Industrial Crops and Products 97 (2017) 639–648
Contents lists available at ScienceDirect
Industrial Crops and Products
journal homepage: www.elsevier.com/locate/indcrop
Role of conventional and biotechnological approaches for genetic
improvement of cluster bean
Sushil Kumar a,∗ , Arpan R. Modi a , Mithil J. Parekh a , Hans R. Mahla b , Ramavtar Sharma b ,
Ranbir S. Fougat a , Devvart Yadav c , Neelam R. Yadav d , Ghanshyam B. Patil a
a
Department of Agricultural Biotechnology, Anand Agricultural University (AAU), Anand, 388 110, India
Division of Plant Improvement, Propagation and Pest Management, Central Arid Zone Research Institute (CAZRI), Jodhpur, 342 003, India
c
Department of Genetics and Plant Breeding, CCS Haryana Agricultural University (CCSHAU), Hisar, 125 004, India
d
Department of Biotechnology and Molecular Biology, CCS Haryana Agricultural University (CCSHAU), Hisar, 125 004, India
b
a r t i c l e
i n f o
Article history:
Received 12 May 2016
Received in revised form 3 January 2017
Accepted 7 January 2017
Keywords:
Breeding
Biotechnology
Cluster bean
Diversity
Guar
Molecular markers
a b s t r a c t
Cluster bean [Cyamopsis tetragonoloba (L.) Taub. (Syn. C. psoraliodes)], commonly known as guar and an
important crop from family Leguminaceae, is grown under resource constrained situations for use as seed,
vegetable and forage in arid and semi-arid regions. The seed of this drought-resilient legume contains
galactomannan polysaccharide, used in wide range of industries, which has made this orphan crop a
high-valued cash crop. Cluster bean shows limited variability for morphological and agronomic traits.
Narrow genetic base of cultivated cluster bean varieties and yield losses due to both biotic and abiotic
stresses has hampered the intensive breeding efforts in cluster bean. Conventional breeding methods
viz. induced mutations, wide-hybridization and induce male sterility have been employed to broaden
the limited genetic base and for genetic improvement of cluster bean. Due to its pivotal role in rainfed
agriculture, research efforts using biotechnological interventions like molecular markers, tissue culture
and transformation have been initiated to boost the varietal improvement but development of these
tools are still at early stage in cluster bean. This article attempts to summarize and discuss the recent
progress made in mutation breeding, distant hybridization, DNA marker studies, development of in-vitro
propagation system and genetic transformation protocols.
© 2017 Elsevier B.V. All rights reserved.
Contents
1.
2.
3.
4.
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 639
Genetic improvement of cluster bean through conventional approaches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 640
2.1.
Genetic diversity and germplasm resources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 640
Induced variation through mutation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 641
2.2.
2.3.
Wide crosses and distant hybridization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 642
2.4.
Male sterility and heterosis breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 642
Role of biotechnology in cluster bean improvement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 642
Molecular markers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 642
3.1.
Tissue culture, regeneration protocols and genetic transformation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 643
3.2.
Conclusion and future prospects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 646
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 646
1. Introduction
∗ Corresponding author.
E-mail addresses: sushil254386@gmail.com, sushil254386@yahoo.com
(S. Kumar).
http://dx.doi.org/10.1016/j.indcrop.2017.01.008
0926-6690/© 2017 Elsevier B.V. All rights reserved.
Globally cluster bean/guar (Cyamopsis tetragonoloba, 2n = 14), a
member of family Leguminaceae, is an important crop of arid and
semi-arid regions (Kumar et al., 2015). It is believed that geographic
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S. Kumar et al. / Industrial Crops and Products 97 (2017) 639–648
centre of origin of cluster bean is India although no wild species has
been reported in country. Gillette (1958) pointed out that tropical
Africa is its probable centre of origin as wild species were found to
occur in that region. Due to deep rooted system, this short duration,
hardy and drought resilient legume is extremely adapted to the
inhospitable environment of rainfed regions. This ancient legume is
mainly cultivated for feed and food (Singh and Dahiya, 2004) therefore providing nutrition to both mankind and animals. It is used
for human consumption and cattle feed (30–40%), industrial purpose (50–55%), medicinal (5%) as well as for soil improvement and
other miscellaneous purposes. Due to multiple usages and presence of gum in its seeds, cluster bean has been emerged as a unique
high-valued commercial crop.
Cluster bean is predominantly autogamous and bears beans or
pods in clusters, hence named as cluster bean. The light-grey, pink,
white or black coloured seed consists of 43–47% germ, 35–42%
endosperm and 14–17% husk (Goldstein and Alter, 1959). In cluster bean seed, nearly 90% (w/w) portion of the spherical-shaped
endosperm is deposited with galactomannan gum −a polysaccharide synthesized in Golgi apparatus (Dhugga et al., 2004). In
cold water, this odourless polysaccharide forms a thick colloidal
solution (Das and Arora, 1978). Galactomannan gum has a genetically controlled ratio (1:2) of galactose to mannose (Sandhu et al.,
2009). The gum consists of a mannose framework with a galactose group attached to it and act as reserve of glucose to be used
by the seed during germination. Cluster bean gum has diversified uses such as in textile, ore-/metal-refining, paper, coal-mining,
petroleum drilling, cosmetic and pharmaceuticals, explosion manufacture, potash purification, tobacco and food enterprises (Punia
et al., 2009; Kuravadi et al, 2013). It is an integral part of the natural gas exploration process known as a hydraulic fracture. Due to
hydrophilic and swelling nature, the gum and its derivatives in various forms such as coatings, matrix tablets, hydrogels, and nano- or
micro-particle have found their use in developing slow releasing
and targeted drug delivery (Prabaharan, 2011). Cluster bean is also
being utilized to cure diabetic and lipoproteins/cholesterol patients
(Kumar et al., 2013).
