Abstract
Bitter gourd is an important vegetable of the family Cucurbitaceae, cultivated mainly in humid and subtropical Asia. Bitter gourd is a vegetable with immense health benefits due to the presence of medicinal compounds such as charantin, vicine, and polypeptide-p, which play essential roles in lessening blood glucose levels. Moreover, bitter gourd fruits are particularly rich in vitamin C, minerals, and carotenes. Here, an effort has been made to critically evaluate the extent of achievements during the enhancement and enactment of bitter gourd breeding programs with the use of latest technologies. Broadening the genetic base of cultivated bitter gourd varieties as a result of enrichment of existing resources by using wild species in breeding programs. Practical seed production technological know-how along with the use of the MS system (male sterility)/chemical-induced sterility procedure is nonetheless vital to cope with market demands. Superior yielding bitter gourd hybrids combining early maturity and resistance to biotic and abiotic stresses are regularly needed to cope with the challenge of bitter gourd production.
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Key words: Bitter gourd, Breeding, Genetic diversity, Genomics, Heterosis, Molecular breeding, Mutation breeding
INTRODUCTION
The domestication was believed to arrive from Africa to Brazil with the slave trade then dispersed to Central America (
Ames 1939;
Marr et al. 2004). Wild relatives of bitter gourd are green to dark blue, spiny, brief and very bitter and abundantly present in Northeastern India (
Gaikwad et al. 2008). Based on historical, scientific studies (
Chakravarty 1990;
Miniraj et al. 1993;
Walters and Decker-Walters 1988) along with molecular analyses (
Dey et al. 2006;
Singh et al. 2007;
Gaikwad et al. 2008), eastern India (Orissa, West Bengal, Assam, Jharkhand, and Bihar) may be con-sidered as the primary center of diversity. Diversity an-alysis based on simple sequence repeat markers among Asian collections of bitter gourd landraces and hybrids showed that South Asia is the center of domestication of bitter gourd (
Dhillon et al. 2016).
M. charantia is a com-mercial crop grown in India, China, Japan, Malaysia, Thailand, Philippines, Australia, Tropical Africa and South America. Here, an effort has been made to critically evalu-ate the extent of achievements during the enhancement and enactment of bitter gourd breeding programs with the use of latest technologies.
GENENTIC DIVERSITY
The importance of genetic diversity is well recognized in providing the basic raw material for selection and hy-bridization program of plant breeding and crop improve-ment. The evolution of transgressive segregants in the hybridization program largely depends on the availability of diverse germplasm in the gene pool of a crop. A great level of diversity is present in both cultivated and wild relatives of
Momordica genus for several important traits. Although interspecific crosses were not successful between these two species (
Joseph 2005), the formation of bivalents during the normal meiotic cycle suggested that these two species are phylogenetically similar (
Singh 1990). Germplasm which leads to partial fertility upon hybridization comes under the secondary gene pool. Gene pool 2 includes
M. dioica,
M. sahyadrica,
M. subangulata subsp.
renigera, and
M. cochinchinensis. Except for two
viz.,
M. cochinchinensis ×
M. dioica and
M. cochinchinensis ×
M. sahyadrica, a suc-cessful, viable seed setting was observed between all pos-sible combinations of dioecious species (
Bharathi 2010).
A lower seed setting and germination were observed when a diploid species was used as a female parent while a proper seed setting was reported when a tetraploid species was used as a female parent.
M. subangulata subsp.
renigera, a tetraploid species, showed high fruit setting when crossed with diploid species
M. sahyadrica and
M. dioica while in the reverse cross of
M. sahyadrica and
M. dioica with
M. subangulata subsp.
renigera, the fruit setting was comparatively low (
Brar et al. 2019;
Behera et al. 2020). Tertiary gene pool includes species which are distantly related to each other and gene exchange between them is impossible except by using some special techniques such as embryo rescue, somatic hybridization, bridging cross and genetic engineering. Tertiary gene pool includes
M. cymbalaria, which did not show any seed setting when hybridized with the rest of the species of the
Momordica genus (
Behera et al. 2020).
M. charantia shows a tremendous amount of diversity for various morphological and quality traits. Fruits of wild type plants are generally smaller in size, pointed at both ends and produced grey to black seeds while cultivated plants have fruits of different shapes and sizes with lengths up to 60 cm and produced large brown seeds (
Walters and Decker-Walters 1988;
Yang and Walters 1992). Significant variation was observed for earliness-related traits by
Dey et al. (2009) and
Khan et al. (2015). Diversity for ascorbic acid (60.20 mg to 122.07 mg/100 g) among bitter gourd accessions was observed in
Dey et al. (2005). Chemical and nutritional composition study of dif-ferent colour and size of bitter gourd varieties revealed that the highest carbohydrate and fiber contents were found in the light green small types (8.22 and 1.21 g) (
Krishnendu and Nandini 2016).
The critical traits for bitter gourd are earliness, number of fruits per plant, fruit length, fruit girth, number of female flowers per plant, number of days taken for first female flower emergence, fruit weight, fruit colour, bitterness, fruit yield per plant, disease and insect resistance etc. Light brown seed coat colour is recessive to deep brown (
Srivastava and Nath 1972;
Ram et al. 2006); large seed size is re-cessive to small seed size (
Srivastava and Nath 1972); white coloured epicarp is recessive to eco-friendly (
Suribabu et al. 1986;
Vahab 1989); spiny fruit is dominant over sleek (
Vahab 1989); Green fruit colour is dominant over yellow (
Hu et al. 2002) and is managed by 2 genes (
Liu et al. 2005). The bitterness of fruit is a monogenic characteristic, with much more bitterness is dominant over less (
Suribabu et al. 1986; Dalamu
et al. 2012). The monogenic inheri-tance was described for fruit colour, fruit lustre, fruit area system, and stigma colour, but a digenic form of inheri-tance was suggested for seed colour (
Kole et al. 2012).
Srivastava and Nath (1972) discovered immature fruit colour in bitter gourd was managed by one nuclear gene without any cytoplasmic element involved. The light green colours were most likely influenced by incomplete domin-ance or maybe modifier genes (
Hu et al. 2002). The con-stant perturbation in fruit length indicated its quantitative inheritance, and over 4 genes had been reported to be en-gaged in managing this particular trait (
Kumari et al. 2015). The quite short fruit length is partly dominant, and cur-viness and tubercles of fruits in bitter gourd are, governed by a single pair of a nuclear gene as well as the tubercles trait (
Kumari et al. 2015). Expression of gynoecious trait in the bitter gourd was discussed by
Iwamoto and Ishida (2006) as partly dominating in Japanese germplasms of bitter gourd while in Indian germplasms of bitter gourd
Ram et al. (2006) observed gynoecy (gy1) is, in fact, recessive to monoecy. Whereas, the genetic analysis by
Kim and Kim (1990) showed that a small fruit size was partially domin-ant to large fruit size. Complementary epistasis and domin-ance × dominance gene interaction were important deter-minants of fruit yield in bitter gourd (
Singh and Ram 2005). Inheritance of resistance to melon fruit fly indicates that fruit fly resistance is dominant over susceptibility (
Tewatia and Dhankhar 1996).
HETEROSIS BREEDING
Degree of heterosis depends on the genetic divergence between parents involved in the cross, their modes of reproduction, traits to be studied and the plant develop-mental stage. This hypothesis suggests that heterozygosity at individual loci as leading to heterosis. Heterosis is main-ly utilizing dominance variation (
Kumar et al. 2020). The exploitation of heterosis is more feasible in bitter gourd due to its cross-pollinated nature. The crucial points for the exploitation of heterosis are the identification of superior and divergent inbred lines, information on the combining ability of inbred lines and production of hybrid based on the pedigree and combining ability of inbred lines. It will be hugely beneficial to the breeder if a correlation could exist between the genetic distances of hybrid parents and the yield obtained from their respective cross or hybrid. Molecular markers can be used for the prediction of heterosis based on genetic diversity of parental lines as was demonstrated in several other crops, such as rice (
Xie et al. 2014). The preferable parameters to satisfy the consumer demands are soft fruits with a smaller number of seeds, minimized ridges, uniform green colour, high vigour, a more significant number of female flowers, good fruit setting and high yield (
Al-Mamun et al. 2015). However, consumer preference is varied depending on the region.
