heredia and
Sánchez Martín, 2008).
These studies not only highlight the potential of M. oleifera as an
efficient and environmentally friendly coagulant and biosorbent for
bioremediation of polluted water and land, but open new gateways for
developing commercially viable coagulants, flocculants and biosorbents
for remediation of xenobiotics.
3.2. Nutritional value
M. oleifera has been used as a dietary supplement due to its rich
nutritional content. It is an outstanding indigenous source of vitamins,
proteins and minerals (Fahey, 2005). The tree contains digestible pro-
teins, iron, magnesium, calcium, vitamins (B6, B2 and C), and car-
otenoids. M. oleifera is one of the richest natural sources of provitamin
A (Maheshwari et al., 2014; Mehta et al., 2011; Seshadri and Nambiar,
2003).
Every part of this tree has been found to possess many nutrients.
Flowers, leaves, young shoots and immature pods are good sources of
methionine and are also rich in phosphorus, calcium and iron. The
refined seed oil is an acceptable substitute to olive oil, due to the pre-
sence of all the essential fatty acids as in olive oil (Morton, 1991). The
lipid composition of seeds is even greater than that of soyabean, making
it nutritionally important. The high content of essential amino acids in
the seeds makes it an excellent substitute to legumes, which are poor in
sulphur containing amino acids (Ferreira et al., 2008).
The leaves of the plant are also significant in terms of their use as a
nutritional supplement to combat malnutrition (Fahey, 2005; Seshadri
and Nambiar, 2003; Thurber and Fahey, 2009).
According to Botany of the Plant Industry as well as Trees of Life (an
NGO), “leaves of M. oleifera possess: 4 times more calcium and two
times more protein than milk, 7 times more Vit. C than oranges, 3 times
more potassium and iron than banana and spinach respectively and 4
times more Vit. A than carrots” (Thurber and Fahey, 2009) – and
therefore, this plant is unique as it is very rare to find such a diversified
nutrient profile in a single plant species (Razis et al., 2014).
S. Gupta et al.
This tree serves as a sustainable and economically sound nutrient
rich food supplement, for those suffering from malnutrition, especially
in developing countries. In addition, it is also being used to boost food
security, foster rural development and support sustainable land.
3.3. Pharmaceutical value
The medicinal virtues of M. oleifera are well known through its use
as a traditional medicine for a number of diseases and disorders since
ancient times (Ramachandran et al., 1980). This tree has been popu-
larly used as a folk medicine for anaemia, arthritis and rheumatism,
asthma, constipation, diarrhea, stomach pain, ulcers, intestinal spasms,
headache and sore gums (Mehta et al., 2011; Pandey et al., 2012; Razis
et al., 2014). The tree possesses abortifacient, anti-rheumatic, anti-in-
flammatory, bactericidal and diuretic activities and also serves as an-
tidote, emetic, purgative, stimulant, tonic, vermifuge (anthelmintic
medicine) and vesicant (Fatima et al., 2014; Mehta et al., 2011; Patel
et al., 2010a,b). The gum exudates are not only reported to be used in
treating various chronic disorders, but are also being used as a potential
substitute to synthetic binders and suspending agents in drug delivery
systems (Jarald et al., 2012; Panda and Ansari, 2013; Panda et al.,
2008; Varma et al., 2014). Various performulation studies ascertained
the suitability of M. oleifera gum as a mucoadhesive polymer, disin-
tegrant, binder (Patel et al., 2012) and as a sustained release polymer
(Varma et al., 2014), while the drug dissolution and disintegration
studies showed that the rate of drug release can adequately be mon-
itored by varying the gum concentration in the tablet formulation
(Basawaraj et al., 2010; Panda et al., 2008). Later, in vitro toxicity as-
says of Moringa gum confirmed its non-toxic nature, thus rendering it
safe for both topical and non-topical formulations (Jarald et al., 2012;
Panda et al., 2006). Shah et al. (2011), evaluated the efficiency of
Moringa gum as a pharmaceutical excipient based on various physico-
chemical parameters and concluded that the gum obtained from M.
oleifera serves as an excellent natural source of pharmaceutical ex-
cipient. Therefore, Moringa gum exudates can efficiently serve as a
natural substitute to the toxic and synthetic excipients, leading to en-
hanced therapeutic efficiency of the drug by improvising its stability,
rate of dissolution and disintegration. Similarly, Moringa coagulant can
be used as an efficient amorphous state stabilizer in many drug for-
mulations such as ibuprofen, meloxicam and felodipine (Bhende and
Jadhav, 2012). The salt extract of Moringa seeds has also been parti-
cularly used with Povidone (polyvinylpyrrolidone, PVP) to improve
solubility of the partially soluble NSAIDS (Noolkar et al., 2013).