Cluster bean requires reasonably warm weather and moderate
rainfall for its growth and therefore, it is mainly cultivated as a
cash crop in the Indian subcontinent (India and Pakistan). Though,
to a narrow extent, the crop is also under cultivation in Australia,
Bangladesh, Myanmar, USA, South Africa, Brazil, Congo, Sri Lanka
(Boghara et al., 2015). Due to abrupt and unexpected rise in demand
of guar and its gum in recent years, its cultivation has extended to
resource rich regions and alternative seasons under proper management (Kumar et al., 2015). The chief importer of guar gum and
its derivatives is Australia, Austria, Brazil, Canada, China, Chile,
Germany, Greece, Ireland, Italy, Japan, Mexico, Portugal, Sweden,
UK and USA (NRAA, 2014). The annual world’s total cluster bean
gum and its derivatives production is around 0.75–1.0 million
tonnes. Globally, India ranks first in production as producing about
75–82% of the world’s cluster bean followed by Pakistan (10–12%).
In India, with 75% of total production, Rajasthan is the top cluster
bean producing state.
In spite of its importance, the productivity of this crop is very low
and hence there is need to enhance the yield of cluster bean. Lack
of suitable early maturing high yielding varieties, an incidence of
many fungal and bacterial diseases in rainfed cultivation, improper
time of sowing, inadequate fertilization and improper agronomic
practices are major production constraints for adequate production of this versatile crop. Therefore, there is an urgency to design
breeding approaches aiming at developing proper plant architecture coupled with tolerance to abiotic and biotic stresses in order
to stabilize the yield of guar at higher level of production. At the
morphological level, cluster bean germplasm showed variability
(Saini et al., 1981; Mishra et al., 2009; Pathak et al., 2011c) but this
variability is not sufficient for stress tolerance and gum content.
Therefore, broadening the genetic base and developing stress tolerant varieties with high galactomannan content is restricted due
to inadequate variability available for these traits. Thus, along with
various conventional breeding approaches like mutation breeding, distant hybridization, male sterility etc, there is urgent need
to explore non-conventional genetic improvement approaches like
molecular marker, tissue culture and genetic transformation technologies. This article reviews the conventional breeding work done,
highlights the current role of biotechnology and its future prospects
to enhance production and productivity of cluster bean through
genetic improvement.
2. Genetic improvement of cluster bean through
conventional approaches
2.1. Genetic diversity and germplasm resources
The availability of genetic diversity and its successful collection, maintenance, utilization and conservation is pre-requisite
for crop improvement program (Poehlman and Sleper, 1995). The
genetic variability existing in cluster bean germplasm has been
evaluated by using various morphological and biochemical traits.
Conspicuous morphological variations are displayed for branching
(branched/unbranched), pubescence (hairy/smooth), pod shape
(straight/sickle), growth habit (determinate to indeterminate), pod
bearing pattern (regular/irregular) (Saini et al., 1981). In India, the
exhaustive catalogue by Dabas et al. (1989) has provided information on fifteen morpho-physiological and yield traits along with
place of collection for all 3580 accessions. Many important traits
like plant height (46.75–239 cm), clusters per plant (1.75–64.5),
pods per plant (2.25–262.35), pod length (1.85–19.3 cm), seeds
per pod (4.15–13.0) days to maturity (128–185 d), seed yield
(0.95–59.7) and 100 seed weight (1.9–4.75 g) showed a high degree
of diversity. Though, there was paucity of early flowering and
maturing genotypes in this collection but was later on enriched
by many collections evaluated at Regional Station, National Bureau
of Plant Genetic Resources (NBPGR), Jodhpur, India where certain genotypes flowered as early as 28 days and matured in 70
days. Many other studies involving comparatively limited genotypes though reported considerable diversity towards desirable
direction but with reduced range (Mishra et al., 2009; Pathak et al.,
2011a, 2011c). Plant height and branches per plant on a higher
side are important as they bear more clusters and branches (Dabas
et al., 1989). High level of diversity for yield traits viz. number of
pods, clusters etc. have also been reported by many studies along
with other morphological traits (Mittal et al., 1977; Dabas et al.,
1982; Henry and Mathur, 2005; Mahla and Kumar, 2006; Pathak
et al., 2009; Pathak et al., 2010a; Kumar et al., 2014; Girish et al.,
2012; Sultan et al., 2012; Manivannan and Anandakumar, 2013;
Manivannan et al., 2015; Kumar and Ram, 2015). Morris (2010)
characterized 73 accessions collected from India, Pakistan and USA;
and reported enough genetic variability pod length (32–110 mm),
100 seed weight (2.3–4.8 gm) and number of days to 50% maturity
(96–185 days).
Despite of commercial importance, biochemical studies on
gum content are limited. The remaining part of seed comprising of seed coat and protein rich germ makes good animal feed.
A few studies have indicated some extent of diversity in these
biochemical and nutritional traits. Pathak et al. (2009) reported
considerable variation for endosperm proportion (30.4–46.3%),
gum content (23.9–34.2%) along with crude fibre (4.1–8.0%), oil
(1.8–5.2%), crude protein (28.3–35%), carbohydrate (38.8–59.1%)
and minerals (3.5–6.0%). However, comparatively narrow range
was obtained by Kays et al. (2006) for dietary fibre (52.4–57.7%),
S. Kumar et al. / Industrial Crops and Products 97 (2017) 639–648
protein (22.9–30.6%), fat (2.88–3.45%) and ash (3.04–3.53%). In
addition to above primary phytochemicals, Wang and Morris
(2007) observed variations for seed derived secondary metabolites viz. daidzein, genistein, quercetin and kaempferol in guar
germplasm.
A number of studies have indicated significant correlations
among economically important traits resulting in simultaneous
improvement of various traits (Henry and Mathur, 2008; Mahla
and Kumar 2006; Shekhawat and Singhania, 2005; Singh et al.,
2005a, 2005b). Many economically important traits like pods per
plant, pod length, clusters per plant, number of branches and hundred seed weight were found positively associated with seed yield
(Sidhu et al., 1982; Vijay, 1988; Gresta et al., 2016). Positive correlations among grain yield and gum content were not followed for
seed weight and gum percentage (Menon et al., 1970; Mittal et al.,
1971). However, percent endosperm was positively correlated with
gum percentage (Menon et al., 1970; Lal and Gupta, 1977).
In India, cluster bean improvement program was started in
1950s. The initial breeding emphasis was on vegetable type genotypes but later, due to realization of cluster bean as industrial crop,
research on seed type genotypes for gum content was intensified.