Light green, medium-long, spindle fruits are preferred in Vietnam, while long cylindrical and smooth fruit and light-green are preferred in Thailand. South Asian consumers like small-to-medium size, dark-green, spindle-shaped fruits with spiny exterior surfaces. In Taiwan and India, white fruit is used in soups (
Dhillon et al. 2016). A lot of variabilities were noticed for vegetative and fruit figures in bitter gourd. Due to the presence of large variability, mono-ecious characteristics, convenient and conspicuous flowers along with a substantial selection of seeds per fruit, bitter gourd work as a prospective crop for the exploitation of heterosis (
Thangamani and Pugalendhi 2013). As bitter gourd does not show or very negligible inbreeding depres-sion, the homozygous inbred lines are developed after 6 to 7 generations of continuous selfing. Before making the crosses between inbred lines directly, information regarding the general and specific combining ability will help the breeder in getting the superior hybrids (
Haripriya 1991). High general combining ability effects for different charac-teristics may help identify better parents with favourable alleles for different components of yield (
Acharya et al. 2019). Thus, a substantial general combining capability of the parents appears to be a dependable criterion for the prediction of certain combining ability (
Brar and Sidhu 1977). In bitter gourd, heterosis continues to be noticed for days to the very first female flowering, fruit per yield and vine per vine (
Vahab 1989), vine measurements, fruit mea-surements, as well as yield per vine (
Devdass 1993), fruit fat as well as yield per vine (
Richard et al. 1995), vine length, days to first female flowering, fruit length and fruit number (
Celine and Sirohi 1996), fruit flesh thickness and fruits per vine (
Jadhav et al. 2009), node at which first female flower appears and fruit length (Thangmani and Pugalendhi 2013), days to first harvest and number of fruits per plants (
Table 1) (
Al-Mamun et al. 2015).
MUTATION BREEDING
Mutation may occur spontaneously or can be induced artificially. Artificial mutations can be induced by physical mutagens such as X-rays, y-rays and neutrons, and by chemical mutagens such as ethyl methanesulfonate (EMS) (Ahloowalia and Malugzgnslia 2001). Physical mutagens are used more frequently than chemical mutagens, and among physical mutagens, y-rays are used more commonly than X-rays (
Beyaz and Yildiz 2017). Mutation breeding has become a powerful technique for developing novel plant genotypes (
Penna et al. 2012). Mutation screening is the selection of individuals from a sizeable mutated popu-lation that meet specific selection criteria, and mutant con-firmation is re-evaluating (
Oladosu et al. 2016). A Bitter gourd landrace MC 013 was treated with gamma radiation and developed a new cultivar MDU 1 which possesses improvement for yield, long greenish white fruits, toler-ance to pumpkin beetle, fruit fly and leaf spot diseases (
Rajasekharan and Shanmugavelu 1984). Similarly, a white fruited type mutant variety Pusa Do Mausami was developed from a natural population of green fruited variety Pusa Do Mausami variety (
Behera et al. 2010).
INTERSPECIFIC HYBRIDIZATION
Inter-specific and intra-specific hybridization have es-sential roles in tracing the genomic relationship and im-proving crops by transferring desirable agronomic characters and some specific traits such as disease, pests and stress resistance from wild relatives to cultivated ones (Bowely and Taylor 1987). In
Momordica, studies were carried out on inter-specific hybridization to establish phylogenetic relationships (
Vahab 1989). Crossability studies among different species of
Momordica such as
M. dioca,
M. cochinchinensis and
M. subtangulata subsp.
renigera were reported by
Bharathi et al. (2011). The hybrids were ob-served as intermediate for ovary colour, fruit shape and colour, and the number of seeds and distribution of trichomes on stem and leaves (
Hassena and Suhara 2012). High cross-ability and pollen fertility were observed between inter-varietal cross (
M. charantia var.
charantia ×
M. charantia var.
muricata).
In contrast, low crossability and moderate pollen fertility were observed in the inter-specific cross (
M. charantia ×
M. balsamina). Similarly,
Rathod et al. (2019) attempted inter-specific crossing for the characterization of plant morphology, pollen-pistil compatibility cytology and molecular relationships among parents and hybrids. Both direct and reciprocal crosses showed an approximate 90% cross-ability with a good percentage of pollen viability.
An interspecific cross resulted in a hybrid between
M. charantia variety Pusa Aushadi (female) and
M. balsamina (male) variety Pusa were selfed and backcrossed with Pusa Aushadi and
M. balsamina. Inheritance studies showed that a single dominant gene governed fruit tubercle, dis-continuous ridges and green colour of fruit, seed coat colour was governed by semi-dominant gene, and gynoecia was governed by a single recessive gene (Annual report IARI-2018-19, India,
www.iari.res.in). Crossability per-centage among interspecific crosses in the bitter gourd is presented in
Table 2.
BIOTECHNOLOGY
Traditional breeding techniques for any crop improve-ment is time-consuming. Due to the increasing importance of bitter gourd, the improvement and the development of new varieties are necessary, which could be done through the modern application of biotechnology (
Sinha et al. 2019). The application of transgenic technology through gene manipulation opens the way for producing genotypes carrying important traits such as resistance against biotic and abiotic stresses. Plant tissue cultures are used for plant breeding, commercial production and basic biological research (
Agarwal 2015). Types of explants, media com-position, growth conditions, genotypes and physiological conditions of the explants affect callus induction and plant regeneration.
Wang et al. (2008) established
in vitro plant regeneration method from cotyledon node. Overall, it was determined that age of seedlings, along with the different combination of hormones, results differently in the
in vitro morphogenesis of
M. charantia. In this direction, the effect of various growth regulators was studied by
Malik et al. (2007) on callogenesis and organogenesis of
M. charantia. It was determined that the leaf explants produced a maxi-mum callus percentage as compared to stem and cotyledons.
Among the generally used approaches of gene transfer, the
Agrobacterium-mediated gene transfer strategy is seen as most effective for the healthy integration of genes into the host plant genome.
Agrobacterium-mediated b-glucu-ronidase gene expression was recognized in explants of immature cotyledonary nodes in
M. charantia (
Sikdar et al. 2005) as well as in the leaf disc (Thriruvengadam
et al. 2012).
GYNOCISM IN BITTER GOURD
Gynoecious lines have a better genetic combining ability, and gynoecious × monoecious hybrids mature early with higher yield potential (
Dey et al. 2010). Thus, gynoecious lines can easily be utilized for the hybrid seed generation in bitter gourd as it stays away from the mechanical emas-culation and pollination. The main sex type in the bitter gourd is monoecious; however, gynoecious sex type has been reported from India, China and Japan (
Behera et al. 2006). For the very first time, bitter gourd crops with an entire phrase of gynoecious flowering habit (only pistillate blossoms on a plant) have been put in 3 gynoecious col-lections, namely Gy23, Gy63 and Gy263B (
Ram et al. 2002). And then, many populations with an extremely high pro-portion (more than 90 %) of pistillate blossoms have been designed (
Ram et al. 2002b). 2 gynoecious lines known as DBGy-202 and DBGy-201 are isolated from its vivid rela-tives
M. charantia var.
muricata L. (
Behera et al. 2006). These lines have been recognized for the inheritance of development and gynoecia of hybrid cars (
Behera et al. 2009). They hold great potential in future breeding plans for the improvement of earliness and yield in bitter gourd (
Varalakshmi et al. 2014). Gynoecium is actually under the command of an individual recessive gene (
gy 1) (
Ram et al. 2006;
Behera et al. 2009) whereas
Iwamoto and Ishida (2006) found that gynoecious sex expression in bitter gourd is partly dominant. In this direction, 2 pairs of genes were also determined by
Cui et al. (2018). Moreover, the sub-sequent decades (including F
1) utilizing gynoecious as a single parent showed an extremely high percent of pistillate (female) blossoms (
Ram et al. 2002;
Behera et al. 2006;
Iwamoto and Ishida 2006).