Recent studies indicate that the tree gains its exceptional ability to
treat such a wide range of medical conditions from its rich phyto-
chemical composition. An overview of some of the important medicinal
properties and associated phytochemicals is as follows:
3.3.1. Antioxidant activity
The antioxidant activity of different parts of M. oleifera is mainly
attributed to the presence of ascorbic acid (Vit. C), β-carotene (Kumar
et al., 2012; Mahajan and Mehta, 2007), quercetin, kaempferol (Gupta
et al., 2012), and phenolic acids (Anwar et al., 2007; Mahajan and
Mehta, 2007; Sreelatha et al., 2011). Also, isothiocyanates, polyphenols
and rutin from leaves (Bajpai et al., 2005; Tumer et al., 2015); toco-
pherols, myricetin and lectins from seeds (Mahajan and Mehta, 2007;
Santos et al., 2005; Singh et al., 2013); procyanidins from bark
(Atawodi et al., 2010) and palmitic acid, phytosterols and 9-octadece-
namide from the flowers (Inbathamizh and Padmini, 2012) have been
regarded as potent antioxidants. Monopalmitic acid, oleic acid, tri-oleic
triglycerides present in seed oil are also reported to exhibit antioxidant
activity (Lalas and Tsaknis, 2002; Mahajan and Mehta, 2007).
Antioxidant activities of different extracts obtained from different
plant parts have been reported in several studies. Extracts of leaves in
organic solvents such as methanol and acetone have been found to
exhibit antioxidant activities (Atawodi et al., 2010; Charoensin, 2014;
4
Journal of Herbal Medicine 11 (2018) 1–11
Moyo et al., 2012b; Ogbunugafor et al., 2012; Siddhuraju and Becker,
2003; Vongsak et al., 2013). Similarly, the antioxidant activities of
aqueous extracts of roots (Satish et al., 2013) and methanolic extracts of
stems and pods (Gupta et al., 2012; Kumbhare et al., 2012) have also
been ascertained using both in vitro as well as in vivo assays.
Sreelatha and Padma (2010), reported that the diverse antioxidant
profile of leaves also corresponds to the cryoprotective nature of the
plant. The plant parts can also be used as natural preservative for fat
(Patel et al., 2010a,b).
3.3.2. Antitumor activity
Extracts of the stem and seeds are reported to exhibit cytotoxic,
anticancer and antitumor activities (Araújo et al., 2013; Onsare and
Arora, 2015; Shaban et al., 2012). Similarly, leaf extracts prepared in
different solvents have also been found to exhibit anticancer, cytotoxic
(Berkovich et al., 2013; Nair and Varalakshmi, 2011), antiproliferative
(Pamok et al., 2012), antimyelomic (Parvathy and Umamaheshwari,
2007), antileukemia, antihepatocarcinoma (Khalafalla et al., 2010) and
chemoprotective (Anwar et al., 2007; Murakami et al., 1998) activities.
Phytochemical analysis revealed that these activities are credited to
3-O-6x-oleoyl-β-D-glucopyranosyl-β-sitosterol, β-sitosterol-3-O-β-D-glu-
copyranoside, 4-(α-L-rhamnosyloxy) phenylacetonitrile, 4-hydro-
xyphenylacetonitrile and 4-hydroxyphenyl-acetamide in seeds
(Guevara et al., 1999; Villasenor et al., 1989); cis-9-hexadecenal, quinic
acid, 3,5-dihydroxy-6-methyl-2,3-dihydro-4H-pyran-4-1, 9-octadece-
namide, methyl octadecenoate in flowers (Inbathamizh and Padmini,
2012) and quercetin, kaempferol (Krishnamurthy et al., 2015; Sreelatha
et al., 2011), 4-(4′-O-acetyl-α-L-rhamnosyloxy) benzyl isothiocyanate,
O-ethyl-4-(α-L-rhamnosyloxy) benzyl carbamate, 4-(L-rhamnosyloxy)
benzyl isothiocyanate, niaziminin and niazimicin (Anwar et al., 2007;
Murakami et al., 1998) in leaves.
Recently, it was found that the antitumor activities of leaf extracts is
mediated through the antioxidant and apoptosis inducing activities of
the plant (Jung, 2014; Tiloke et al., 2013).
3.3.3. Antimicrobial activity
Antibacterial activities of different components of the plant in-
cluding leaves (Abalaka et al., 2012; Doughari et al., 2007; Kekuda
et al., 2010; Mandal et al., 2014; Moyo et al., 2012a; Oluduro, 2012;
Rahman et al., 2009; Thilza et al., 2010; Valarmathy et al., 2010), seeds
(Auwal et al., 2013; Jabeen et al., 2008; Oluduro et al., 2010; Viera
et al., 2010) and pods (Arora and Onsare, 2014) have been reported
against E. coli, S. typhi, P. aeruginosa, E. cloace, P. vulgaris, S. aureus, M.
kristinae, E. aerogenes, Shigella, B. cereus, Streptococcus-B-haemolytica, B.
subtilis, K. pneumonia, B. megaterium, S. lutea, B. sterothermophilus, S.
pyogenes, V. cholera, S. entridis, enteropathogens and wound bacteria.