Therefore, in 1961, systematic development of genetic resources
was initiated by Division of Plant Introduction, Indian Agricultural
Research Institute (Now NBPGR), New Delhi by collecting diverse
guar germplasm sources from prime areas (Gujarat, Rajasthan and
Punjab) of variability (Thomas and Dabas, 1982). Real momentum
was picked up during 1965–70 by launching a scheme entitled: Collection and isolation of superior genotypes for gum purposes from,
under Public Law 480 in 1954 (Agricultural Trade Development
and Assistance Act). Consequently, two catalogues were published
in succession containing information on1150 accessions in 1981
(Dabas et al., 1981) and 3580 accessions in 1988 (Dabas et al., 1989).
In addition to this, independent collections were made and maintained at various research centres of state agriculture universities
especially in the North-western Indian states (Rajasthan, Gujarat,
Haryana and Punjab).
Presently National Gene Bank at NBPGR, India is maintaing
about 5000 accessions of cluster bean along with C. serrata
and C. senegalensis −two close wild relatives of guar. Cluster
bean improvement work started at Durgapura (Rajasthan), Hisar
(Haryana) and CAZRI (Rajasthan) with collection and assessment of
germplasm which resulted selection/identification of a number of
high yielding genotypes (Paroda and Saini, 1978; Henry et al., 1992;
Singh et al., 1995; Mahla et al., 2011). According to report of Mishra
et al. (2009), NBPGR, India is maintaining 4878 indigenously collected accessions in medium term storages while efforts have also
been made for long term ex situ conservation of 3714 accessions.
Till now, using traditional breeding techniques like mass selection
and pedigree methods nearly 40 varieties for vegetable as well as
seed purposes has been released in India (Pathak, 2015). Though
morphological variability exists in cluster bean but it is inadequate
for various economic traits (Jukanti et al., 2015); therefore cluster bean breeding is slower and there is urgency to broaden the
genetic base through germplasm introduction, mutagenesis, distant hybridization (Kumar et al., 2015). Similar to NBPGR, India
the United States Department of Agriculture, Agricultural Research
Service, Plant Genetic Resources Conservation Unit (USDA, ARS,
PGRCU) also conserves 1298 accessions originating from India,
Pakistan, and breeding lines from the USA (Morris 2010).
2.2. Induced variation through mutation
Natural genetic diversity coupled with induced variability forms
the basis for improvement of all major food crops (Parry et al.,
2009). The utilization of artificial and/or induced mutation to create new elite alleles is a powerful tool in crops like cluster bean
641
where variability at genetic level is low. However, in comparison
to induced mutation, frequency of desired natural mutation is very
low for accelerated plant breeding. Hence, artificial mutation using
physical and chemical mutagens is the best way to expand genetic
variability in short time period (Auti, 2012). Though, the success
of mutation either natural or induced depends on efficiency to
create desirable changes with least undesirable changes (Harten,
1998; Pathak, 2015). Artificial mutagenesis could be better option
for enriching variation in guar having small cleistogamous flowers
difficult for emasculation and further artificial crossing (Arora and
Pahuja, 2008).
Induced mutation by gamma rays in cluster bean was for the first
time carried out by Vig (1965) where 60 Co was a as source of radiation. He reported low fertility mutant in a gamma (20 kR/200 Gray;
1 kR = 10 Gray) irradiated population of cluster bean cv. Punjab G2.
Semi-sterile variants reported by him were outcome of reciprocal translocations and compared with fertile plants, these variants
were taller and low yielder and showed more racemes perplant
with prolonged vegetative period. Study of Vig (1969) also pointed
out that the guar chromosomes are radio-resistance as a dose as
30,000 kR as showed no detectable effect on the chromosomes. This
might be a delayed effect of radiation due to small chromosomes
size. Adverse effects of irradiation doses (10, 20, 40, 60, 80, 100,
150 and 200 kR) on morphology of guar was recorded by Lather
and Chowdhury (1972) where germination rate, seedling mortality rate and pollen fertility were inversely proportional to radiation
dosage. Similarly, chromosomal abnormalities such as translocation, anaphase bridges and laggards were found in the progenies
obtained from treated seeds. Irradiation at higher dose (100–200
kR/1000-2000 Gray) completely repressed the germination hence
was demonstrated as lethal. Though, mutants generated at low
doses (2, 5, 10, 15 and 20 kR) of gamma rays showed increments
for number of seed per plant, protein and galactomannan in seeds
of M2 generation (Chaudhary et al., 1973). A high yielding stable
early-flowering mutant of Pusa Navbahar was recorded by Rao and
Rao (1982) in population created by irradiation from 60 Co source
of X-rays (10 kR/100 Gray).
Indeterminate growth habit where vegetative and reproductive
stage occurs simultaneously is one of important reasons for low
yield of cluster bean crop. To avoid such yield loss, Singh et al. (1981)
irradiated genotype B-5-54 with gamma rays (20, 40, 60, 80, 100,
150 and 200kR) and identified a macro-mutant exhibited determinate growth habit with early flowering. Using gamma-rays (80
kR/800 Gray), ethyl methanesulfonate (EMS) (0.1%) and N-NitrosoN-methylurea (NMU) (0.01%) alone or in combination, Singh and
Aggarwal (1986) produced many mutants of PLG143 genotypes
also showed early flowering, high and long pod, better yields and
gum percentage (17%). In a mutation study with gamma rays and
EMS treatment in two cultivars (Pusa Navbahar and VRS-culture),
Babariya et al. (2008) found that apart from variability creation
mutagens also altered the correlation between traits studied hence
selection in desired direction for trait(s) of interest is be possible. Mahla et al. (2005, 2010) and Velu et al. (2007, 2012) carried
out physical (gamma-rays) and chemical (EMS) mutagenesis and
detected a gradual decrement in plant height as well as many other
agronomic traits with the increased mutagen dose in cluster bean
cultivars. Similar results were also obtained previously by Bhosale
and Kothekar (2010) in the cluster bean. They recorded increase in
the mutation frequency with increased doses of gamma rays and
chemical mutagens EMS and sodium azide (SA). Polyploidization
was also explored as means to increase the variability by Vig (1963)
and Bewal et al. (2009) however, no significant achievement was
reported.