In Okinawa, a gynoecious line (OHB61-5) was identified, which was supposed to be a spontaneous mutant. This line, when crossed with OHB95-1A (monoecy) and F2 pro-genies studied, indicating that the trait was governed by a single recessive gene (
Matsumura et al. 2014). High yielding hybrids between gynoecious × monoecious have recently become available (Behra
et al. 2009). For example, the popular hybrid bitter gourd (VNR 28), after transplanting in the last week of March in Chhattisgarh, India in 2012, produced only pistillate flowers continuously from the first week of May through the end of May. Still, staminate flowers did not appear until the end of May (
Dhillon et al. 2017).
Similarly, Kumaken BP1, an early high yielding hybrid of bitter gourd in Japan developed using a gynoecious inbred line as the seed parent, needs a pollenizer for more initial fruit setting (
Iwamoto et al. 2009). Thus, adapted monoecious lines when blended with gynoecious × mono-ecious hybrids that produce pistillate flowers earlier than staminate flowers (
Dhillon et al. 2017).
For transgressive segregants form, NBGH-167 cultivar was isolated. The lines development through progeny row method takes four consecutive generations of selfing for obtaining complete gynoecious expression (
Jadhav et al. 2018). For the maintenance of gynoecious line, the induction of male flower is essential. For the effective induction staminate flower, the gynoecious lines were treated with silver nitrate, silver thiosulphate and Gibberellic acid. Silver nitrate (AgNO
3) performs well as compared to GA3 and silver thiosulphate [Ag(S
2O
2)
2] for induction of male flowers in gynoecious line (
Jadhav et al. 2018). Silver nitrate inhibits the synthesis of ethylene and thus induce staminate flower (
Krishnamoorthy 1975;
Saha and Behera 2015).
Identification of molecular marker associated with gynoecy trait is of great importance to determine the cost-effective hybrid seed production (
Gaikwad et al. 2014). In this direc-tion, using the F2 population from a linkage map was constructed in bitter gourd, and five single nucleotide poly-morphism (SNP) loci were found linked to gynoecy. One of the SNP markers GTFL-1 was located at 5.46 cM distance (
Matsumura et al. 2014). Similarly,
gy-1, a gynoecious gene flanked by bTP_54865, and TP_54890 markers were mapped on LG 12 in the bitter gourd genetic map (
Gangadhara Rao et al. 2018).
MOLECULAR BREEDING
Traditional plant breeding techniques are primarily de-pending on the phenotypic screening of germplasm lines. These techniques require a more extended period of time for diversity analysis, identification of desirable parents and took a long time for the improvement of essential traits. Genetic diversity grounded on quantitative characteristics has been completed in a bitter gourd by
Mishra et al. (1998),
and Ram et al. (2000). But genetic diversity among dif-ferent lines can also be done by molecular markers. Mole-cular markers are independent of environmental conditions and show a higher level of polymorphism. The initial study on diversity analysis in bitter gourd was done by
Dey et al. (2006). They evaluated 38 bitter gourd lines, including commercial cultivars collected from different parts of India using RAPD primers. Out of 116 primers, 29 were found to be polymorphic and informative.
The primary reason for mismatch might be that the majority of the quantitative traits are actually managed by a big selection of genes and are highly affected by locations (
Dey et al. 2006).
Ferriol et al. (2003) additionally did not find some correlation between morphological and RAPD characterization in strawberry. The inter-simple sequence repeat (ISSR) markers had been implemented to expose polymorphism among thirty-eight lines of
M. charantia collected from various agro-ecological zones of India. The dendrogram analysis showed that 38 genotypes were grouped into 2 major groups with 36 genotypes in one group and 2 genotypes in the second group. RAPD markers were used by
Rathod et al. (2008) for determining the genetic re-lationships among twenty genotypes of
M. charantia. The genotypes were divided into two clusters, cluster A with one genotype and cluster B with 19 genotypes. Cluster B was further divided into two sub-clusters B1 (one genotype and B2 (18 genotypes). The RAPD and ISSR marker (
Dey et al. 2006;
Singh et al. 2007) used for diversity analysis could not provide complete insights into the cultigens examined (
Gaikwad et al. 2008). The discriminating power of AFLP marker is generally higher than that of RAPD and ISSR (
Vos et al. 1995;
Powell et al. 1996) due to high polymorphic nature, broad distribution throughout the genome and high multiplex ratio (
Milbourne et al. 1997).
The genetic relationships among 38 bitter gourd lines originating from different geographical regions were studied by
Gaikwad et al. (2008). The results indicated that these lines could be used directly as parents in hybridization or as germplasm for selection to improve economically important trait (
Fazio et al. 2003;
Fan et al. 2006;
Gao et al. 2010). The genotypes were divided into three main clusters and six sub-groups using RAPD markers and three main clusters and seven subgroups based on ISSR markers. The characterization by high polymorphism, simple sequence repeats (SSRs) are ideal genetic markers and have gained signifi-cant importance in plant breeding (
Akkaya et al. 1992;
Morgante and Olivieri 1992;
Gupta et al. 1996;
Peakall et al. 1998).
Guang-guang et al. (2013) studied genetic di-versity and relationship among 50 bitter gourd varieties using 16 pairs of SSR primers. The genotypes were classi-fied into 6 groups using UPGMA methods (
Saxena et al. 2014). Since marker-assisted selection (MAS) is useful in the breeding of crops, genetic mapping of agronomically important traits directly contribute to the breeding pro-grams. The first genetic map in bitter gourd was developed
Kole et al. (2012) in the F
2 population between Taiwan White × CBM12 using AFLP markers. Subsequently,
Wang and Xiang (2013) developed a linkage map in F
2 population of Z-1-4 × 189-4-1 with the use of different kinds of mole-cular markers such as simple sequence repeat (SSR), AFLP, and sequence-related amplified polymorphism (SRAP). A list of QTLs identified for important traits in bitter gourd (
Table 3).
BITTER GOURD GENOME
The bitter gourd draft genome was generated with the inbred line, OHB3-1, by employing Illumina sequencing with an estimated size of 339 Mb (
Urasaki et al. 2017). Overall, 45,859 protein-coding genes were determined, and based on the synteny bitter gourd was determined to be more closely related to watermelon (
Citrullus lanatus) as compared to cucumber or melon. Interestingly, it was determined that trypsin-inhibitor and ribosome-inactivating genes were notable characteristics in the bitter gourd genome (
Yilmaz and Khawar 2020). These genes provide medicinal benefits properties to the bitter gourd. The bitter gourd genome is further assembled to chromosome-level, and is also available with a maximum contig N50 close to 10 Mb. The new version of bitter gourd genome based on the Nanopore long-read assembler was published with a claim of the complete assembly of the Cucurbitaceae family (
Matsumura and Urasaki 2020). Based on the preference at a particular location phenotypic variation exists in the bitter gourd cultivars, e.g., in South-East Asia, small size fruits with bitter taste are preferable. Based on the genome as-sembly, it was formulated that the South Asian varietal group diverged from the progenitors around 6000 years ago (
Matsumura et al. 2019).
CONCLUSIONS AND FUTURE PROSPECTS
Bitter gourd produces large flowers and the techniques used for the hybrid development are well established in the bitter gourd and is similar to the ones for melon and cucumber. Further inbreeding or selfing is performed for several generations to produce uniformity in the inbreds before their crossing to develop them into hybrids. More-over, there is no significant inbreeding depression in the bitter gourd; therefore, inbreds are regularly maintained by selfing. Further inbred testing is performed for their com-bining ability via employing several matting designs Line by Tester analysis, diallel analyses, etc.
