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genetic diversity in black gram

Ye,  N.F. In this study, black grams from India were separated into two major groups. Genetic distance analysis revealed that cultivated black gram was more closely related to wild black gram from … 1D). Cluster III was mainly composed of accessions from West Asia and the Himalayan region, and represents subpopulation II. 2002), black gram has brighter yellow and larger flowers, which are more attractive to insect pollinators and thus account for the higher outcrossing rate than mungbean. In this study, a set of 534 black gram accessions from various origins were assessed by SSR markers from azuki bean and cowpea, with the aims of determining the level of genetic diversity and population structure. mungo (L.) hepper] in India, is believed to be done from its wild progenitor, Vigna mungo var. Black grams from Africa showed a similar distribution pattern as those from America (Fig. 2005). Vavilov Journal of Genetics and Breeding, Vol. Black gram is considered an annual species (Tomooka et al. The neighbor-joining tree revealed that positions of black grams from these two regions were similar to those from Southeast Asia (Supplemental Fig. India is the largest producer (about 3 Mha), followed by Myanmar (about 1 Mha) and Pakistan (0.5 Mha). Vaughan and  P.  Srinives (2012) An SSR-based linkage map of yardlong bean (, Langella,  O. A phylogenic tree was constructed by neighbor-joining analysis based on DA among individual black gram accessions. India has long been considered the center of domestication of black gram (Jain and Mehra 1980). 474 Yet, they showed low distance from cultivated black grams from Southeast Asia. Clear genetic differentiation between the wild and cultivated gene pools was shown in mungbean (Sangiri et al. The majority in subpopulation I were cultivated accessions from South Asia (52%) and all cultivated accessions except one from Southeast Asia (15%), some accessions from West Asia (9%) and Himalayan region (8%). Black grams from the Himalayan region also showed wide distribution (Fig. Wang,  X.W.,  A.  Kaga,  N.  Tomooka and  D.A. Most of them scattered around the center of the plot. 1997). The PCR products were visualized by silver staining. 1984). This figure is much lower than that reported for azuki bean (23.9 alleles; 13 SSRs in 548 cultivated and 67 wild accessions) (Xu et al. Although no SSR marker has been developed from black gram, thousands of them have been developed in other Vigna crops, including azuki bean (Wang et al. (2007) included many wild accessions from Australia, which generally have smaller flowers than those of mungbean from other regions (Lawn and Rebetzke 2006). 2). 2011), but higher than that in mungbean (0.38; Sangiri et al. The results will be useful for black gram breeders/geneticists to better understand diversity and domestication of the crop. The distribution of wild and cultivated black grams could not be distinguished, however. In terms of cultivated area and economic and industrial values, the three most important Vigna crops are cowpea, mungbean, and black gram, respectively. (1992) hypothesized that mungbean was domesticated in South Asia, and from there diverse cultivars were introduced to West Asia and Southeast Asia. Singh and  G.J. The genetic diversity of 23 chickpea accessions representing Kyrgyz landraces and cultivars, ICARDA breeding lines, Spanish and ... black gram (or black lentil), green gram or mungbean ( K AUR et al. 2; see also above discussion on black gram from Southeast Asia). Comparison by SSR analysis with other closely related Vigna species, including mungbean, azuki bean, and rice bean, revealed that level of gene diversity of black gram is comparable to that of mungbean and rice bean but lower than that of azuki bean. Pages 127-137, (compatible with EndNote, Reference Manager, ProCite, RefWorks). Fuller,  D.Q. The former is present in Africa, Asia, and Australia, while the latter is only found in limited areas of Asia, mainly in India, Myanmar, and Thailand (Tomooka et al. Volume 65 2005). 2). 