However, work on mutation breeding of guar is limited and
only a few mutants carrying one or two useful attributes have
been obtained so far. Therefore, there is a need to carry out exten-
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S. Kumar et al. / Industrial Crops and Products 97 (2017) 639–648
sive mutation work in guar to get desirable variability for further
exploitation in breeding (Arora and Pahuja, 2008). Moreover, still
basic information regarding choice of perfect mutagen(s) and correct dose(s) need to be identified for proper application of mutation
technology for guar improvement.
2.3. Wide crosses and distant hybridization
Wild crop gene pool is a rich reservoir of rare alleles.
Wide/distant hybridization in crop plants, especially in crops with
the limiting variability, is an integral part of breeding to widen
the genetic base and to incorporate various desirable traits into
the agronomically desirable species (Singh et al., 2015). Besides,
wide hybridization is also a useful method in taxonomic and
phylogenic studies (Sandhu, 1988). Gillette (1958) alienated the
genus Cyamposis of tribe Indigofereae (Leguminosae) into three
species viz., C. tetragonoloba, C. senegalensis and C. serrata. As
per hypothesis of Hymowitz, gene pool-1A (domesticated) consists of the cultigen C. tetragonoloba and the gene pool–1 B (wild)
of C. senegalensis. Both wild relatives are identified as reservoir
for many desirable traits like tolerance to disease and drought
(Orellana, 1966; Menon, 1973), insensitivity to light and temperature (Anonymous, 1984). Thus, interspecific hybridization between
cluster bean and C. senegalensis or C. serrata can be exploited to
generate hybrids possessing early maturity, disease resistance and
photo-and thermo-insensitivity (Sandhu 1988). Distant hybridization though might have potential in genetic improvement, has so
far remained unexploited and less successful, due to incompatibility in species combinations probably because of fragile pollen
germination, abnormal pollen tube development or incompatible
interaction of pollen with stigma and style. A complete failure of
conventional wide crossing between cluster bean and C. serrata has
been revealed by Sandhu (1988) and this might be due to the unreceptiveness of stigma for foreign pollens. To make stigma receptive
Sandhu (1988) also deployed other approaches like bud pollination,
stigma/style amputation, use of organic solvents but was unsuccessful to get the stigma-pollen compatibility. Thus, in cluster bean,
interspecific hybridization is limited because of pre-and/or postfertilization barriers (Ahlawat et al., 2013a,b).
Mathiyazhagan (2007) reported failure in achieving interspecific hybridization between cultivated guar genotypes and wild
species. Very few pollen grains [5–7] of wild species were seen on
the stigma of cultivated species. Scope for interspecific hybridization is limited in cluster bean (Pathak, 2015). However, use of
two simple techniques viz., bud and stump pollination ensured
and increased the fertilization frequency. Virender (2008) exemplified five pods out of 300 crosses attempted from interspecific
crosses between HG 563 (C. tetragonoloba) and C. serrata. Similarly,
in another 250 interspecific crosses between FS 277 × C. serrata,
Virender (2008) could harvest merely three pods. To discover the
barriers of interspecific hybridization, Ahlawat et al. (2013c) formulated a study with three species of Cyamopsis and made crosses
between C. tetragonoloba × C. serrata and C. tetragonoloba × C. senegalensis. In this study, the length of the style of C. tetragonoloba
and C. serrata was almost similar (2.6 mm) though C. senegalensis
possessed longest style (3.8 mm). Studies deploying conventional
method of plant breeding revealed no pod setting in the above
said crosses. In addition to the conventional breeding method,
Ahlawat et al. (2013a,b) attempted non-conventional methods like
stub pollination with or without smearing with pollen germination medium (PGM), in-vivo placental pollination were attempted.
In this interspecific hybridization study, through stub smeared with
pollen germination medium (PGM), they found 10.43% of pod setting (83 out of 792 crosses) between C. tetragonoloba × C. serrata
cross. The low success rate in their study indicates that systematic
intense efforts to identify pre- as well as post- fertilization barriers
and future studies on sexual incompatibility phenomenon between
cultivated and wild species might aid in overcoming the failure of
interspecific hybridization.
2.4. Male sterility and heterosis breeding
Compared with soybean, cowpea and pigeonpea where malesterility has been successfully used (Saxena et al., 2010, 2015),
male sterility phenomenon was not exploited much in cluster bean
(Arora and Pahuja, 2008). This is either due to non-availability of
natural outcrossing system, or an efficient male-sterility system or
both. A number of natural mutations have been reported in guar for
male sterility (Stafford, 1989; Mittal et al., 1968; Vig, 1965). Spontaneous complete as well as partial male sterility in cluster bean was
observed by Mittal et al. (1968). He also mentioned that this trait
was inherited monogenically where pollen fertility was recorded
dominant over sterility. During induced mutagenesis, Vig (1965)
and Kinnmann et al. (1969) found that reciprocal translocations
phenomenon caused semi-sterility in guar and was responsible for
setting of fewer seeds in pod, more racemes per plant and extra
plant height than fertile plants. Later on in South Africa, Stafford
(1989) recorded a partially sterile mutant in cluster bean having rosette-type inflorescence (raceme). During the study, Stafford
(1989) also described that partial mate-sterility in cluster bean is
governed by two genes, of which one is dominant epistatic while
another is incomplete dominant. As per Stafford (1989) this partial
male-sterility (pms) was totally unique from genetic male-sterility
(ms) observed by Mittal et al. (1968).
Several workers described an extensive range of heterosis in
cluster bean (Chaudhary et al., 1981; Saini et al., 1990; Arora et al.,
1998). However, for commercial exploitation of heterosis, stable
male sterility along with complete fertility restoration is a prerequisite. Due to absence of stable source of male sterility a limited
work on heterosis breeding has been reported in cluster bean.
The opportunity of heterosis breeding is practically low in the
absence of methods for economic production of large quantities
of hybrid seed (Pathak, 2015). Owing to unavailability of stable
male sterility, efforts have been made to develop stable and operational male sterility by male gametocides or chemical hybridizing
agents (CHAs) and physical agents. Nisha and Chauhan (2006)
found benzotriazole, maleic hydrazide (MH) and ethrel (ethephon)
to be useful in induction of male sterility in cluster bean. Various concentration and treatments of all three chemicals induced
pollen sterility between 92 and 100%. Although ethrel (0.3%) was
more sensitive and induce complete sterility but associated significantly in yield reduction. Chauhan and Nisha (2006) reported that
foliar spray of benzotriazole induced 93–100% pollen sterility in
cluster bean. A comparative light and transmission electron microscopic study showed abnormal behaviour of tapetal mitochondria
in pollen caused pollen abortion. Shinde and More (2010) analysed
the effect of different mutagenic treatments on the pollen sterility while working on gamma rays mutagenesis. Gamma rays at
400 gray (40 kR) dosage were found to be most effective as induced
maximum sterility (20.83%).