It is further based on the obtained information regarding the general and specific combining abilities, most promising parents are chosen for the hybrid production. With the availability of bitter gourd genome and detailed genetic map, the selection for heterosis related alleles can be easily achieved in the bitter gourd improvement program.
Moreover, to exploit heterosis in bitter gourd, it should not be only for yield but in line with the interest of other important traits such as those related to climate change and also a mix of traits like insect pest and disease resistance. In this direction, crop wild relatives (CWR) can play an im-portant role as they are the storehouse of important genes desired for the improvement of yield as well as stress tolerance. Bitter gourd is especially vital to countries like in South East Asia where the population is still poor and cannot afford the medications for the treatment of their chronic illness. There is considerable reliance on the plants with therapeutic potential, and bitter gourd has been proved as a cure for many chronic diseases. We hope the use of bitter gourd likewise will increase in the western world and feature some significant research roads which ought to be organized. Failure to generate and provide a qualified amount of hybrid seed, specifically of general public bred hybrids warrants urgent notice.
Table 1Review of literature on magnitude of heterosis (%) over the mid parent, and better parent values for yields and its component traits in bitter gourd.
Table 1
|
Characters |
Mid parent |
Better parent |
References |
|
Vine length |
1.66 to 23.37 |
|
, Sirohi and Choudhary (1978)
|
|
2.10 to 22.30 |
|
, Singh and Joshi (1980)
|
|
4.26 to 57.81 |
|
, Chaudhari and Kale (1991b)
|
|
0.99 to 20.98 |
1.41 to 4.21 |
, Munshi and Sirohi (1993)
|
|
‒8.31 to 29.81 |
‒24.51 to 2.20 |
, Yadav et al. (2009)
|
|
‒42.56 to 111.74 |
‒45.10 to 49.08 |
, Talukdar et al. (2010)
|
|
‒6.00 to 15.90 |
|
, Behera et al. (2009)
|
|
|
‒34.60 to 30.72 |
, Talekar et al. (2013)
|
|
|
‒32.57 to 57.48 |
, Verma and Singh (2014)
|
|
Number of primary branches/vine |
7.80 to 37.00 |
|
, Singh and Joshi (1980)
|
|
3.30 to 25.85 |
|
, Singh et al. (1997)
|
|
‒30.46 to 251.77 |
‒11.21 to 293.65 |
, Yadav et al. (2009)
|
|
|
‒26.23 to 13.70 |
, Talekar et al. (2013)
|
|
‒47.37 to 182.35 |
‒50.00 to 140.00 |
, Talukdar et al. (2010)
|
|
Internodal length |
‒21.92 to 20.66 |
‒34.34 to 16.44 |
, Yadav et al. (2009)
|
|
|
‒21.13 to 20.00 |
, Talekar et al. (2013)
|
|
‒39.31 to 22.36 |
‒42.53 to 19.16 |
, Talukdar et al. (2010)
|
|
Days to opening of the first male flower |
‒8.29 to ‒26.8 |
|
, Singh et al. (2000)
|
|
‒1.09 to 12.09 |
‒7.43 to 8.51 |
, Yadav et al. (2009)
|
|
‒27.22 to 9.80 |
‒29.79 to 7.33 |
, Talukdar et al. (2010)
|
|
|
‒17.83 to 10.64 |
, Talekar et al. (2013)
|
|
Days to opening of the first female flower |
54.00 to 66.00 |
|
, Lawande and Patil (1989)
|
|
|
‒15.72 to 7.48 |
, Ram et al. (1997)
|
|
|
‒0.12 to ‒6.50 |
, Singh et al. (2001)
|
|
|
‒0.22 to ‒29.51 |
, Kandasamy (2015)
|
|
‒17.65 to 3.08 |
‒23.00 to ‒2.22 |
, Yadav et al. (2009)
|
|
|
‒19.34 to 14.47 |
, Talekar et al. (2013)
|
|
‒18.46 to 10.77 |
‒23.89 to 3.94 |
, Talukdar et al. (2010)
|
|
|
‒9.17 to 34.86 |
, Verma and Singh (2014)
|
|
Days to first fruit harvest |
‒0.42 to 15.06 |
|
Sing et al. (1997) |
|
‒6.19 to ‒22.20 |
|
, Singh et al. (2000)
|
|
|
|
, Chaubey and Ram (2004)
|
|
‒26.82 to ‒17.90 |
|
, Behera et al. (2009)
|
|
|
‒1.4 to ‒26.3 |
, Al-Mamun et al. (2015)
|
|
|
‒17.27 to 11.20 |
, Talekar et al. (2013)
|
|
‒11.91 to 4.66 |
‒16.42 to ‒0.14 |
, Talukdar et al. (2010)
|
|
‒19.80 to 3.60 |
|
, Dey et al. (2010)
|
|
|
‒19.33 to 7.69 |
, Verma and Singh (2014)
|
|
Number of fruits per plant |
0.86 to 44.44 |
0.39 to 35.02 |
, Munshi and Sirohi (1993)
|
|
|
‒66.67 to 30.61 |
, Ram et al. (1997)
|
|
|
0.0 to 104.7 |
, Al-Mamun et al. (2015)
|
|
|
‒32.23 to 22.99 |
, Talekar et al. (2013)
|
|
74.12 to 100.23 |
|
, Dey et al. (2010)
|
|
|
39.86 to 32.43 |
, Verma and Singh (2014)
|
|
Fruit length |
|
0.90 to 17.75 |
, Khattra et al. (1994)
|
|
|
1.40 to 25.46 |
, Singh et al. (2001)
|
|
31.27 to 37.00 |
|
, Behera et al. (2009)
|
|
|
‒37.7 to 6.8 |
, Al-Mamun et al. (2015)
|
|
|
1.43 to 39.79 |
, Kandasamy (2015)
|
|
‒37.57 to 4.87 |
‒30.95 to 20.76 |
, Yadav et al. (2009)
|
|
‒18.17 to 15.01 |
‒20.24 to 12.38 |
, Talukdar et al. (2010)
|
|
|
‒38.08 to 39.11 |
, Talekar et al. (2013)
|
|
4.65 to 13.80 |
|
, Dey et al. (2010)
|
|
|
29.72 to 17.99 |
, Verma and Singh (2014)
|
|
Fruit diameter |
|
‒11.7 to 13.4 |
, Al-Mamun et al. (2015)
|
|
|
1.07 to 25.34 |
, Kandasamy (2015)
|
|
20.90 to 37.60 |
|
, Behera et al. (2009)
|
|
‒24.19 to 23.88 |
‒22.73 to 39.53 |
, Yadav et al. (2009)
|
|
2.30 to 10.50 |
|
, Dey et al. (2010)
|
|
‒5.58 to 7.09 |
‒8.37 to 4.95 |
, Talukdar et al. (2010)
|
|
|
‒36.57 to 41.08 |
, Verma and Singh (2014)
|
|
Fruit weight |
5.00 to 14.50 |
|
, Dey et al. (2010)
|
|
|
‒18.60 to 9.38 |
, Verma and Singh (2014)
|
|
26.75 to 46.38 |
|
, Behera et al. (2009)
|
|
‒10.41 to 29.97 |
‒29.17 to 28.67 |
, Talukdar et al. (2010)
|
|
Flesh thickness |
|
|
, Ranpise et al. (1992)
|
|
6.13 to 16.26 |
1.47 to 6.27 |
, Celine and Sirohi (1996)
|
|
‒17.11 to 29.66 |
‒25.19 to 26.87 |
, Mallikarjunarao et al. (2018)
|
|
Fruit yield per plant |
|
4.35 to 64.28 |
, Khattra et al. (1994)
|
|
|
19.7 to 102.0 |
, Al-Mamun et al. (2015)
|
|
132.00 to 142.20 |
|
, Behera et al. (2009)
|
|
‒8.35 to 113.01 |
‒22.80 to 94.25 |
, Talukdar et al. (2010)
|
|
|
0.38 to 60.38 |
, Kandasamy (2015)
|
|
‒34.75 to 23.10 |
‒10.29 to 58.51 |
, Yadav et al. (2009)
|
|
‒30.22 to 136.43 |
‒43.64 to 98.75 |
, Mallikarjunarao et al. (2018)
|
|
|
‒58.96 to 51.14 |
, Talekar et al. (2013)
|
|
38.22 to 97.49 |
|
, Dey et al. (2010)
|
|
|
‒39.41 to 35.23 |
, Verma and Singh (2014)
|
|
Vitamin C content (mg/100 g) |
‒11.60 to 29.03 |
‒16.15 to 24.75 |
, Mallikarjunarao et al. (2018)
|
|
|
‒33.33 to 18.52 |
, Kumar and Pathak (2018)
|
|
Iron content (mg/100 g) |
‒23.43 to 52.34 |
‒34.62 to 36.97 |
, Mallikarjunarao et al. (2018)
|
|
TSS (°Bri×) |
‒23.50 to 27.37 |
‒34.44 to 20.18 |
, Mallikarjunarao et al. (2018)
|
|
Carotene content (mg/100 g) |
|
‒79.61 to 52.92 |
, Kumar and Pathak (2018)
|
|
Total sugar (g/100 g) |
|
‒57.73 to 81.13 |
, Kumar and Pathak (2018)
|
|
Reducing sugar (g/100 g) |
|
‒59.07 to 79.77 |
, Kumar and Pathak (2018)
|
Table 2Crossability percentage among interspecific crosses in bitter gourd.