Extracts of seeds (Chuang et al., 2007; Jabeen et al., 2008; Oluduro
et al., 2010) and leaves (Chuang et al., 2007; Kekuda et al., 2010;
Oluduro, 2012) are also reported to exhibit significant anti-fungal ac-
tivity against T. mentagrophyte, Pullarium sp., A. flavus, Penicillium sp., A.
niger, A. oryzae, A. terreus, A. nidulans, F. solani, R. solani, C. clados-
porioides, P. sclerotigenum and Dermatophytes (T. rubrum, E. xoccosum,
M. canis).
Significant antimicrobial activity of root extracts against E. coli, S.
aureus, P. aeruginosa, P. mirabilis, Penicillium sp., Mucor sp., A. niger and
C. albicans have also been reported.
In addition, the antiviral activities of seeds against HSV-1 (Ali et al.,
2004) and that of leaves against Epstin Barr Virus (EBV), HIV and HSV-
1 (Lipipun et al., 2003; Nworu et al., 2013; Sudha et al., 2010) have also
been tested successfully.
Further investigations showed that these antimicrobial activities are
possibly attributed to the presence of 4-(α-L-rhamnopyranosyloxy)
benzyl isothiocyanate, methyl N-4-(α-L-rhamnopyranosyloxy)benzyl
carbamate, 4-(α-D-glucopyranosyl-1→4-α-L-rhamnopyranosyloxy)-
benzyl thiocarboxamide (Oluduro et al., 2010; Pandey et al., 2012) and
4-(α-L-rhamnopyranosyloxy)benzyl glucosinolate (Goyal et al., 2007),
S. Gupta et al.
together with proanthocyanidins and glucomoringine (Maldini et al.,
2014; Singh et al., 2013). The presence of cardiac glycosides in pods
(Arora and Onsare, 2014), kaempferol, rhamnetin, kaempferitin, iso-
quercitrin and pterygospermin in flowers, spirochin and anthonine in
roots (Farooq et al., 2012; Mehta et al., 2011; Raj et al., 2011) and
aglycon of deoxy-niazimicine (N-benzyl,S-ethyl thioformate) in bark
(Nikkon et al., 2003) also contribute to their antimicrobial nature.
3.3.4. Anti-inflammatory activity
The presence of potent anti-inflammatory agents including 4-[(α-L-
rhamnosyloxy)benzyl] isothiocyanate, 4-[(4′-O-acetyl-α-L-rhamnosy-
loxy)benzyl]isothiocyanate (Morton, 1991; Stohs and Hartman, 2015;
Waterman et al., 2014), quercetin, kaempferol glucosides, (Coppin
et al., 2013), 4-(2-O-acetyl- α-L-rhamnosyloxy)benzyl isothiocyanate, 4-
(3-O-acetyl-α-rhamnosyloxy)benzyl isothiocyanate (Cheenpracha et al.,
2010), 3,5-dihydroxy-6-methyl-2,3-dihydro-4H-pyran-4-1, 9-octadece-
namide (Inbathamizh and Padmini, 2012), aurantiamide acetate and
1,3-dibenzyl urea (Maheshwari et al., 2014; Pandey et al., 2012;
Sashidhara et al., 2009) in different parts of M. oleifera have been re-
ported. The reported anti-inflammatory and anti-arthritic activities of
seed extracts are ascribed to the presence of glycosides (Gupta et al.,
2005; Hamza, 2010; Mahajan et al., 2007).
Further, Sulaiman et al. (2008), demonstrated that anti-in-
flammatory activity of leaves is mediated through inhibition of in-
flammation signaling pathways. The role of leaf and root extracts in
treatment of inflammation (Ezeamuzie et al., 1996; Pal et al., 1995) and
that of pod extracts in amelioration of inflammation associated dis-
orders such as cancer, asthma, allergic rhinitis, atopic dermatitis and
rheumatoid arthritis (Lee et al., 2013; Muangnoi et al., 2012) has also
been studied.
3.3.5. Cardio-protective activity
The presence of O-[2′-hydroxy-3′-(2″-heptenyloxy)]propyl un-
decanoate, O-ethyl-4-(α-L-(rhamnosyloxy)-benzyl carbamate, methyl p-
hydroxybenzoate, (Faizi et al., 1998; Pandey et al., 2012; Stohs and
Hartman, 2015) and 4-(α -L-rhamnosyloxy benzyl)-O-methyl thio-
carbamate (Farooq et al., 2012; Pandey et al., 2012) correspond to the
hypotensive nature of the plant as a whole, while N- α-L-rhamnopyr-
anosyl vincosamide from the bark is reported to exhibit cardioprotec-
tive effects (Morton, 1991).
