Indian Journal of Agricultural Sciences 79 (8):575-86 August 2009 Genetic diversity in jute (Corchorus spp) and its utilization: a review C S KAR1, A KUNDU2, D SARKAR3, M K SINHA4 and B S MAHAPATRA5 Central Research Institute for Jute and Allied Fibres, Barrackpore, Kolkata, West Bengal 700 120 Received: 25 January 2009 ABSTRACT Natural fibres of commercial importance are obtained from the bark of two jute species (Corchorus capsularis L. and C. olitorius L.), and they are cultivated in different south-east Asian countries including India and Bangladesh. High-yielding varieties of both species evolved around a very narrow genetic base, supported by pedigree relationship, morphological analysis including distinctness, uniformity and stability (DUS) testing and molecular marker analysis. A number of microsatellite markers developed recently has shown sufficient transferability between these two species and effectiveness in elucidating polymorphism. Molecular analysis of germplasm and released elite varieties using different marker systems indicated more narrowness of C. capsularis as compared to C. olitorius. The differentiation and separation of both species in the evolutionary pathway may not be recent, although cultivation for fibre use has a short history. Intra-specific hybridization utilizing diverse genotypes and inter-specific hybridization between C. capsularis/ C. olitorius and wild species having quality fibre characteristics and resistance to biotic and abiotic stresses should be emphasized to widen the narrow genetic base of both species. Key words: Corchorus, Diversity, Germplasm, Hybridization, Inter-specific, Jute, Microsatellite, Molecular marker Jute comprises two cultivated species of the genus Corchorus, viz C. capsularis L. and C. olitorius L. Roxburg during the last part of 18th century, identified jute (Corchorus spp; 2n = 2x = 14) as an alternative to European hemp (Cannabis sativa L.) (Ghosh 1983). The cultivation of jute dated back about 200 years ago in the tropics having high rainfall and availability of retting water. As jute was domesticated recently, many mutants have not yet been accumulated in endemic Corchorus population resulting from little human selection pressure in course of evolutionary history. Both cultivated species are distinguished by growth habitat, branching habit and leaf, flower, fruit, seed, bast fibre characteristics and photosensitivity. However, they are characterized by narrow genetic variability for adaptability to not only various agronomic environments, but also fibre yield, quality and susceptibility to diseases and pests. In Indian subcontinent, jute is the second most important natural fibre crop after cotton (Gossypium hirsutum L. and G. arboreum L.). For manufacturing diversified value-added industrial products and packaging materials, jute has a great potential. It is largely a self-pollinated crop, however, its natural genetic variability is very narrow making it difficult for the plant breeders to develop new varieties through conventional breeding. In this context, it is to be noted that plant breeding essentially leads to qualitative rather than quantitative shifts in registered cultivars. Besides, germplasm collections are required to be optimized for maximal redundancy with regard to genotypes, gene complexes or possibly even genes. In view of above considerations, jute breeders have become more aware of the needs of maintaining genetic diversity among varieties and improving the management of genetic resources through the conservation of traditional land races and germplasm. These populations exist with high genetic variability and fitness to the natural and anthropological environments where they have been originated and domesticated (Brandolini 1969). They represent not only a valuable source of potentially useful traits, such as resistance or tolerance to biotic and abiotic stresses, but also an irreplaceable repository of highly co-adapted genotypes. However, this crop demands the immediate attention of plant breeders. The available elite cultivars are essentially derived either through pure line selection or hybridization, followed by selection from a few common accessions. Earlier reports indicated that limited success was obtained in crossing between two