Table 2
|
Inter-specific combinations |
% crossability |
References |
|
M. charantia var. charantia × M. charntia var. muricata
|
100% |
, Hassena and Suhara (2012)
|
|
M. charntia var. muricata × M. charantia var. charantia
|
88.89% |
|
|
M. charantia var. charantia × M. charntia var. muricata
|
97% |
, Bharathi et al. (2012)
|
|
M. charntia var. muricata × M. charantia var. charantia
|
85% |
|
|
M. charntia var. muricata × M. balsamina
|
6% |
|
|
M. dioca × M. sahyadrica
|
75% |
|
|
M. dioca × M. subangulata subsp. renigera
|
53% |
|
|
M. dioca × M. cochinchinensis
|
60% |
|
|
M. subangulata subsp. renigera × M. dioca
|
65% |
|
|
M. subangulata subsp. renigera × M. sahyadrica
|
84% |
|
|
M. subangulata subsp. renigera × M. cochinchinensis
|
2% |
|
|
M. sahyadrica × M. subangulata subsp. renigera
|
50% |
|
|
M. sahyadrica × M. dioca
|
81% |
|
|
M. sahyadrica × M. cochinchinensis
|
65% |
|
|
M. cochinchinensis × M. subangulata subsp. renigera
|
5% |
|
|
M. charantia var. charantia × M. charntia var. muricata
|
90.98% |
, Rathod et al. (2019)
|
|
M. charntia var. muricata × M. charantia var. charantia
|
84.43% |
|
Table 3List of QTLs identified for important traits in bitter gourd.
Table 3
|
Traits |
Number of QTLs |
Linkage group |
References |
|
Fruit colour |
1 |
LG 7 |
, Kole et al. (2012)
|
|
Seed colour |
2 |
LG 3 |
|
|
Fruit length |
2 |
LG 2 and LG 7 |
|
|
Fruit diameter |
1 |
LG 1 |
|
|
Fruit weight |
1 |
LG 1 |
|
|
Fruit number |
4 |
LG 1, LG 2 and LG 5 |
|
|
Fruit yield |
4 |
LG 1, LG 2 and LG 3 |
|
|
Sex ratio (male:female) |
9 |
LG 9, LG 13, LG 14 and LG 16 |
, Gangadhara Rao et al. (2018)
|
|
Days to first pistillate flower appearance |
8 |
LG 3, LG 5, LG 14 and LG 16 |
|
|
The node at first pistillate flower appearance |
5 |
LG 5, LG 9 and LG 14 |
|
|
Gynoecy |
1 |
LG 12 |
|
|
Female flower ratios |
3 |
LG 4, LG 5 and LG 9 |
, Wang and Xiang (2013)
|
|
First female flower node |
3 |
LG 4, LG 5 and LG 9 |
|
|
Fruit length |
4 |
LG 1, LG 2, LG 5 and LG 9 |
|
|
Fruit diameter |
5 |
LG 1, LG 9 and LG 11 |
|
|
Flesh thickness |
2 |
LG 1 |
|
|
Fruit shape |
5 |
LG 4, LG 5, LG 9 and LG 11 |
|
|
Fruit pedicel length |
3 |
LG 4, LG 8 and LG 9 |
|
|
Fruit pedicel length ratios |
5 |
LG 4, LG 6 and LG 8 |
|
|
Fruit weight |
4 |
LG 4, LG 5, LG 6 and LG 12 |
|
|
Fruit numbers per plant |
3 |
LG 1 and LG 5 |
|
|
Yield per plant |
2 |
LG 5 and LG 9 |
|
|
Stem diameter |
2 |
LG 2 and LG 4 |
|
|
Internode length |
2 |
LG 2 and LG 5 |
|
|
Gynoecy |
2 |
MC01 |
, Cui et al. (2018)
|
|
First female flower node |
2 |
MC01 |
|
|
Female flower number |
2 |
MC01 |
|
|
Fruit wart |
1 |
MC04 |
|
|
Width of ridge |
1 |
MC10 |
|
|
Hue angle |
1 |
MC10 |
|
|
Lightness variable |
1 |
MC10 |
|
|
Bitterness |
3 |
LG 3 and LG 4 |
, Shang et al. (2020)
|
References
- Acharya SK, Kaushik RA, Ameta KD, Dubey RB, Upadhyay B. 2019. Heterosis and combining ability in bitter gourd (Momordica charantia L.). Int. J. Bioassays. 8: 5692- 5711.
- Agarwal M. 2015. Tissue culture of Momordica charantia L.: A review. J. Plant Sci.. 15: 24-32.
- Ahloowalia BS, Maluszynski M. 2001. Induced mutations-A new paradigm in plant breeding. Euphytica. 118: 167-173.
- Akkaya MS, Bhagwat AA, Cregan PB. 1992. Length poly-morphisms of simple sequence repeat DNA in soybean. Genetics. 132: 1131-1139.
- Al-Mamun MH, Rashida MH, Uddin MN, Islam MR, Asaduzzaman M. 2015. Heterosis studies in bitter gourd. Int. J. Veg. Sci.. 22: 442-450.
- Ames O. 1939. Economic annuals and human cultures. Botanical Museum of Harvard University. Cambridge, MA, U.S.A..
- Behera TK, Dey SS, Munshi AD, Gaikwad AB, Pal A, Singh I. 2009. Sex inheritance and development of gynoecious hybrids in bitter gourd (Momordica charantia L.). Sci. Hortic.. 120: 130-133.
- Behera TK, Dey SS, Sirohi PS. 2006. DBGy-201 and DBGy-202: two gynoecious lines in bitter gourd (Momordica charantia L.) isolated from indigenous source. Indian J. Genet. Pl. Br.. 66: 61-62.
- Behera TK, Dey SS, Datta S, Kole C. Kole C., Matsumura H., Behera TK., 2020. Genetic resources and diversity in bitter gourd. editors. The bitter gourd gen-ome, Compendium of plant genomes. Springer Nature Switzerland AG. Switzerland: pp. 45-59.
- Beyaz R, Yildiz M. 2017. The use of gamma irradiation in plant mutation breeding. Plant Engineering. Snjezana Juric, IntecOpen.:
- Bharathi LK, Munshi AD, Behera TK, Vinod John JK, Bhat KV, Das AB, et al. 2012. Production and preliminary characterization of novel inter-specific hybrids derived from Momordica species. Curr. Sci.. 103: 178-186.
- Bharathi LK, Vinod , Munshi AD, Behera TK, Shanti-Chandrashekaran , Kattukunnel JJ, et al. 2010. Cyto-morphological evidence for segmental allopolyploid origin of Teasel gourd (Momordica subangulata subsp. renigera). Euphytica. 176: 79-85.