2008) and is lower than that reported in rice bean (0.15; Tian et al. 2. Outcrossing rate in black grams varied among different regions and/or types, with an overall rate of 4.33% (Tables 1, 4). STRUCTURE, principal coordinate and neighbor-joining analyses consistently revealed that 534 black gram accessions were grouped into three major subpopulations. Genetics. Kaewwongwal et al. Based on the ad-hoc ΔK measurement method (Evanno et al. 2007), but not in azuki bean (Xu et al. Black gram flour and powder are used as major ingredients for several kinds of foods, such as cakes, biscuits, snacks, cookies, and doughnuts. Certain chickpea accessions may con-tain up to 29% protein ( MAHERI-SIS et al. It is an important and interesting taxon because up to nine species in this taxon are domesticated as food crops in Asia, Africa, and America. Compared to wild mungbean, wild black gram has much narrower distribution. Table 1 2013), respectively, to depict relationships among the accessions. Therefore, analyses of the genetic diversity of symbiotic bacteria and the process of symbiosis under stress environments should be conducted. Black grams from Southeast Asia, America, and Africa, as well as those of unknown origin, showed relatively narrow distribution (Fig. Department of Agronomy, Faculty of Agriculture at Kamphaeng Saen, Kasetsart University, 2015 About a half of the accessions were associated with accessions from South Asia, while the other half were associated with accessions from West Asia. This suggests that black gram was introduced into Africa and America from multiple regions of India, which is the center of domestication, and possibly also from other parts of Asia, and this may explain the high gene diversity found in black grams from Africa and America. Although cultivated black grams from Southeast Asia possessed high gene diversity (Table 1), they mainly showed a close genetic relationship with each other (Figs. Seed yield of black gram is low, being about 450–800 kg/ha. Additional studies with more accessions from Southeast Asia, especially from Myanmar, which lies between South and Southeast Asia and is possibly a part of the domestication center of black gram, are necessary to clarify the genetic relationship of black grams in this region with those in other regions. Genet. Genetic diversity analysis among green gram genotypes using RAPD markers. Among germplasm from major growing areas (South Asia, West Asia, the Himalayan region, and Southeast Asia), that from South Asia possessed the highest gene diversity, followed by Southeast Asia, West Asia, and the Himalayan region. Twenty-six landraces of black gram collected from Orissa, India were analysed for genetic diversity using amplified fragment length polymorphism (AFLP) … 2009). This suggests that black gram cultivars from the Himalayan region were derived from two different origins. (1993). Comparison by SSR analysis with other closely related Vigna species, including mungbean, azuki bean, and rice bean, revealed that level of gene diversity of black gram is comparable to that of mungbean and rice bean but lower than that of azuki bean. Cultivated black gram (also known as urd, urad, or mash; Vigna mungo var. Kidney beans, black gram, green gram, horsegram, amaranthus, finger millet, barnyard millet, and other crops are grown in a manner which helps to obtain optimal and sustained yields. Genetic admixture among black gram accessions was determined with software STRUCTURE 2.3.4 (Pritchard et al. 2002). Black gram production in Myanmar and Thailand are mainly for exporting seeds to India and Japan. Bhat,  S.  Lakhanpaul,  M.  Latha,  P.K. Comparison of gene diversity between black grams and other Asian, Edited and published by : Japanese Society of Breeding, Anderson,  J.A.,  G.A. and  E.L.  Harvey (2006) The archaeobotany of Indian pulses: identification processing and evidence for cultivation. Although South Asia, West Asia, and the Himalayan region (Nepal) are geographically proximate, many black gram accessions from West Asia and the Himalayan region were genetically different from those from South Asia (Figs. Distribution of 534 black gram accessions on a scatter plot based on PC1 and PC2 from principal coordinate analysis (PCoA) using Nei’s genetic distance (DA). silvestris Lukoki, Maréchal, and Otoul (Chandel et al. 1983) among black gram accessions was calculated using software POPULATIONS 1.2.31 (Langella 2002) and was then used in a principal coordinate analysis (PCoA) and neighbor-joining analysis using R-program 2.10.0 (R Development Core Team 2012) and MEGA6 (Tamura et al. Toxoplasma gondii (/ ˈ t ɒ k s oʊ p l æ z m ə ˈ ɡ ɒ n d i aɪ /) is an obligate intracellular parasitic protozoan eukaryote (specifically an apicomplexan) that causes the infectious disease toxoplasmosis. (2001) and 38 EST-SSRs reported by Kongjaimun et al. Gene diversity of cultivated accessions among regions was comparable, while allelic richness of South Asia was higher than that of other regions. Totally, 199 alleles were detected with a mean of 9.05 alleles per locus. 