cultivated jute species, possibly because of the presence of a strong sexual incompatibility barrier (Patel and 1 Scientist(SS) (E-mail: chandanskar@rediffmail.com); Associate (E-mail: kundu_avi@rediffmail.com); 3 Principal Scientist (Biotechnology) (E-mail: debabrata_s @yahoo.com); 4 Principal Scientist, Incharge, All India Network Project on Jute and Allied Fibres (E-mail: mohitsinha48 @hotmail.com); 5 Director 2 Research 3 576 KAR ET AL. Datta 1960, Swaminathan et al. 1961). Therefore, valuable genetic variability from diverse germplasm sources either at intra- or inter-specific levels should be utilized. [Indian Journal of Agricultural Sciences 79 (8) and ‘JRC 212’ and C. olitorius (tossa jute) cv ‘JRO 632’ increased the fibre yield from 1.0 to 1.25 tonnes/ha during the sixties (Saha and Hazra 2008). White jute varieties occupied more than 80% of the jute growing areas till early seventies because they were found suitable for mid-March to mid-April sowing and thus fit for the rice-based multiple cropping sequences. Tossa jute cv ‘JRO 632’, despite being higher yielder, suffered from pre-matured flowering when sown earlier. Subsequently, tossa jute cvs ‘JRO 878’, ‘JRO 7835’ and ‘JRO 524’ resistant to pre-mature flowering were released in 1967, 1971 and 1977, respectively after successfully transferring the early flowering resistance gene from an exotic germplasm ‘Sudan Green’ to Indian elite cvs ‘JRO 620’ and ‘JRO 632’. This resulted in a paradigm shift in jute agriculture as the entire jute growing area of the country was taken over by the tossa jute. These olitorius varieties, particularly ‘JRO 524’, enabled jute to be fitted in the multiple cropping sequences with transplanted Aman paddy mainly because of their shorter duration (120 days) and suitability for mid-March sowing (Das and Maiti 1998). Now a series of new olitorius varieties, like ‘JRO 66’, ‘JRO 8432’, ‘JRO 128’, ‘S 19’ and capsularis varieties, viz ‘JRC 698’, ‘JRC 80’ are available for cultivation. These varieties have wider adaptability to varied agro-climatic situations and were responsible for the quantum jump realized in average fibre yield (2.3 tonnes/ha) at national level. Besides olitorius varieties ‘JBO 2003-H’, ‘JRO 204’ and ‘AUOJ-1’ with higher yield and better fibre quality were released for cultivation in the tossa jute belts. The characteristic features of improved jute varieties are well-documented (Kundu et al. 1959, Ghosh 1983, Basak 1993, Chaudhury et al. 2004, Karmakar et al. 2008). The genetic base of elite jute varieties is quite narrow. The 14 olitorius and 13 capsularis varieties released till date have been developed from 14 and 24 parental accessions, respectively, which make the varieties susceptible to various biotic and abiotic stresses. However, CRIJAF has a collection of 2 899 germplasm accessions belonging to jute only. Many of the germplasm possess desirable traits of tolerance to biotic and abiotic stresses as well as improved fibre quality like fineness, strength, etc. The germplasm available in CRIJAF Gene Bank are in the process of rigorous multilocation field screening, and are being used in the breeding programme to develop suitable jute varieties with desirable agronomic traits. GENETIC DIVERSITY ANALYSIS IN JUTE Germplasm status The Indian Council of Agricultural Research, New Delhi has been maintaining more than 1 450 C. olitorius and about 830 C. capsularis accessions of the International Jute Organization (IJO) since 1993 (Palit et al. 1996). In Bangladesh, the largest gene bank of jute and allied fibres (JAFs) is being maintained at Bangladesh Jute Research Institute (BJRI), with 5 936 accessions comprising 15 species of Corchorus, 22 species of Hibiscus and 15 of allied genera, yet to be characterized (Haque 2007). In India, during 1999–2004, 655 accessions including land races and wild relatives were collected from different agroecological regions. Presently, CRIJAF is maintaining 939 accessions of C. capsularis, 1 647 accessions of C. olitorius and 313 wild species covering eight species (Mahapatra and Saha 2008). Morphological analysis A broad approach using phenotypic and molecular markers is required to analyze diversity and to support conservation, management and development of plant genetic resources (Hammer