- Bharathi LK, Das AB, Ghosh N, Behera TK, Nak G, et al. 2011. Cytomorphological and molecular characterization of interspecific F1 hybrid of Momordica dioica Roxb. x Momordica subangulata subsp. renigera (G. Don) de Wilde. Afr. J. Agric. Res.. 6: 2982-2990.
- Bowley SR, Taylor NL. Christie BR., 1987. Introgressive hybridization. editor. Handbook of Plant Science in Agriculture. CRC Press. Boca Raton, Florida, U.S.A: pp. 23-59.
- Brar JS, Sidhu AS. 1977. Heterosis and combining ability for earliness and quality characters in watermelon (Citrullus lanatus Thunb. mansf.). II J. Res.. 14: 272-278.
- Brar NS, Saini DK, Kaushik P, Chauhan J, Kamboj NK. 2019. Directing for higher seed production in vegetables. Agronomy - Climate change & Food security. Amanullah, IntechOpen..
- Celine VA, Sirohi PS. 1996. Heterosis in bitter gourd (Momordica charantia L.). Veg. Sci.. 23: 180-185.
- Chakravarty HL. Bates DM., Robinson RW., Jeffrey C., 1990. Cucurbits of India and their role in the development of vegetable crops. editors. Biology and utilization of Cucurbitaceae. Cornell University Press. Ithaca, NY, U.S.A: pp. 325-334.
- Chaubey AK, Ram HH. 2004. Heterosis for fruit yield and its components in bitter gourd (Momordica charantia L.). Veg. Sci.. 31: 51-53.
- Chaudhari SM, Kale PN. 1991. Studies on heterosis in bitter gourd (Momordica charantia L.). Maharashtra J. Hort. 5: 45-51.
- Cui J, Luo S, Niu Y, Huang R, Wen Q, Su J, et al. 2018. A RAD-based genetic map for anchoring scaffold se-quences and identifying QTLs in bitter gourd (Momordica charantia). Front. Plant Sci.. 9: 1-10.
- Dalamu , Behera TK, Satyavati CT, Pal A. 2012. Generation mean analysis of yield related traits and inheritance of fruit colour and surface in bitter gourd. Indian J. Hortic.. 69: 65-69.
- de Wilde WJJO, Duyfjes BEE. 2002. Synopsis of Momordica (Cucurbitaceae) in South East Asia and Malaysia. Bot. Zhurn.. 57: 132-148.
- Decker-Walters DS. 1999. Cucurbits, Sanskrit, and the Indo-Aryans. Econ. Bot.. 53: 98-112.
- Degner O. 1947. Flora hawaiiensis, Fifth edition. Privately published. Honolulu, Hawaii, U.S.A: pp. 185.
- Devdass VS. 1993. Genetic studies on fruit and seed yield and quality in bitter gourd (Momordica charantia L.), Ph.D. Thesis. Tamil Nadu Agricultural University. Coimbatore, India.
- Dey SS, Behera TK, Munshi AD, Bhatia R. 2009. Genetic variability, genetic advance and heritability in bitter gourd (Momordica charantia L.). Indian Agri.. 53: 7-12.
- Dey SS, Behera TK, Munshi AD, Pal A. 2010. Gynoecious inbred with better combing ability and earliness in bitter gourd (Momordica charantia L.). Euphytica. 173: 37-47.
- Dey SS, Behera TK, Pal A, Munshi AD. 2005. Correlation and path coefficient analysis in bitter gourd (Momordica charantia L.). Veg. Sci.. 32: 173-176.
- Dey SS, Singh AK, Chandel D, Behera TK. 2006. Genetic diversity of bitter gourd (Momordica charantia L.) geno-types revealed by RAPD markers and agronomic traits. Sci. Hortic.. 109: 21-28.
- Dhillon NPS, Phethin S, Sanguansil S, McCreight JD. 2017. Early staminate flowering monoecious lines have po-tential as pollenizers for gynoecious hybrid bitter gourd cultivars. Pak. J. Agri. Sci.. 54: 27-33.
- Dhillon NPS, Sanguansil S, Schafleitner R, Wang Y, McCreight JD. 2016. Diversity among a wide Asian collection of bitter gourd landraces and their genetic relationships with commercial hybrid cultivars. J. Am. Soc. Hortic. Sci.. 141: 475-484.
- Fan Z, Robbins MD, Staub JE. 2006. Population development by phenotypic selection with subsequent marker-assisted selection for line extraction in cucumber (Cucumis sativus L.). Theor. Appl. Genet.. 112: 843-855.
- Fazio G, Chung SM, Staub JE. 2003. Comparative analysis of response to phenotypic and marker-assisted selection for multiple lateral branching in cucumber (Cucumis sativus L.). Theor. Appl. Genet.. 107: 875-883.
- Ferriol M, Pico B, Nuez F. 2003. Genetic diversity of a germplasm collection of Cucurbita pepo using SRAP and AFLP markers. Theor. Appl. Genet.. 107: 271-282.
- Gaikwad AB, Behera TK, Singh AK, Chandel D, Karihaloo JL, Staub JE. 2008. Amplified fragment length poly-morphism analysis provides strategies for improvement of bitter gourd (Momordica charantia L.). HortScience. 43: 127-133.
- Gaikwad AB, Saxena S, Behera TK, Archak S, Meshram SU. 2014. Molecular marker to identify gynoecious lines in bitter gourd. Indian J. Hortic.. 71: 142-144.
- Gangadhara Rao P, Behera TK, Gaikwad AB, Munshi AD, Jat GS, Boopalakrishnan G. 2018. Mapping and QTL analysis of gynoecy and earliness in bitter gourd (Momordica charantia L.) using genotyping-by-sequencing (GBS) technology. Front. Plant Sci.. 9: 1-14.
- Gao S, Lin BY, Xu DX, Fu RQ, Lin F, Lin YZ, et al. 2010. Genetic diversity of bitter gourd (Momordica charantia L.) based on RAPD and ISSR. J. Plant Genet. Resour.. 11: 78-83.
- Guang-guang LI, Yan-song Z, Xiang-yang LI, Zhang H, Pei-guo G, Hong-di H. 2013. Genetic diversity analysis in bitter gourd germplasm resources based on SSR mole-cular markers. J. Southern Agri.. 44: 6-11.
- Gupta PK, Balyan HS, Sharma PC, Ramesh B. 1996. Micro-satellites in plants: A new class of molecular markers. Curr. Sci.. 70: 45-54.
- Haripriya K. 1991. Heterosis and combining ability in water-melon (Citrullus lanatus Thumb. Mansf.). M.S. Thesis. Tamilnadu Agricultural University. Coimbatore, India.
- Hassena NB, Suhara SB. 2012. Characterization of intra-specific F1 hybrids of Momordica charantia L. based on morphology, cytology and palynology. Cytologia. 77: 301-310.
- Hu KL, Fu QM, Wang GP, Hu ZH. 2002. Studies on the genetics of fruit color in bitter gourd (Momordica charantia L.). China Vegetables. 6: 11-12.
- Iwamoto E, Hayashida S, Ishida T, Morita T. 2009. Breeding and seasonal adaptability of high-female F1 hybrid bitter melon (Momordica charantia L.) 'Kumken BP1' using gynoecious inbred line for the seed parent. Hort. Res. (Japan). 8: 143-147.
- Iwamoto E, Ishida T. 2006. Development of gynoecious inbred line in balsam pear (Momordica charantia L.). Hort. Res. (Japan). 5: 101-104.
- Jadhav BP, Patil MB, Patil SY, Halakude IS, Rajput JC. 2018. Development and maintenance of novel source of gy-noecy in bitter gourd (Momordica charantia). Int. J. Curr. Res.. 10: 63802-63805.