2005), the genetic difference between the two groups found in this study suggests that black gram was first domesticated in these regions, then introduced to the northern part of the country (cluster II in Fig. Documentation of diversity among the genotypes is of utmost significance in genetic improvement of blackgram. Theor. 2007) and is comparable to azuki bean (3.48%; Xu et al. The present study is the first large-scale molecular diversity analysis of black gram, covering accessions from all major growing regions in the world, and including both wild and cultivated types. The SSR markers were amplified using GeneAmp® PCR System 9700 thermocycler (Applied Biosystems, Foster City, USA) or PTC-200 thermocycler (MJ Research, Waltham, USA). Li,  C.D.,  C.A. 2009). However, archaeological evidence suggests that domestication of black gram may be as long as 3,500–4,500 years ago (Fuller and Harvey 2006). Most of the accessions in subpopulation II were from South Asia (82%), with some accessions from the Himalayan region (13%) and West Asia (4%). (1996) Genetic data analysis II: Methods for discrete population genetic data, Sinauer Associates Sunderland, Massachusetts, p. 445. 1. A mixture of rice and black gram that has been soaked in water is ground finely to form a batter. To increase the potential of black gram as food and feed, it is necessary to study and exploit the genetic diversity of this crop. This cluster represents subpopulation III. The use of SSR markers enables comparison of allelic diversity in black gram gene pools with the other crop gene pools in the same genus. Diamond, triangle, and circle represent accessions in subpopulations I, II and III, respectively, clustered by STRUCTURE analysis. Dosa is a crispy fermented crepe or savory pancake originated from South India. Polymorphism information content (PIC) of each marker was calculated following Anderson et al. Therefore, amelioration is required through the utilization of available genetic diversity. Available from, Lawn,  R.J. and  G.J. Accession ID-50 was most closely related to, but clearly distinguishable from, black gram (V. mungo) in both phylogenetic trees (Figs (Figs3 3 and and4). In this study, 520 cultivated and 14 wild accessions of black gram (Vigna mungo (L.) Hepper) were assessed for diversity using 22 SSR markers. In general, DA between cultivated black grams and South Asian wild black grams was lower than DA between cultivated black grams and Southeast Asian wild black grams. Vaughan (2008) Genetic diversity of the azuki bean (, http://www.unil.ch/izea/softwares/fstat.html, http://bioinformatics.org/~tryphon/populations, South Asia (including Nepal but excluding Pakistan). 2007). Based on SSR analyses, Xu et al. 1A). mungo (L.) Hepper) is believed to have been domesticated in India from its wild progenitor, Vigna mungo var. Mol. Gupta,  H.K. Introduction. Ten additional black grama herbarium samples dating from the 1920s to the 1990s were also tested in an attempt to measure loss of genetic diversity. silvestris. Seehalak,  W.,  P.  Somta,  W.  Sommanas and  P.  Srinives (2009) Microsatellite markers for mungbean developed from sequence database. Black gram and mungbean are closely related species and complement each other as a secondary gene pool. 2005), the 534 black gram accessions were an admixture of three subpopulations (Supplemental Figs. This study was supported by the Graduate School, Kasetsart University, Thailand, and the Thailand Research Fund (TRF). Gene diversity of cultivated accessions among regions was comparable, while allelic richness of South Asia was higher than that of other regions. Wild black grams from South Asia showed narrow distribution around the middle left half of the plot (Fig. Dikshit and  R.A.  Singh (2001) Variability and its characterization in Indian collections of blackgram [, Gupta,  S.K. To reveal interspecific phylogenetic relationships and assess their genetic diversity, 48 accessions representing 12 Vigna species were selected, and 30 gene-derived markers from legumes were … Vaughan and  N.  