The hypotensive and bradycardiac activities of leaves are attributed
to niazinin, niazimicin, niaziminin, niazimin, niazirin, niazicin, niazir-
inin, 4-(4′-O-acetyl-α-rhamnosyloxy) benzyl isothiocyanate (Bose,
2007; Faizi et al., 1994a,b), and glucomoringine is reported to have a
hypertensive effect (Dangi et al., 2002; Maheshwari et al., 2014).
Leaves are also reported to exert a positive effect on the circulatory
system and reduce mortality and morbidity as a consequence of cor-
onary heart diseases, due to the presence of gossypetin, quercetagenin
and proanthocyanidins (Kumar et al., 2012; Seshadri and Nambiar,
2003). Nandave et al. (2009), demonstrated that the cardioprotective
activity of leaves is mediated through its antioxidant, antiperoxidative
and myocardial preservative effects. Later, Abdulazeez et al. (2016)
reported that the blood pressure effect of alkaloids and flavonoids
present in seeds and leaves is due to their inhibitory effect on angio-
tensin converting enzymes.
3.3.6. Immunomodulatory activity
Leaf extracts have been found to exhibit both immunosuppressive
and immunostimulatory activities (Rachmawati and Rifa’i, 2014). The
immunomodulatory effect of leaves is mediated through reduction in
cyclophosphamide induced immunosuppression by stimulating both
cellular and humoral immunity (Gupta et al., 2010), which is attributed
to the presence of compounds like isothiocyanates and glycoside cya-
nides (Sudha et al., 2010).
5
Journal of Herbal Medicine 11 (2018) 1–11
3.3.7. Neuroprotective effect
Hannan et al. (2014), revealed that leaves exert a neuroprotective
effect by promoting neuronal survival and neurite outgrowth. Leaf ex-
tracts are also reported to exhibit a protective effect against Alzheimer’s
disease by altering brain monoamine levels and electrical activity
(Ganguly and Guha, 2008). Further, it was found that the CNS de-
pressant action of leaves is mediated through its analgesic and antic-
onvulsive activities (Bakre et al., 2013; Gupta et al., 1999; Ray et al.,
2003).
3.3.8. Treatment of metabolic disorders
Leaf extracts have been reported to possess hypocholesterolemic
(Ghasi et al., 2000), antihyperglycemic, antihyperlipidemic and hepa-
toprotective activities (Divi et al., 2012; Jain et al., 2010; Jaiswal et al.,
2009; Kumar and Pari, 2003; Ndong et al., 2007; Pari and Kumar,
2002).
Later, Tahiliani and Kar (2000) and Chumark et al. (2008) affirmed
the presence of anti-hyperthyroidism and anti-atherosclerotic activities
in leaves. Also, extracts of roots have been found to possess estrogenic,
anti-estrogenic, progestational and anti-progestational activities (Bose,
2007; Shukla et al., 1988) along with liver and kidney protective effects
(Mazumder et al., 1999).
Phytochemical analysis showed that β-sitosterol and 4-[α-(L-rham-
nosyloxy) benzyl]-o-methyl thiocarbamate (trans) are potent choles-
terol lowering, anti-diarrheal, antispasmodic, hypolipidemic agents
(Anwar et al., 2007; Jain et al., 2010; Maheshwari et al., 2014), while
quercetin-3-glycoside is responsible for the antidiabetic (Farooq et al.,
2012), antidyslipidemic (Mbikay, 2012) and hypoglycemic
(Maheshwari et al., 2014) effects. Also, moringine and moringinine are
reported to have a role in antihypoglycemic properties of the plant (Kar
et al., 1999; Maheshwari et al., 2014; Mehta et al., 2011). The presence
of insulin like plant proteins in leaf extracts has been recently reported
for treatment of diabetes (Paula et al., 2016).
Stohs and Hartman (2015), reported the presence of a novel com-
pound, niaziridin, which enhances the gastrointestinal absorption of
vitamins and nutrients. The presence of quercetin and kaempferol in
flowers and pods has been reported to be associated with their anti-
diabetic and hepatoprotective activities (Farooq et al., 2012; Gupta
et al., 2012).
3.3.9. Other activities
M. oleifera has also been reported to have wound healing activities.
Potent wound healing agents such as vicenin-2 (Muhammad et al.,
2013), quercetin, kaempferol and phytosterols (Hukkeri et al., 2006)
have been reported in ethyl acetate extract of the leaves. Recently,
Lambole and Kumar (2012) reported dexamethasone present in bark as
an efficient wound healer. It mediates the activity by suppressing anti-
healing agents. The wound healing activity has also been reported in
the seeds of M. oleifera (Bhatnagar et al., 2013; Parwani et al., 2016).