- Jadhav KA, Garad BV, Dhumal SS, Kshirsagar DB, Patil BT, Shinde KG. 2009. Heterosis in bitter gourd. (Momordica charantia L. ). Agric. Sci. Dig.. 29: 7-11.
- Joseph JK. 2005. Studies on ecogeography and genetic diversity of the genus Momordica L. in India. Ph.D. Thesis. Mahatma Gandhi University. Kottayam, Kerala, India.
- Kandasamy R. 2015. Heterosis in bitter gourd (Momordica charantia L.). Asian J. Hort.. 10: 158-160.
- Kaushik P, Andújar I, Vilanova S, Plazas M, Gramazio P, Herraiz FJ, et al. 2015. Breeding vegetables with in-creased content in bioactive phenolic acids. Molecules. 20: 18464-18481.
- Khan MH, Bhuiyan SR, Saha KC, Bhuyin MR, Ali ASMY. 2015. Variability, correlation and path co-efficient analysis of bitter gourd (Momordica charantia L.). Bangladesh J. Agr. Res.. 40: 607-618.
- Khattra AS, Singh NJ, Thakur JC. 1994. Studies on combining ability in bitter gourd. Veg. Sci.. 21: 158-162.
- Kim ZH, Kim YR. 1990. Inheritance of fruit weight in bitter-gourd (Momordica charantia L.). J. Korean Soc. Hort. Sci.. 31: 238-246.
- Kole C, Olukolu BA, Kole P, Rao VK, Bajpai A, Backiyarani S, et al. 2012. The first genetic map and positions of major fruit trait loci of bitter melon (Momordica charantia). J. Plant Sci. Mol. Breed.. 1: 1-6.
- Krishnamoorthy HN. Krishnamoorthy HN., 1975. Role of gibberellins in juvenility flowering and sex expression. editor. Gibberellins and plant growth. Wiley Eastern Limited. New Delhi, India: pp. 115-143.
- Krishnendu JR, Nandini PV. 2016. Nutritional composition of bitter gourd types (Momordica charantia L.). Int. J. Adv. Eng. Res. Sci.. 3: 96-104.
- Kumar A, Sharma V, Jain BT, Kaushik P. 2020. Heterosis breeding in eggplant (Solanum melongena L.): Gains and provocations. Plants. 9: 403
- Kumar D, Pathak M. 2018. Estimation of heterosis and combining ability for biochemical traits in bitter gourd (Momordica charantia L.). Int. J. Chem. Stud.. 6: 2579- 2585.
- Kumari M. 2015. Physiological and molecular characterization of gynoecious line and genetics of fruit traits in Bitter gourd (Momordica charantia L.). Ph.D. Thesis. Division of Vegetable Science Indian Agricultural Research In-stitute. New Delhi, India.
- Lawande KE, Patil AV. 1989. Studies on heterosis as in-fluenced by combining ability on bitter gourd. Veg. Sci.. 16: 49-55.
- Liu ZG, Long MM, Qin RY, Wang XY. 2005. Studies on genetic variation, correlation and path analysis in bitter gourd (Momordica charantia L.). Guan. Bot.. 25: 426-430.
- Malik S, Zia M, Rehman R, Chaudhary F. 2007. In vitro plant regeneration from direct and indirect organogenesis of Momordica charantia. Pak. J. Biol. Sci.. 10: 4118-4122.
- Mallikarjunarao K, Das AK, Nandi A, Baisakh B, Tripathy P, Sahu GS. 2018. Heterosis and combining ability of quality and yield of bitter gourd (Momordica charantia L.). J. Pharmacogn. Phytochem.. 7: 5-9.
- Marr KL, Mei XY, Bhattarai NK. 2004. Allozyme, morpho-logical and nutritional analysis bearing on the domesti-cation of Momordica charantia L. (Cucurbitaceae). Econ. Bot.. 58: 435-455.
- Matsumura H, Miyagi N, Taniai N, Fukushima M, Tarora K, Shudo A, et al. 2014. Mapping of the gynoecy in bitter gourd (Momordica charantia L.) using RAD-Seq an-alysis. PLoS One. 9: e87138
- Matsumura H, Hsiao MC, Toyoda A, Taniai N, Tarora K, Urasaki N, et al. 2019. Long-read bitter gourd (Momordica charantia) genome and the genomic architecture of do-mestication. bioRxiv. 822411:
- Matsumura H, Urasaki N. Kole C., Matsumura H., Behera T., 2020. Genome sequence of bitter gourd and its comparative study with other Cucurbitaceae genomes. The bitter gourd genome. Compendium of Plant Genomes, Springer, Cham, Switzerland: pp. 113-123.
- Milbourne D, Meyer R, Bradshaw JE, Baired E, Bonar N, Provan J, et al. 1997. Comparison of PCR-based marker systems for the analysis of genetic relationships in cultivated potato. Mol. Breed.. 3: 127-136.
- Miniraj N, Prasanna KP, Peter KV. Kalloo G., Bergh BO., 1993. Bitter gourd (Momordica spp.) . editors. Genetic improvement of vegetable crops. Pergamon Press. Oxford, U.K: pp. 239-246.
- Mishra HN, Mishra RS, Parhi G, Mishra N. 1998. Diallel analysis for variability in bitter gourd (Momordica charantia). Indian J. Agr. Sci.. 68: 18-20.
- Morgante M, Olivieri A. 1992. PCR-amplified microsatellites as markers in plant genetics. Plant J.. 3: 175-182.
- Munshi AD, Sirohi PS. 1993. Studies on heterosis in bitter gourd (Momordica charantia L.). Veg. Sci.. 20: 147-151.
- Oladosu Y, Rafii MY, Abdullah N, Ghazali H, Asfaliza R, Rahim HA, et al. 2016. Principle and application of plant mutagenesis in crop improvement: A review. Biotechnol. Biotechnol. Equip.. 30: 1-16.
- Peakall R, Gilmore S, Keys W, Morgante M, Rafalski A. 1998. Cross-species amplification of Soybean (Glycine max) simple sequence repeats (SSRs) within the genus and other legume genera: Implications for the transfer-ability of SSRs in plants. Mol. Biol. Evol.. 15: 1275-1287.
- Penna S, Vitthal SB, Yadav PV. 2012. In vitro mutagenesis and selection in plant tissue cultures and their prospects for crop improvement. Biorem. Biodiv. Bioavail.. 6: 6-14.
- Powell W, Morgante M, Andre C, Hanafe M, Vogel J, Tingey SV, et al. 1996. The comparison of RFLP, RAPD, AFLP and SSR (microsatellite) markers for germplasm analysis. Mol. Breed.. 2: 225-238.
- Rajasekharan KR, Shanmugavelu KG. 1984. MDU-I bitter gourd. South Indian Hortic.. 32: 47.
- Ram D, Kalloo G, Singh M. 1997. Heterosis in bitter gourd (Momordica charantia L.). Veg. Sci.. 24: 99-102.
- Ram D, Kumar S, Banerjee MK, Kalloo G. 2000. Occurrence, identification and preliminary characterization of gynoe-cium in bittergourd (Momordica charantia L.). Indian J. Agri. Sci.. 72: 348-349.
- Ram D, Kumar S, Banerjee MK, Singh B, Singh S. 2002. Developing bitter gourd (Momordica charantia L.) popu-lations with very high proportion of pistillate flowers. Cucurbit. Genet. Coop. Rep.. 25: 65-66.
- Ram D, Kumar S, Singh M, Rai M, Kalloo G. 2006. Inheri-tance of gynoecism in bitter gourd (Momordica charantia L.). J. Hered.. 97: 294-295.
- Ranpise SA, Kale PN, Deysale GY, Desai UT. 1992. Heter-osis in bitter gourd. South Indian Hort.. 40: 313-315.
- Rathod V, Behera TK, Munshi AD, Vinod , Jat GS. 2019. Crossability studies among Momordica charantia var. charantia and Momordica charantia var. muricata. Indian J. Agri. Sci.. 89: 1900-1905.