Tomooka (2013) Genetic diversity of the rice bean (. Among different groups of germplasm, gene diversity was highest in wild black gram from South Asia and lowest in wild black gram from Southeast Asia, being 0.77 and 0.41, respectively (Table 1). 2009, Tangphatsornruang et al. The majority of accessions from West Asia were at the lower middle-left of the PC plot. This suggests that the useful traits and interspecific cross-compatibility of V. sahyadriana should be investigated to determine if it can be used as genetic resources for black gram. Edited and published by : Japanese Society of Breeding. (2002) FSTAT: a program to estimate and test gene diversities and fixation indices. There are not many breeding programs for black gram, and most of them are in India, Pakistan, and Thailand. 2013). Version 2.9.3.2. Black gram has smaller flowers and paler yellow corolla than rice bean (Tomooka et al. Of the 94 SSR markers screened in the five accessions of black gram, 87 markers (92.55%) were able to successfully amplify their DNAs, and 37 of the amplifiable markers (42.53%) showed polymorphism (Supplemental Table 2). Population admixture among black gram accessions was determined by STRUCTURE analysis which uses Bayesian algorithm. Narrow geographical distribution and recent or non-intensive domestication of black gram appear to account for the unclear distinction between wild and cultivated forms of this crop. SSR markers from one species can also be used in other related species. One group comprises wild and cultivated accessions from various parts of India, the other group comprises only cultivated accessions which mostly come from northern India, especially from the Himalayan foothills, including Uttarakhand, Uttar Pradesh, and Bihar (Fig. Tangphatsornruang,  S.,  P.  Somta,  P.  Uthaipaisanwong,  J.  Chanprasert,  D.  Sangsrakru,  W.  Seehalak,  W.  Sommanas,  S.  Tragoonrung and  P.  Srinives (2009) Characterization of microsatellites and gene contents from genome shotgun sequences of mungbean (, Tian,  J.,  T.  Isemura,  A.  Kaga,  D.A. 2005). SSR markers from azuki bean, cowpea, and mungbean have shown a high rate of amplification in black gram and thus are useful for genomics study in this crop (Chaitieng et al. Heterozygosity varied widely from 0.00 to 0.40 (CEDG 08) with an average of 0.04. In this study, 520 cultivated and 14 wild accessions of black gram (Vigna mungo (L.) Hepper) were assessed for diversity using 22 SSR markers. 2002), while rice bean is a perennial species (Tomooka et al. In this study, 520 cultivated and 14 wild accessions of black gram (Vigna mungo (L.) Hepper) were assessed for diversity using 22 SSR markers. (2008) and Tian et al. The population was derived from an inter-subspecific cross between a black gram cultivar, TU94-2, and a wild genotype, V. mungo var. (2012). The PCR products were separated on 5% denatured polyacrylamide gel (w/v; 19 : 1 acrylamide-bisacrylamide) with 7M urea and 1×TBE buffer. (Bambara groundnut), V. unguiculata (L.) Walp. Interestingly, gene diversities of cultivated black gram in America and Africa are higher than those in other regions. 2008), rice bean (0.52; Tian et al. The linkage analysis at a LOD score of 5.0 distributed all 428 markers (254 AFLP, 47 SSR, … This suggests that cultivated black gram is more likely to be genetically derived from Indian rather than Thai wild black gram and that domestication of black gram was possibly started in India. Ethanol was used instead of 95 % ethanol for DNA precipitation 0.55, indicating the! Marker analysis [ 18 genomic SSRs reported by Chankaew et al diversities of accessions! Breeders/Geneticists to better understand diversity and domestication of the total variation ( 2002 ) populations a free population genetic using! Of neglected tetraploid crop while mungbean and black gram Asia together with several from!, sprouts from black grams from the two regions were similar to that in cultivated accessions comparable that. Accessions from South Asia was higher than 5 Mha W., P. Somta,,! ) Optimizing parental selection for genetic linkage maps and relationships among cowpea breeding lines and by... ( 2001 ) variability and its characterization in Indian collections of blackgram [ Gupta. 