3. Role of biotechnology in cluster bean improvement
3.1. Molecular markers
Success of any breeding strategy depends on the available
genetic diversity and its precise assessment. Plant breeders conventionally used morphological characters (both quantitative and
qualitative) for diversity analysis as well as for selection (Henry and
Mathur, 2005; Pathak et al., 2011b; Kumar et al., 2013). But variability assessment based on such morphological traits usually fail
S. Kumar et al. / Industrial Crops and Products 97 (2017) 639–648
to reflect accurate genetic variation and are susceptible to environmental factors, and at many times make selection efforts ineffective
(Pathak et al., 2011d; Kumar et al., 2015). Hence, for some time
DNA based molecular markers have been found suitable and convenient for the assessment of genetic diversity, cultivar/hybrid purity
test, variety/genotype identification and selecting the diverse genotypes for hybrid development. Moreover, molecular markers could
also be exploited to trace a gene of interest, before its phenotypic
expression (Lübberstedt and Varshney, 2013). Though, the reports
on extent and pattern of genetic variability especially at DNA level
in cluster bean are insufficient (Sharma et al., 2014b).
Despite variability at morpho-agronomic level, isozymes variation study through electrophoretic technique in guar germplasm
has not been employed in depth for genetic diversity analysis.
Mauria (2000) made primary attempt in cluster bean to understand
domestication through isozyme diversity in primitive landrace
accessions from India along with released cultivars from USA, and
two wild relatives (C. serrata and C. senegalensis). Later on, Brahmi
et al. (2004) made attempt to resolve the diversity in cluster bean
germplasm with allozyme markers and reported greater interpopulation diversity as compared with the overall genetic diversity
in guar. Because of environmental sensitivity, inconsistency, fewer
loci and low polymorphism of protein markers (isozymes and
allozymes), marker analysis inclined towards DNA-based marker
system. Previous studies indicated that employing DNA markers in
crop improvement can economize both time and resources (Shah
et al., 2015a). In the last decade or so, studies have been conducted for assessing genetic diversity and phylogenetic of cluster
bean using DNA markers (Table 1). Among various DNA based
marker system, Random Amplification of Polymorphic DNA (RAPD)
has been more frquently used in cluster bean for diversity analysis and to study genetic relationships (Weixin et al., 2009; Punia
et al., 2009; Pathak et al., 2011d; Rodge et al., 2012; Sharma et al.,
2013, 2014a, b; Sharma and Sharma, 2013; Kuravadi et al., 2013;
Ajit and Priyadarshani, 2013; Kumar et al., 2013; Kalaskar et al.,
2014; Patel et al., 2014). Though, RAPD markers show low reproducibility and not repeatable but have been extensively used for
assessment of genetic diversity, germplasm characterization, cultivar identification, genetic purity testing, and gene tagging. Sharma
et al. (2014b) reported that Inter-Simple Sequence Repeat (ISSR)
are more powerful than RAPD due to their higher capacity to reveal
polymorphisms and greater potential to determine intra and intergenomic diversity. Similar to other crops, for molecular diversity
analysis in cluster bean, ISSRs have been preferred over (Kuravadi
et al., 2013; Sharma et al., 2014a,b). Evaluation of cluster bean with
both RAPD and ISSR indicated the presence of substantial diversity
in cluster bean.
Though, RAPD and ISSR are relatively simple to use, highly effective and independent to prior sequence knowledge but are poor
in reproducibility and stability. Therefore, in order to resolve this
issue, Paran and Michelmore (1993) converted the RAPD markers into sequence-characterized amplified region (SCAR) marker
by molecular cloning (Cheng et al., 2015). SCAR markers, a derivation of RAPD, ISSR and Amplified fragment length polymorphism
(AFLP), are co-dominant, locus-specific and; consistent and highly
reproducible for molecular level detection (Dhawan et al., 2013;
Shah et al., 2015b). Sharma et al. (2014a) first time developed
three polymorphic and one region specific SCAR from RAPD as
well as genotype (RGC-1031) specific SCAR from ISSR to elevate
the polymorphism reproducibility and specificity of RAPD and ISSR
markers.
Evolution pressure may cause variation in the number and
arrangement pattern of nucleotides in Internal Transcribed Spacer
(ITS) regions of ribosomal DNA (rDNA; Sharma et al., 2002). Due
to easy amplification and better polymorphism, ITS spacers are
considered as versatile genetic markers for phylogenetic and diver-
643
sity analysis (Powers et al., 1997; Beltrame-Botelho et al., 2005).
Nuclear ribosomal ITS region (ITS-1 & 2; and 5.8S rDNA) primers
reported in fungi by White et al. (1990) have also been applied by
Pathak et al. (2011d) to assess the genetic diversity in cluster bean.
DNA sequence analysis of rDNA ITS demonstrated high frequencies
of single nucleotide polymorphisms in amplified conserved DNA
stretch at seven positions and indicated a close lineage of diverse
genotypes.
However, as compared to other legumes, sequence based DNA
markers especially microsatellites or simple sequence repeats
(SSRs) are inadequate in cluster bean (Kumar et al., 2015) consequently the molecular breeding efforts in cluster bean are not
implemented swiftly (Kuravadi et al., 2014). As the initial development cost of microsatellite markers is high, publically available
data base expressed sequence tags (dbESTs) is a cheaper source to
identify SSRs. Though, cluster bean is a crop with high industrial
demand, but adoption of various “omics” approaches like transcriptomics and genomics are slow; hence merely 16,476 ESTs,
submitted by Naoumkina et al. (2007), are available in the National
Center for Biotechnology Information (NCBI) database. Kuravadi
et al. (2014) and Kumar et al. (2015) mined these ESTs and identified
187 and 100 SSR markers, respectively. EST-SSR based diversity analysis of 32 cluster bean genotypes of displayed narrow
genetic diversity as mean polymorphic information content (0.13)
of markers and average dissimilarity co-efficient (0.09) was low
(Kumar et al., 2015). The high level of similarity reported by Kumar
et al. (2015) was inconsistent from earlier reports (Pathak et al.,
2010a,b; Kumar et al., 2013; Sharma et al., 2014b) which may be
because of deployment of different marker system. Thus, on the
basis of SSR based results it is advisable that there is necessity to
develop genomic SSR markers or recent high throughput marker
systems like single nucleotide polymorphism (SNP) or genotypingby-sequencing (GBS; Elshire et al., 2011), for effective deployment
of markers for cluster bean improvement.