Leaves, seeds and roots of the plant also possess various other activities,
such as schizonticidal, antiulcerogenic (Dahiru et al., 2006; Patel et al.,
2010a,b), larvicidal (De Oliveira et al., 2011; Prabhu et al., 2011),
antiurolithiatic and anti-ulcerative colitis (Gholap et al., 2012; Karadi
et al., 2006), whereas the gum exudates of plant are reported to exhibit
macrofilaricidal activity (Kushwaha et al., 2011).
Extracts of the leaves contribute significant radioprotective effects
as estimated by Rao et al. (2001) using comparative scoring of aber-
rations in bone marrow metaphase chromosomes between mice pre-
treated with leaf extracts and those without any pre-treatment prior to
irradiation. Seeds are reported to exhibit antipyretic (Hukkeri et al.,
2006), aphrodisiac (Zade et al., 2013), antihypersentitivity and anti-
anaphylactic (Mahajan and Mehta, 2007) activities along with being a
potent antitoxidant to fluoride toxicity (Ranjan et al., 2009).
The anti-asthamatic and anti-rhinitis activities of seed extracts were
attributed to the presence of moringine and moringinine, which play a
significant role in bronchiole relaxation and treatment of respiratory
S. Gupta et al. Journal of Herbal Medicine 11 (2018) 1–11

tract infections (Agrawal and Mehta, 2008; Mahajan and Mehta, 2008). 5. Need for M. oleifera biotechnology
Mehta et al. (2003), reported the presence of anti-atherosclerotic ac-
tivity in pods, while the protective effect of the phenolic compounds of In the past few years, Moringa has gained the attention of both
the tree against oxidative stress was recently reported by Ramabulana scientific and business communities due to its economic importance in
et al. (2016). terms of food, medicines, waste management and sustainable agri-
Evidently, drumstick tree serves as a good source of neutraceuticals culture practices. The extensive research and attempt for commercia-
which are very rare to find altogether in a single plant in sufficient lization of drumstick tree and its products has led to an urgent need for
quantities. Thus, the tree can be regarded as medicinally important for its conservation – from dietary, pharmacological, ethnobotanical and
prevention as well as cure. biotechnological perspectives.
One solution to the increasing demand for M. oleifera based products
is biotechnological approaches, which include in vitro propagation for
3.4. Other economic utilities
mass multiplication, enhanced biosynthesis of secondary metabolites
through metabolite profiling, pathway engineering and recombinant
Apart from the above discussed properties, the tree has also been
technology.
studied for some other economic utilities such as feed value and a
source of natural gas. M. oleifera has emerged as an important cattle
5.1. Problems associated with M. oleifera propagation
feed that can be used as a substitute to conventional animal feeds such
as sunflower seed cake and alfalfa (Babiker et al., 2016; Sarwatt et al.,
The Drumstick tree is easily propagated by stem cuttings as well as
2002) due to its high nutritional value. Use of Moringa silage as a feed
seeds, but propagation by cutting reduces the growth of the mother
has been reported to exhibit a significantly positive effect on milk
plant and also affects the yield and sometimes can cause death of the
quality and quantity in dairy cows (Mendieta-Araica et al., 2011), and
mother plant (Islam et al., 2005). Plants obtained by seed propagation
milk yield and growth performance in goats and lambs (Babiker et al.,
vary in genotypes leading to variations in phenotypes and nutritional
2016). Further, Aregheore (2002) suggested that use of the tree foliage
values (Riyathong et al., 2010). The tree also has low seed viability and
as a part of dairy forage for goats serves as an economical source of
low seed germination rates, which in turn results in slow population
protein supplementation. The cost of beef production and nutritional
growth rates (Stephenson and Fahey, 2004).
quality of roughages was estimated to be optimum when M. oleifera was
The need for large areas of agricultrural land for mass cultivation of
used as a feed, thereby serving as an economical forage for cattle (Roy
M. oleifera, to meet the increasing demand of industries for production
et al., 2016).
of finished goods is somewhat impractical in a society where scarcity of
The tree has also been reported to be an efficient feedstock for
cultivable land is a major concern due to increasing population and
biofuels such as biogas and biodiesel (Kafuku and Mbarawa, 2010;
industrialization.