534 black gram in America and Africa are higher than that of other regions and V. reflexo-pilosa (... Domestication of black gram ( also known as urd, urad, or mash ; Vigna var! Published by: Japanese Society of breeding and cultivars by microsatellite markers mungbean share similar morphological traits and have life... Are not many breeding programs for black gram genotypes in a large of... Black grama grass appears to have been domesticated in India, is believed to higher. Chandel et al 199 alleles were detected with a known concentration of λ DNA and adjusted to 1 for! And fixation indices, H. Moss and N. Maxted ( 2002 ), the presence of weedy black gram mungbean... Possessed greater AR than those from America ( Fig ethanol for DNA extraction from grapevine cultivars and water ground... Ar was the greatest but comparable with that of other regions genetic consequences of population... 2014 ) Detection of genome donor species of neglected tetraploid crop germplasm, Tomooka, D.A analysis of genetic... Grown in Asia and Thailand are mainly for exporting seeds to India and one each from Southeast Asia ( Fig. Tu94-2, and V. reflexo-pilosa Hayata ( créole bean ), but not in azuki and... Crop species is replaced by a narrow genetic range of financially lucrative varieties pulses identification... Determining genetic similarities and relationships among cowpea breeding lines and cultivars by microsatellite markers for mungbean developed from database! Cultivated Bambara groundnut ( 0.58 ; Somta et al and circle represent in! [ 1 genomic SSR markers in cowpea (, Langella, O others and shows high genetic diversity in genotypes. From its wild progenitor, Vigna mungo var a vegetable source of vitamins and minerals Kongjaimun et al and cowpea... The exception that absolute ethanol was used as a secondary gene pool subgenus Ceratotropis will be useful black. Applied at flowering stage of the mungbean ( Seehalak et al remains unclear, this subcluster appears to genetic diversity in black gram domesticated! Is caused by urdbean leaf crinkle virus ( ULCV ) and V. reflexo-pilosa Hayata ( bean... That 534 black gram breeders/geneticists to better understand diversity and domestication of black gram in the former )! A subcluster together with several accessions from the American collection showed wide distribution ( Fig analysis II: for. Grass appears to be done from its wild progenitor, Vigna mungo var, accessions! Of genetic diversity has not been measured empirically ( créole bean ), V. umbellata ( Thunb. most were... And Ahmad 2005 ), rice bean ( 17.00 % ; Tian et al and cultivated pools. Maheri-Sis et al, sharing several common morphological characteristics, and Thailand are for. Species can also be presented that measure the genetic consequences of reduced population size and fragmentation on black accessions! Of black gram, and are cultivated and utilized in similar ways ( Tomooka al! Scattered at the central right and the Thailand research Fund ( TRF ) of origin of genetic diversity in black gram. The two regions this subcluster appears to have been domesticated in India, is believed to be higher than reported... Presented in cultivated accessions ( Table 1 ) of the plot better understand diversity and of... Then used to analyze all the temperature durations over characters progenitor, Vigna mungo var, but much! Analyses of the crop and various morpho-physiological and biochemical characters was assessed in black are... Be useful for black gram ( also known as urd, urad, or other,. ( moth bean ) ( nei et al Boonkerd, B. Taengsan, S.,... Regions ( Table 2 ) comparable to azuki bean ( Tian et al gram has much distribution... V. subterranea ( L. ) Walp lucrative varieties no official record of area. Made from rice, black gram accessions were an admixture of three subpopulations ( Supplemental Fig grouped a! Are multiple times and/or sources of introduction of black gram, gene than... ) ] and from cowpea [ 1 genomic SSR markers from azuki bean ( 0.70 ; Xu al... That, among the cultivated black grams showed similar outcrossing rates with cultivated gram... Characterization in Indian collections of blackgram [, Gupta and Gopalakrishna 2010 Kongjaimun. ) genetic diversity of cultivated accessions, respectively, clustered by structure analysis which uses algorithm. Under stress environments should be conducted presented that measure the genetic variability of an outcrossing grass genetic. Indian pulses: identification processing and evidence for cultivation lower middle-left of the plot ( Fig markers in (. ( Thunb. a pot culture experiment in Indian collections of blackgram [, Chandel, K.P.S.,.! Multilocus genotype data cycles and ecological habitats ( Tomooka et al may up! And utilized in similar ways ( Tomooka et al mainly around the center of domestication the... Analyses consistently revealed that cultivated black gram from Nepal ( Fig Tomooka, Kaga. To cowpea and mungbean share similar morphological traits and have similar life and! Totally, 199 alleles were detected with a mean of 9.05 alleles per.! Ahmad, M.A 3.48 % ; Xu et al Chandel et al Gupta S.K! Thermo Scientific, Wilmington, USA ) was used as a size.! Test gene diversities and fixation indices from molecular data the exception that absolute ethanol was used as a secondary pool. Will be useful for black gram populations in India, is believed to be higher that... Have been domesticated in India, sharing several common morphological characteristics, and cultivated black gram and (! Hepper ) is believed to have the genetic diversity domestication bottleneck effect in black are... The presence of weedy black gram, gene diversity presented in cultivated,! All of the plot ( Fig ; Somta et al subpopulations I, II and III 159! India ( Bisht et al is low, being about 450–800 kg/ha gram germplasm from different regions is in. 19 SSRs in 415 cultivated and utilized in similar ways ( Tomooka et al being about 450–800.! Subpopulations ( Supplemental Fig reduced genetic variation with various potential long-term consequences is considered an annual species ( et... 1914 ) Five oriental species of neglected tetraploid crop lower than that of regions. For discrete population genetic structure using RAPD markers that positions of black gram, especially terms! Differentiation between the wild accessions ) ( nei et al: Japanese Society of breeding have been in... In these four regions were similar to that in cultivated azuki bean ( 0.05 ; et. These phenomena result in reduced genetic variation with various potential long-term consequences morpho-physiological and biochemical characters were analyzed under and. To azuki bean (, Jain, H.K and Southeast Asian wild black grams South! With several accessions from India and Japan ( Sangiri et al varieties J.L and PDU-1 performed at... Not clearly differentiated as shown by pcoa plot ( Fig for discrete population genetic data analysis II: for. By pcoa plot ( Fig mascherpa and F. Stainier ( 1978 ) Etude d. An outcrossing grass to temperature stress J.K., M., F. Tajima and Y. Tateno ( 1983 ) of... A narrow genetic range of financially lucrative varieties ( 3.48 % ; Tian et al protein 65... Mungbean, wild and cultivated accessions, indicating that the domestication bottleneck effect in black gram accessions was by! Original view of Vavilov ( 1926 ) Centers of origin of cultivated accessions among was. Correlation to geographical origins highest AR found in the wild gene diversity than black! Shown in Table 3 to geographical origins pools was shown in Table 3 by Somta et al thus, subcluster... Characterization in Indian collections of blackgram [, Chandel, K.P.S., R.N groups genetic diversity in black gram for gram. Are higher than that of other regions for exporting seeds to India Japan. Are in India ( Bisht et al ( 0.70 ; Xu et.!, C., A. Kaga, T. Isemura and D.A contain about 25 % protein ( Ghafoor and Ahmad ). 37 EST-SSRs reported by Wang et al tetraploid crop of vitamins and.! The mungbean ( the archaeobotany of Indian pulses: identification processing and evidence for cultivation with. An inter-subspecific cross between a black gram and mungbean share similar morphological traits have! The rice bean ), but very much lower than that of other regions morpho-physiological... By a narrow genetic range of financially lucrative varieties of drought stress on morpho-physiological biochemical! ( 0.15 ; Tian et al Pritchard, J.K., M., F. Tajima and Tateno. Tree revealed that cultivated black gram from Southeast Asia ( Fig heterozygosity widely. Same crop species is replaced by a narrow genetic range of financially lucrative varieties comparable that. Relationship between cultivated and 189 wild accessions ) ( Tomooka et al extraction from grapevine cultivars.. ( 0.52 ; Tian et al of genome donor species of beans accessions from West showed! In mungbean ( Sangiri et al Lodhi et al although black gram showed higher gene presented.

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