3.2. Tissue culture, regeneration protocols and genetic
transformation
Tissue culture based technique such as micro-propagation and
genetic transformation offer a supplementary approach to marker
assisted and conventional breeding strategies for the improvement
of cluster bean. Although there are very few constraints associated
with conventional propagation methods, tissue culture approach is
still in the flow. Generally, micro-propagation is used for avoidance
of post fertilization barriers, production of large scale single gendered true-to-type plants being dioecious in nature (Kumar et al.,
2010a, 2010b), to produce plants having less germination percentage (Modi et al., 2012), to enhance secondary chemicals and, most
importantly, to establish regeneration protocol for genetic transformation as far as crop improvement is concerned (Karupussamy,
2009).
Success of in-vitro regeneration of any plant species depends
on several factors like explants and explant source, medium composition, type of hormones, media composition, culture condition,
etc (Kalia et al., 2014). In most studies, Murashige and Skoog (MS)
medium was used for cluster bean regeneration. Several attempts
have been made to establish protocol for micropropagation of cluster bean for both production and crop improvement which are
summarized in Table 2. In context to in-vitro regeneration and
tissue culture, the work on cluster bean was started by Ramulu
and Rao, (1987). Both these researchers reported many in vitro
regeneration protocols on MS and B5 media for establishment and
enhancement of callus in cluster bean (Ramulu and Rao, 1989, 1991,
1993, 1996). However, attempts for callus induction were not just
limited to unorganized mass, but reports are available on direct differentiation from cotyledonary nodes using cytokinins (Prem et al.,
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Table 1
Deployment of various DNA markers in cluster bean for diversity analysis and genetic relationships studies.
Name/number of genotypes with collection region
Country
Type of
markera
Number and list of primers
Reference
HG365, HG563, RGC936, HG75, PNB, PLG225, GG1, Naveen,
HGS854, RGC1017, RGC1022, GUAG9002, GAUG9009, Kiran,
IC102827, IC116525, IC116529, IC116874
Genotypes: 32
Regions: Rajasthan, Gujarat and Haryana
RGC 1080, RGC 1092, RGC 1038, RGC 1059, RGC 1002, RGC
1030, CAZG 6, CAZG 50, Vikas 35, WSP 50, HGS 365, HGS
02-29, HG 155-156, HGS 26-01, GAUG 9703, GAUG 9808,
GAUG 01, GAUG 13
Landraces: 29
Commercial varieties: 19
Regions: Gujarat, Rajasthan, Haryana and Delhi
IC-102828, IC-103019, IC-102853, PNB, IC-103020, HVG-2-30,
HG-2-4, HG-563, HG-2-20, RGC-471, PRT-15, HG-365
AVT-G-1, AVT-G-2, G-6, G-7, IVT-G-8, G-9, IVT-G-11, G-12,
IVT-G-12, IVT-G-14, IVT-G-16, G-17, IVT-G-18, G-19, G-20
HGS 867, RGC 471, RGC 986, HGS, 870, RGC 936, RGC 197, GG
1, HG 365, RGC 1025, HG 563, HFG 119, RGC 1003, Pusa
Navbhar, M 83, RGC 1002, HG 75, Naveen, RGC 1017, FS 277,
Kiran, HG 182, HGS 884, HG 258
India
RAPD
37 random decamers
Punia et al. (2009)
India
RAPD
OPA-16, OPP-7, OPB-12, OPP-9, OPA-14
Pathak et al. (2010b)
India
RAPD
OPA-1, OPA-4, OPA-6, OPA-11, OPA-17,
OPA-18, OPB-10, OPB-12, OPB-18
Pathak et al. (2011d)
India
RAPD
Kuravadi et al. (2013)
India
RAPD
OPA-1, OPD-12, OPM-2, OPM-12,
OPM-15, OPN-1 to 5, OPQ-9, OPU-15,
OPX-12
OPA-1 to 5, OPA-7 to 12, OPB-5
India
RAPD
OPQ-5, OPQ-7, OPQ-11
Ajit and Priyadarshani (2013)
India
RAPD
Kumar et al. (2013)
RGC 936, RGC 1002, RGC 1003, RGC 1031, RGC 1017
RGC 1017, RGC 1003, RGC 1066, RGC 1002, RGC 936, RGC 1031,
Pusa Navbhar, PNB, Neelam 51, Pusa Selection I, Swati 55
Landraces: 29
Commercial varieties: 19
Regions: Gujarat, Rajasthan, Haryana and Delhi
Number of genotypes: 30
Regions: Bahawalpur, Bhakkar, Pakpattan
India
India
RAPD
RAPD
OPD-2, OPD-3, OPD-5, OPD-7, OPD-8,
OPE-1, OPAB-13, OPAS-6, OPAA-16,
OPAS-10, OPAR-5, OPAO-3, OPAM-7,
OPAH-3, OPAF-4, OPAF-10, OPZ-9,
OPI-3, OPI-5, OPI-6, OPI-9
Numbers of primers: 15
Numbers of primers: 15
India
ISSR
UBC 808, UBC 818, UBC 820, UBC 854,
UBC 856, UBC 868, UBC 879
Kuravadi et al. (2013)
Pakistan
RAPD
Sultan et al. (2013)
Rajasthan − 14 genotypes
Haryana − 7 genotypes
Gujarat − 4 genotypes
Uttar Pradesh − 5 genotypes
Punjab − 3 genotypes
Madhya Pradesh − 2 genotypes
Numbers of genotypes: 35
Locations: Rajasthan, Haryana, Gujarat, Uttar Pradesh,
Punjab, Madhya Pradesh
Numbers of genotype: 12
M-83, RGC1066, RGC1002, C. serrata, C. senegalensis
India
RAPD
OPB-1, OPB-3, OPB-4, OPB-5, OPB-11,
OPB-14, OPB-16, OPB-17, OPB-18,
OPB-19, OPB-20, OPA-13
15 RAPD primers
India
ISSR
Numbers of primers: 10
Sharma et al. (2014b)
India
India
Numbers of primers: 19
Numbers of primers: 224
Kalaskar et al. (2014)
Kuravadi et al. (2014)
Commercial varieties: 31
Regions: Gujarat, Rajasthan, Haryana and Delhi
Numbers of genotype: 31
India
RAPD
ESTSSR
ESTSSR
ESTSSR
AFLP
Numbers of primers: 39
Kumar et al. (2015)
Numbers of primers: 17
Boghara et al. (2015)
EcoRI + A and MseI + C
Gresta et al. (2016)
Numbers of genotypes: 8
Regions: South Africa, India and the United States
India
Italy
Patel et al. (2014)
Sharma and Sharma (2013)
Sharma et al. (2013)
Sharma et al. (2014b)
a
RAPD: Random Amplification of Polymorphic DNA; ISSR: Inter-Simple Sequence Repeat; AFLP: Amplified fragment length polymorphism (AFLP); EST-SSR: Expressed
Sequence Tag-Simple Sequence Repeat.