Rashid et al., 2008). The vegetable oil obtained from the seeds of tree
M. oleifera is highly susceptible to infestations with flies, aphids,
has been reported to be of superior quality than that of Jatropha curcas
thrips and other insects, leading to foliar damage which in turn results
(Kibazohi and Sangwan, 2011). These findings provide new horizons
in decline in total biomass and thereby making it unfit for consumption
that could be investigated through sustainable development and green
as food and medicine. Additionally, the frost sensitive nature of the tree
technology.
makes it almost impossible to grow it in open lands of temperate zones
In addition to culinary uses, the oil obtained from the seeds has also
(Förster et al., 2013).
been used for a variety of purposes such as lighting, hair dressing, lu-
Being the sole cultivable species of family Moringaceae, the tree has
brication in watches, soap making and perfume production (Morton,
to meet the growing demands of local population for its use as food,
1991). The seed cake remaining after oil extraction is commonly used
medicine and water clarifiers which may lead to gradual decline in its
as a biofertilizer due to its rich nutrient content.
biodiversity due to reducing natural tree resources (Steinitz et al., 2009;
Pontual et al. (2012) reported the caseinolytic and milk coagulating
Stephenson and Fahey, 2004).
activities of proteases present in flowers, while in a recent similar study,
Apart from these issues, one of the major drawbacks that usually
milk clotting enzymes were purified and characterized from seeds and
holds back the commercialization of such medicinal plant species is
were successfully employed in cheese making, thus suggesting an al-
fluctuations in metabolic constituents due to inconsistent environ-
ternative to conventional animal rennets (Ahmed, 2016).
mental conditions, harvesting conditions and other variations which in
Gum exudates obtained from excisions on the bark have been used
turn causes difficulties in standardization of commercial formulations.
in leather tanning and calico printing (Morton, 1991).
Moreover, the geographical distribution of the tree also results in var-
Lee et al. (2016), recently developed an antimicrobial, antioxidative
iations in morphological, metabolic and genetic characteristics as a
and biodegradable packaging material from puffer fish skin gelatin
consequence of different climatic conditions prevailing in different
containing M. oleifera leaf extract.
zones (Bhatia et al., 2013; Chatterjee et al., 2010).
Therefore, the tree is not only important from medicinal and nu-
tritional perspectives, but also has a vast range of other potential uti-
5.2. Biotechnological strategies: potential solution
lities.
Biotechnology is the only field of science that not only aids in
4. Commercial value of the plant gaining insights into the actual mechanisms of living bodies but also
provides numerous techniques which can be employed for their further
The considerable medicinal potential has led to the development of improvisation. Plant biotechnology is one such specialized field of
various commercial formulations, such as Shrigu, Rumalaya, Septilin biotechnology in which biotechnological tools and techniques enable
(The Himalaya Drug Company, Bangaluru, India), Livospin (Herbals one to explore plant systems and to understand mechanisms associated
APS Pvt. Ltd., Patna, India) and Orthoherb (Walter Bushnell Ltd. with different activities followed by their improvisation such as en-
Mumbai, India), while Ratnagiri Rasa, Sarasvata Ghrta, and Sudarsana hanced growth and yield, rapid mass multiplication, increased meta-
churna are some of its ayurvedic formulations. bolite production and secretion. Plant tissue culture, metabolite pro-
The gum exudates of the tree are valued in the Indian as well as filing, genetic and pathway engineering are some of the common tools
global markets for its dark colored texture and rich medicinal value. of plant biotechnology which are being used extensively to achieve
The increasing global demand and limitations in the availability of the different objectives of crop improvement in terms of yield, disease re-
gum has also resulted in its increasing costs. sistance and enhanced nutritional value.

6
S. Gupta et al.
Plant tissue culture via in vitro propagation helps in the conservation
of endangered, medicinally and economically important plant species.
It also provides scope for improvisation by generation of /virus free
plants, maintaining supply of seasonal crops throughout the year and
rapid bulk propagation. Plants possess a variety of phytochemicals
which exhibit numerous medicinal and non-medicinal properties, but
lack of knowledge about the phytocomposition and pathways involved
makes it difficult to exploit their properties at commercial scale.
Metabolite profiling aids in identification of key enzymes involved in
biosynthesis of commercially important phytochemicals. Genetic en-
gineering leads to generation of new varieties possessing the desired
characteristics and are commonly referred to as genetically modified
(GM) plants.
In addition, the analysis of morphological, biochemical and genetic
variability using suitable molecular markers not only helps in the se-
lection of superior varieties of interest for future breeding programs and
mass multiplication for commercial purposes, but also provides insights
into the effects of different biotic and abiotic factors on the morpho-
logical, phytochemical and genetic characteristics (Goyal et al., 2015).
Such analysis facilitates the selection and identification of superior
varieties of plants with particular desired characteristics, which can be
further subjected to mass multiplication and breeding programs.
5.2.1. Micropropagation
In recent years, in vitro micropropagation based techniques have
proved a fruitful approach for mass multiplication and germplasm
conservation of endangered as well as economically and medicinally
important plants (Goyal et al., 2015; Jain et al., 2009; Steinitz et al.,
2009; Stephenson and Fahey, 2004).