- Rathod V, Narasegowda NC, Papanna N, Simon L. 2008. Evaluation of genetic diversity and genome fingerprinting of bitter gourd genotypes (Momordica charantia L.) by morphological and RAPD markers. Int. J. Plant Breed.. 2: 79-84.
- Richard KR, Arumugam R, Kandasamy G, Suresh S. 1995. Heterosis studies in bitter gourd. Madras Agril. J.. 82: 121-123.
- Robinson RW, Decker-Walters DS. 1997. Cucurbits (CABI Publishing, Cambridge, MA).
- Saha S, Behera TK. 2015. Standardization of techniques for in vitro multiplication of gynoecious line in bitter gourd. Int. J. Plant Res.. 28: 48-53.
- Saxena S, Singh A, Archak S, Behera TK, Joahn JK, Meshram SU, et al. 2014. Development of novel simple sequence repeat markers in bitter gourd (Momordica charantia L.) through enriched genomic libraries and their utilization in analysis of genetic diversity and cross-species transfer-ability. Appl. Biochem. Biothechnol.. 175: 93-118.
- Schaefer H, Renner SS. 2010. A three-genome phylogeny of Momordica (Cucurbitaceae) suggests seven returns from dioecy to monoecy and recent long-distance dispersal to Asia. Mol. Phylogenet. Evol.. 54: 553-560.
- Shang J, Kong S, Li N, Wang J, Zhou D, Li N, et al. 2020. Genetic mapping and localization of major QTL for bitterness in melon (Cucumis melo L.). Sci. Hortic.. 266: 109286
- Sikdar B, Shafiullah M, Chowdhury AR, Sharmin N, Naha S, Joarder OI. 2005. Agrobacterium-mediated GUS expres-sion in bitter gourd (Momordica charantia L.). Biotech-nology. 4: 149-152.
- Singh AK. Bates DM., Robinson RW., Jeffrey C., 1990. Cytogenetics and evolution in the Cucur-bitaceae. editors. Biology and utilization of Cucurbitaceae. Comstock Publishing Associates, Cornell University Press. Ithaca, New York, U.S.A: pp. 10-28.
- Singh AK, Behera TK, Chandel D, Sharma P, Singh NK. 2007. Assessing genetic relationships among bitter gourd (Momordica charantia L.) accessions using inter simple sequence repeat (ISSR) markers. J. Hortic. Sci. Biotech-nol.. 82: 217-222.
- Singh AK, Pandey UB, Singh M. 2000. Studies on heterosis in bitter gourd. Veg. Sci.. 27: 158-161.
- Singh B, Joshi S. 1980. Heterosis and combining ability in bitter gourd. Indian J. Agric. Sci.. 50: 558-551.
- Singh NJ, Khattra AS, Thakur JC. 1997. Heterobeltosis in bitter gourd. Haryana J. Hort. Sci.. 26: 291-296.
- Singh NJ, Khattra AS, Thakur JC. 2001. Studies on heterosis in bitter gourd (Momordica charantia L.). Haryana J. Hort. Sci.. 30: 224-227.
- Singh SK, Ram HH. 2005. Seed quality attributes in bitter gourd (Momordica charantia L.). Seed Res.. 33: 92-95.
- Sinha S, Sandhu K, Bisht N, Naliwal T, Saini I, Kaushik P. 2019. Ascertaining the paradigm of secondary metabolism enhancement through gene level modification in thera-peutic plants. J. Young Pharm.. 11: 337-343.
- Sirohi PS, Choudhary B. 1978. Heterosis in bitter gourd (Momordica charantia L.). Veg. Sci.. 4: 107-115.
- Smith AC. 1981. Flora vitiensis nova. SB Printers. Honolulu, Hawaii, U.S.A..
- Srivastava VK, Nath P. 1972. Inheritance of some qualitative characters in Momordica charantia L. Indian J. Hortic.. 29: 319-321.
- Suribabu B, Reddy NE, Ramarao M. 1986. Inheritance of certain quantitative and qualitative characters in bitter-gourd (Momordia charantia L.), South Indian Hort. 34: 380-382.
- Talekar NS, Vaddoria MA, Kalkaran GV. 2013. Heterosis studies for quantitative traits in bitter gourd (Momordica charantia L.). Progressive Res.. 8: 650-653.
- Talukdar MB, Rahman MM, Islam MK, Rahman KAMM, Hossen MA. 2010. Estimate of heterosis in bitter gourd. J. Sher-e-Bangla Agric. Univ.. 4: 73-79.
- Tewatia AS, Dhankar BS. 1996. Inheritance of resistance to melon fruit fly (Bactroceracucurbitae) in bitter gourd (Momordica charantia L.). Indian J. Agri. Sci.. 66: 617- 620.
- Thangamani C, Pugalendhi L. 2013. Heterosis studies in bitter gourd for yield and related characters. Int. J. Veg. Sci.. 19: 119-25.
- Thiruvengadam M, Praveen N, Chung IM. 2012. An efficient Agrobacterium tumefaciens-mediated genetic transfor-mation of bitter melon (Momordica charantia L.). Aust. J. Crop Sci.. 6: 1094-1100.
- Urasaki N, Takagi H, Natsume S, Uemura A, Taniai N, Miyagi N, et al. 2017. Draft genome sequence of bitter gourd (Momordica charantia), a vegetable and medicinal plant in tropical and subtropical regions. DNA Res.. 24: 51-58.
- Vahab MA. 1989. Homeostatic analysis of components of genetic variance and inheritance of fruit colour, fruit shape, and bitterness in bitter gourd (Momordica charantia L.). Ph.D. Thesis. Kerala Agri University. India.
- Varalakshmi B, Pitachaimuthu M, Rao ES, Krishnamurthy D, Manjunath KSS. SuchithaY2014. Identification preliminary characterization and maintenance of gynoecious plants, IIHRBTGy-491 and IIHRBTGy-492 in bitter gourd In: In the international Bitter gourd Conference (BiG2014) organized by AVRDC at ICRISAT, Hyderabad in March, 36.
- Verma RS, Singh MK. 2014. Studies on heterosis for yield and its components of bitter gourd (Momordica charantia L.). Asian J. Hort.. 9: 217-223.
- Vos P, Hogers R, Bleeker M, Reijans M, van de Lee T, Hornes M, et al. 1995. AFLP: a new technique for DNA finger-printing. Nucleic Acid Res.. 23: 4407-4414.
- Walters TW, Decker-Walters DS. 1988. Balsam-pear (Momordica charantia, Cucurbitaceae). Econ. Bot.. 42: 286-292.
- Wang GL, Fan HY, Lin RY, Li XT. 2008. Establishment of in vitro regeneration system of Momordica charantia L. with different genotype. J. An. Hui Agric. Sci.. 36: 12125- 12127.
- Wang Z, Xiang C. 2013. Genetic mapping of QTLs for horti-culture traits in a F2-3 population of bitter gourd (Momordica charantia L.). Euphytica. 193: 235-250.
- Xie FM, He ZZ, Esguerra MQ, Qiu FL, Ramanathan V. 2014. Determination of heterotic groups for tropical Indica hybrid rice germplasm. Theor. Appl. Genet.. 127: 407-417.
- Yadav M, Chaudhary R, Singh DB. 2009. Heterosis in bitter gourd (Momordica charantia L.). J. Horti. Sci.. 4: 170-173.
- Yang SL, Walters TW. 1992. Ethnobotany and the economic role of the Cucurbitaceae of China. Econ. Bot.. 46: 349- 367.
- Yen DE. Allen JG., Jones R., 1977. The evidences and questions from Northwest Thailand. editors. Sunda and Sahel. Prehistoric studies in South-east Asia, Melanesia and Australia. Academic Press. London, U.K: pp. 567-599.
- Yilmaz SS, Khawar KM. Kole C., Matsumura H., Behera TK., 2020. Tissue culture, genetic en-gineering, and nanotechnology in bitter gourd. editors. The bitter gourd genome, Compendium of Plant Genomes. Springer Nature Switzerland AG. Switzerland: pp. 83-89.