2003) and shoot organogenesis in cluster bean via callus culture
(Prem et al., 2005). To induce endosperm callus from embryo or
cotyledon seed explants, Bhansali (2011) reported that MS media
supplemented with 2, 4-D, IAA, NAA in combination with BAP is
the most suitable. Till 2013, the work was carried out on cultivated
species but Ahlawat et al. (2013a) successfully induced callusing in
cultivated as well as two wild species viz. C. serrata and C. senegalensis using cotyledon explant. Whilst, most of the research proved
2,4-D and BAP as excellent plant growth regulators to induce callus
and regeneration, Gargi et al. (2012) showed multiple shoot induction with gibberellins along with 2,4-D and BAP as PGRs whereas,
Meghwal et al. (2014) showed that the combination of BA, kinetin
and gibberellins was most suitable for multiple shoot generation.
In order achieve to prompt shoot multiplication from cotyledonary
node explants, Ahmad and Anis (2007) and Ahmad et al. (2012) used
thidiazuron (TDZ-phenyl-urea derivatives) instead of conventional
cytokinin activity of BAP.
Plant regeneration protocol, either direct or indirect, have been
the priority to most of the researcher groups working with the
aim of crop improvement in cluster bean. Among them, Verma
et al. (2013) found regeneration with the same explant placed
on medium supplemented with various combinations of indole3-butyric acid (IBA), BAP and gibberellin and they also observed
rooting on the same medium after 10 days of shooting. They transferred these rooted shoot plantlets for the acclimatization on
cocopeat mixture and showed successful hardening.
Progress was also seen in the establishment of the regeneration protocol where Mathiyazhagan et al. (2013) reported
somatic embryogenesis on MS medium fortified with 2 mgl−1 NAA,
0.5 mgl−1 and 3gl−1 charcoal using mature embryo as an explant in
cultivated species of cluster bean. They also reported direct shoot
regeneration from cotyledonary node placed on MS medium containing 1 mgl−1 Kn, 0.5 mgl−1 and 1 mgl−1 Zeatin on both cultivated
and wild species. The information on somatic embryogenesis in
cluster bean is still absent. The race for the establishment of simple
and efficient regeneration protocol was started from the work done
by Prem et al. (2003) who tried to regenerate plant from cotyledonary node explant and showed successful plantlet formation and
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Table 2
Tissue culture protocols for culture establishment and micropropagation in cluster bean.
Objective
Explant source
Optimized/Used Media
compositiona
Results
Reference
Establishment of callus
culture
Enhancement of callus
yield
Epicotyls, hypocotyls
and cotyledons
Hypocotyls
Maximum callus with 1.5 mg/l 2, 4-D and
0.5 mg/l Kinetin
Fresh and dry weight of callus were
enhanced 3.7 and 3.9 times than control,
respectively
Ramulu and Rao (1987)
Establishment of callus
culture
Effect of different media on
establishment of callus
cultures
Study of genotypic
response to various
auxins
Direct organogenesis
Leaf
MS + IAA + NAA + 2,4D + Kinetin
B5 + 1.5 mg/l
2,4-D + 1 mg/l
Kinetin + 2.5 mg/l yeast
extract/15% coconut
milk/2.5 gm/l casein
hydrolysate
B5 + NAA + BAP
Ramulu and Rao (1989)
B5 medium supplemented with 2.5 mg/l
NAA and 1.25 mg/l BAP
MS/B5/SH/Blayde’s/White’s Highest callus frequency in B5 with
2.5 mg/l 2, 4-D and 0.75 mg/l kinetin
Ramulu and Rao (1991)
Leaf
2,4D/NAA/IAA/Picloram
Picloram was found most effective for
callus growth
Ramulu and Rao (1996)
Embryo, Cotyledons,
Cotyledonary nodes
MS + BAP + IAA
Prem et al. (2003)
Indirect organogenesis
Embryo, cotyledons,
cotyledonary nodes,
shoot tip and
hypocotyls
MS + 2,4-D + BAP
Response of wild types to
auxins
Indirect organogenesis in
wild species
Cotyledon
Auxins
Cotyledon,
cotyledonary node and
hypocotyls
cotyledon,
cotyledonary node,
embryo axis and
hypocotyl
Cotyledonary nodes
MS + B5 vitamis + 2,4D + BAP + NAA
Cotyledonary nodes showed highest
frequency of multiple shoot with 3 mg/l
BAP and 2 mg/l NAA
Embryo or cotyledonary explants on media
with 10.0 M 2,4-D and 5.0 M BAP
showed callus frequency of 82–95%
20-25 shoots per callus was obtained on
MS medium supplemented with 13.0 M
NAA and 5.0 M BAP.
C. serrata and C. senegalensis showed good
callusing exposed to 2, 4-D in the medium.
Cotyledonary explants were found best in
all tested medium.