Plant tissue culture technologies serve as an excellent alternative to
conventional propagation methods, resulting in year round availability
of the tree which involves rapid clonal mass multiplication in a stipu-
lated time and space. This approach enables meeting the huge in-
dustrial demands for drumstick tree as raw materials in variety of drug
and dietary formulations.
According to Jain et al., (2011), in vitro regeneration and multi-
plication techniques are not only being used as a one-stop solution to all
the major concerns such as overexploitation, enhancement of nutri-
tional value and disease resistance but can also be used for commercial
production of metabolites.
Therefore, current biotechnology research is focused on optimiza-
tion of in vitro propagation methods using different parts of M. oleifera
in order to meet the commercial demand and overcome the formerly
discussed issues.
In vitro propagation of M. oleifera was reported by Stephenson and
Fahey (2004), in which they used meristematic stem nodes and im-
mature seeds as explants and reported that the explants showed best
response in full and half strength Murashige and Skoog (MS) medium.
Multiplication of shoots was dramatically increased when the nodal
segments were cultured on membrane raft. The use of isothiazolones for
improved survival rate of the rooted plantlets was also reported in the
study.
In a similar report by Förster et al. (2013), nodes taken from the
sterile seedlings of two different ecotypes were cultured on MS medium
supplemented with different concentrations of 6-benzylaminopurine
(BAP). Results revealed that MS medium without BAP and with 0.5 mg/
L BAP showed best response in terms of shoot length (3.81–4.10 cm)
and number of offshoots (4.34–4.40), respectively. In experiments
conducted using nodal segments as an explant, Islam et al. (2005) re-
ported that at BAP (1 mg/L and 1.5 mg/L) all the explants produced
shoots and maximum number of shoots (4 ± 0.29) were obtained with
1 mg/L BAP. On the other hand, Mathur and Kamal (2012) reported the
best response on MS medium supplemented with BAP:1-Naphthalene
acetic acid (NAA) at 3:3 mg/L. Further, Islam et al. (2005) found that
repeated subcultures on same BAP concentrations resulted in increased
number of shoots (25) and concluded that BAP is the most effective
7
Journal of Herbal Medicine 11 (2018) 1–11
plant growth regulator (PGR) for shoot induction and multiplication.
They also recorded 100% rooting in MS medium without any PGRs, but
maximum number of roots (2.2–3.7 and 2.8) was reported on MS
medium containing Indole-3-acetic acid (IAA) at 0.05, 1 & 2 mg/L and
Indole-3-butyric acid (IBA) at 2 mg/L, respectively (Islam et al., 2005).
Studies based on establishment of an in vitro propagation method
using seeds are also reported. In a study carried out by Marfori (2010),
nodes from seedlings in a sterile environment were subjected to MS
medium supplemented with varying concentration regimes of different
growth hormones and the highest number of axillary shoots per ex-
plants (4.6) was obtained in medium containing 2.5 μM BAP, 5 μM Ki-
netin (Kn) and 2.5 μM Thidiazuron (TDZ). Other cytokinins, i.e. Kn
(10 μM) and TDZ (5–10 μM) showed inhibitory effect. Saini et al. (2012)
also reported that nodal segments obtained from aseptically grown
seedlings, when inoculated on MS medium containing 4.44 μM BAP
showed best response (9 ± 1.0) and highest number of roots (8 ± 0.8)
were observed on IAA (2.85 μM) and IBA (4.29 μM).
Devendra et al. (2012) carried out tissue culture study using zygotic
embryos as explants. Zygotic embryos grown on MS medium containing
10.75 μM NAA resulted in white organogenic callus, which was further
subjected to induction of somatic embryos using MS media supple-
mented with 4.52 μM 2,4-Dichlorophenoxyacetic acid (2,4 D) and
11.09 μM BAP. They concluded that MS medium with 13.31 μM BAP
showed highest somatic embryo induction frequencies with germina-
tion efficiency of 77.2%. Further, maximum rooting efficiency (78.4%)
was reported in MS medium supplemented with 10.75 μM NAA.
Similar studies of callus cultures were also carried out using leaves
and cotyledons as explants.
Highest callus growth was obtained using leaf explants in MS
medium supplemented with 0.1 mg/L 2,4-D (Khalafalla et al., 2011).
Nieves and Aspuria (2011) established callus culture for cotyledons and
observed increased callus formation on MS medium containing 2.5 mg/
L BAP. Also, enrichment of media with TDZ results in compact, nodular
and loose calli (Nieves and Aspuria, 2011).
Riyathong et al. (2010) used stem explants obtained from in vitro
raised seedlings and 100% shoot formation was obtained on MS
medium supplemented with BAP (1–4 mg/L), and the highest number
of shoots per explant (10.2) was achieved with BAP (2 mg/L).
In a recent report by Avila-Treviño et al. (2017), successful micro-
propagation protocol was established using buds and cotyledonary
nodes of the tree without using any plant growth regulators and they
further illustrated the genetic similarity among the regenerated plant-
lets using RAMP markers. The tissue culture studies are summarized in
Table 1.
Thus, it can be concluded that the in vitro propagation studies car-
ried out so far provide a platform for further/future interventions in
order to obtain superior varieties with enhanced medicinal and nutri-
tional value. In one such attempt, Mathur and Kamal (2012) studied the
biosynthesis of “trigonelline – a metabolically active pyridine alkaloid”
in different components of M. oleifera including the callus cultures and
reported that the trigonelline production in in vitro grown cultures can
be significantly enhanced using nicotinic acid as precursor in the
medium.
Additionally, use of plant cell suspension cultures and hairy root
cultures for large scale production of commercially important sec-
ondary metabolites have been suggested by Rao and Ravishankar
(2002), whereas Förster et al. (2013) concluded that in vitro cultures are
successfully being used in medicine and food industries as a source for
important metabolites.
The importance of comprehensive metabolite profiling and en-
gineering was highlighted by Zhou and Wu (2006). They illustrated the
role of plant tissue culture techniques along with plant genomics,
transcriptomics, proteomics and metabolomics for production of im-
portant plant pharmaceuticals and neutraceuticals on a commercial
scale.
Therefore, it is inferred that comprehensive knowledge of the
S. Gupta et al.
Table 1
In vitro propagation studies in M. oleifera.
Explant Used PGRs Used
Meristematic Nodes, Immature Seeds BAP, Ancymidol, NAA, Kinetin
Nodes BAP, IBA, IAA
Seeds BAP, Kinetin, TDZ, NAA
Leaves 2,4-D
Cotyledons BAP, 2,4-D, TDZ
Leaves, Zygotic embryos 2,4-D, NAA, Kinetin, BAP
Nodes IBA, BAP
Nodes Triacontanol (TRIA), BAP, IAA, IBA
Nodes BAP, IAA
Nodes, Leaf No PGR(s)
metabolites present and their biosynthesis pathway is the key behind
converting plant cells and tissues into large bioreactors which are often
referred to as “plant cell factories” for commercial production of phy-
tomolecules/phytochemicals.
6. Conclusion and future prospects
In recent years, great emphasis has been laid on the utilization of
natural resources as a source of food, medicine and waste management
due to the harmful effects posed by the synthetic resources, in addition
to their non-renewable nature. Moringa oleifera Lam. is a tree with
multidimensional utilities which serves as an excellent reservoir for
pharmaceuticals and neutraceuticals. The rich phytochemical profile
and advances in biotechnological techniques has led to generation of
new avenues aimed towards enhancement of overall commercial value
of the tree.
In vitro propagation techniques have emerged as a life-line for ex situ
conservation of endangered and elite germplasm and also lay the
foundation for the concept of “plant cell factories” i.e. the natural
bioreactors for production of plant products. This concept provides new
insights into development of more effective, eco-friendly and biode-
gradable products using mass multiplication and production techni-
ques. Establishment of plant cell factories demands thorough knowl-
edge of the metabolomics of the plant as it facilitates in identification
and alteration/modulation of the key components for enhanced pro-
duction of targeted plant product(s).
Although efficient in vitro propagation protocols for M. oleifera have
been established, comprehensive studies involving the use of metabolite
profiling and engineering for enhanced metabolite production in in vitro
cultures are still lacking. Knowledge of biosynthetic pathways of the
desired compounds in plants and in culture systems will facilitate the
modulation and fabrication of such compounds in in vitro cultures.
Therefore, a comprehensive metabolite profiling followed by biosyn-
thetic pathway identification using high throughput techniques such as
GC–MS and NMR is essential for identification and characterization of
novel neutraceuticals and pharmaceuticals.
The use of biotechnological approaches thereby not only facilitates
selection and conservation of superior varieties but is also a stepping
stone to commercialization of important/targeted plant products.
There is no doubt that the shortcomings in M. oleifera commercia-
lization can be overcome by the use of biotechnological tools, and it is
likely that the future research in this context will make M. oleifera an
important solution for the various health and environmental issues of
the present era/generation.
Conflicts of interest
The authors have no conflict of interest.
8
Journal of Herbal Medicine 11 (2018) 1–11
Method of Propagation Reference(s)
Direct Shoot Induction Stephenson and Fahey (2004)
Direct Shoot Induction Islam et al. (2005)
Direct Shoot Induction Marfori (2010)
Callus Khalafalla et al. (2011)
Callus Nieves and Aspuria (2011)
Somatic embryogenesis Devendra et al. (2012)
Direct Shoot Induction Mathur and Kamal (2012)
Direct Shoot Induction Saini et al. (2012)
Direct Shoot Induction Förster et al. (2013)
Indirect regeneration/callus induction Avila-Treviño et al. (2017)
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