MS + Auxins + Cytokinins
Combinations of various growth regulators
were not found suitable for the
regeneration
Deepika et al. (2014)
Green friable morphogenic callus
Highest frequency of multiple shoot
formation
Gargi et al. (2012)
Green friable morphogenic callus;
Enhanced branching and regeneration of
shoots
Meghwal et al. (2014)
MS + 2 mg/l IBA + 3 mg/l BAP + 1 mg/l GA3
showed highest percentage of shooting
(92%) and rooting (80%)
TDZ at 5.0 M showed successful
regeneration
Established somatic embryo with this
medium.
Highest (34%) shoot morphogenesis with
20 M BAP.
Verma et al. (2013)
Indirect organogenesis
Indirect organogenesis
Hypocotyls
Indirect organogenesis
Cotyledons, hypocotyls
and epicotyls
Establishment of
regeneration protocol
Cotyledons
MS + 5 M BAP + 10 M
2,4-D
MS + 10 M
BAP + 10 M Kn + 5 M
GA3
MS + 1.2 mg/l
2,4-D + 0.6 mg/l BAP
MS + 2 mg/l
BAP + 2 mg/l Kn + 1 mg/l
GA3
MS + IBA + BAP + GA3
Shoot multiplication
Cotyledonary nodes
MS + TDZ
Somatic embryogenesis
Mature embryo
In vitro regeneration
Cotyledons and
cotyledonary nodes
MS + 2 mg/l NAA + 3 g/l
Charcoal
B5 + 2 mg/l
NAA + 2 mg/l BAP
Ramulu and Rao (1993)
Prem et al. (2005)
Ahlawat et al. (2013a)
Ahlawat et al. (2013b)
Ahmad and Anis (2007);
Singh et al. (2015)
Mathiyazhagan et al.
(2013)
Sheikh et al. (2015)
a
MS: Murashige and Skoog (MS) medium; BAP: 6-Benzylaminopurine; IAA: Indole-3-acetic acid; NAA: 1-Naphthaleneacetic acid; Kn: kinetin; IBA: Indole-3-butyric acid;
GA3 : Gibberellic acid; 2, 4D: 2,4-Dichlorophenoxyacetic acid; B5: Gamborg B5 medium; TDZ: Thidiazuron.
subsequent hardening. And still the race is going on to establish “an
efficient” regeneration protocol (Sheikh et al., 2015).
Legumes are considered ‘recalcitrant’ for genetic transformation studies and regeneration of transformed plantlets remains
the major hurdle (Somers et al., 2003). Prem (1999) for the first
time performed an experiment on genetic transformation of cluster bean using two different strains of Agrobacterium tumefaciens.
Amongst the strains used, strain LBA4404 harbouring plasmids
p35SGUSINT:pAL4404gave wide spread GUS expression in 14 days
old co-cultivated explants and found to be more effective than
EHA105. In another study, using construct pBKL4 containing uidA
and nptII genes, Joersbo et al. (1999) has also transformed ˇgluoronidase and neomycin phosphotransferase genes in cluster bean
and produced stable transformants up to 3rd generation. Results of
this study showed30% selection efficiency on kanamycin sulphate
(145 mgl/l) and 1.2% of transformation efficiency with most of the
transformants containing single copy of each gene transformed.
The choice of antibiotic used is a crucial step as these can also hinder the growth of the plant being regenerated. Joersbo et al. (2001)
also made another attempt of transformation of guar where alphagalactosidase gene from Senna occidentalis was transformed. The
cDNA of the gene, isolated from senna, had 75% identical sequences
with the ␣-galactosidase expressed in germinating guar and coffee
beans. They observed reduction in galactomannan content in 30%
of transformants (although it was an era of taking cluster bean as
a model to evaluate the function of other genes of other plants,
still the race for the establishment of “an efficient” regeneration
protocol was not started).
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The transformation of soybean with ManS gene of guar localized a significant amount of gum in soybean seeds and thus offered
avenues for the production of guar gum other species which does
not have the galactomannan gum synthesizing genes (Dhugga et al.,
2004). Even though micropropagation technique can be used to
enhance secondary metabolites at large scale (Karupussamy, 2009),
no attempts were carried out to enhance galactomannan content
through such technique. Indeed, under the influence of strong
promoter, the respective gene, mannan synthase, has been transferred to Medicago truncatula and transformation was attempted to
enhance trehalose phosphate.
An important work regarding promoter, which enhances tissue specific expression of transgene, was carried out by Rasmussen
and Donaldson (2006). They isolated promoter of sucrose synthase gene from rice termed rsus3 and evaluated luciferase and
GUS activity, under its regulation, in guar endosperm. They found
the promoter very strong for endosperm tissue. Considering this
work for the enhancement of galactomannan content through over
expression of respective gene in endosperm, one can have an option
to manipulate this promoter influencing over-expression of downstream transgene. Similar promoter from guar (mannan synthase
promoter) was also isolated by Naoumkina and Dixon (2011).
4. Conclusion and future prospects
Cluster bean crop has experienced a remarkable journey from a
traditional rainfed crop grown on marginal lands mainly for food,
animal feed and fodder to a crop with various industrial usages.
This is a drought resilient high valued crop with diverse and unique
applications. Productivity in terms of yield/hectare as well gum
content is generally very low. The key reasons for low productivity is nonavailability of high yielding varieties, new sources of
genes and diverse germplasm. Moreover, cluster bean is susceptible many foliar diseases like bacterial leaf blight (Xanthomonas
axonopodis pv. Cyamopsidis), leaf spot (Alternaria spp.) and powdery mildew. Therefore, cluster bean breeding should be targeted
to increase and stabilize yield and gum production of varieties
along with inbuilt resistance against these diseases. Useful variants should also identify using intense mutagenesis. To reach to a
desirable level of productivity systematic efforts should be made
to exploit wild species of cluster bean. In spite of its high industrial
value, genomic resources for guar are insufficient. Therefore, next
generation sequencing platform need to be used for transcriptome
analysis to understand metabolic pathways of gum biosynthesis
and for development of DNA based markers to open new avenues
for molecular breeding. Special efforts need to be made to develop
protocol for direct organogenesis and methods for stable transformation to develop tailor made cluster bean plant to improve cluster
bean production and gum quality.
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Mithil Parekh
Ranbir Fougat