et al. 1999). Phenotypic markers are of great value in studies of crop evolution (Gepts P 1993), germplasm evaluation (eg Bretting and Wildrlechner 1995) and in revealing differences between varieties (Gilliland et al. 2000). Estimation of genetic diversity and genetic advance of 192 C. olitorius and 216 C. capsularis accessions of IJO, inclusive of certain induced mutants of Indian cultivars has been reported (Palit et al. 1996). This estimation was based on 4 morpho-physiological attributes, such as plant height, harvest index, cambial activity and fibre strength that are related to yield characters. Because estimations of genetic variability based on morphological traits have the disadvantages of being influenced by both environmental and genetic factors, morphological analysis may not provide proper estimation of genetic diversity. Varietal development in India Efforts for varietal development in jute were initiated in the early part of twentieth century. C. olitorius cv ‘Chinsurah Green’ and C. capsularis cv ‘Dhaka 154’ (D 154) were released during 1915 and 1919, respectively, through pure line selection (Patel and Ghosh 1940). Till date 27 improved jute varieties (14 in C. olitorius and 13 in C. capsularis) have been developed in India. The details of these varieties are given in Tables 1, 2. Plant height and base diameter were identified as the two major criteria for selection as these traits are highly correlated with fibre yield (Kundu et al. 1959). Development of C. capsularis (white jute) cvs ‘JRC 321’ Pedigree relationship among released varieties Pedigree relationship of 14 olitorius varieties released so far in India shows narrow genetic base utilized in developing these varieties. One variety was developed by pureline selection (‘JRO 632’), 12 by hybridization, followed by selection and one by mutation breeding (Fig 1). The two exotic introductions, ‘Sudan Green’ and ‘Tanganyika 1’ were one of the parents for developing 5 varieties each including 4 5 Jute Research Station, Kendrapara, Orissa CRIJAF, Barrackpore, West Bengal ‘KOM 62’ (‘Rebati’) ‘JRO 66’ (‘Golden Jubilee Tossa’) CRIJAF, Barrackpore, West Bengal ‘JRO 524’ (‘Navin’) CRIJAF, Barrackpore, West Bengal CRIJAF, Barrackpore, West Bengal ‘JRO 7835’ (‘Basudev’) ‘JRO 3690’ (‘Savitri’) CRIJAF, Barrackpore, West Bengal ‘JRO 878’ (‘Chaitali Tossa’) BARC, Trombay, Maharashtra CRIJAF, Barrackpore, West Bengal ‘JRO 632’ (‘Baisakhi Tossa’) ‘TJ 40’ (‘Mahadev’) Developing institute Variety 1998 1992 1985 1983 1977 1971 1967 1954 Year of release Multiple crosses involving six parents Gamma-ray derivative of ‘JRO 878’ Tobacco leaf Long internode Selection from intermutant from cross ‘Sudan Green’ × ‘JRO 632’ ‘JRO 632’ בSudan Green’ ‘JRO 620’ בSudan Green’ Selection from indigenous type Parentage Full green Full green Full green Full green Full green Full green Red Full green 180–200 130–140 Resistant to premature 180–200 flowering on mid-March sowing, resistant to root rot and yellow mite, better retting and easy fibre extraction Steel-grey, non-dehiscent on maturity Blackish grey, non-dehiscent on maturity Dehiscent on maturity, steelgrey Susceptible to premature flowering if sown before mid-April, fibre quality TD2 grade Susceptible to premature flowering if sown before mid-April, suitable for early sowing Susceptible to premature flowering if sown before mid-April, better fibre quality, suitable for late sowing 145–155 135–145 130–140 240–250 240–250 220–240 × 220–240 260–280 260–280 260–280 220–240 3.0–3.2 Fibre yield (tonnes /ha) 3.0–33 3.0–3.5 Mid April DIVERSITY IN JUTE AND ITS UTILIZATION (Continued) 3.5–4.0 Mid March to 3.0–3.5 late April Mid April to end April Mid April to end April Mid March to 3.4–3.6 end April End April 3.2–3.4 Mid March to end April Mid March to 3.0–3.2 end April Mid April to end April Days to Seed to seed Optimum 50% flower maturity sowing (days) (days) time Resistant to premature 180–200 sowing, flowering on mid-March waterlogging tolerance at late growth stage Resistant to premature flowering on mid-March sowing, better fibre fineness Susceptible to premature flowering if sown before midApril, suitable for late sowing Salient features Dehiscent on Susceptible to premature 140–150 maturity, brownish- flowering if sown before grey mid-April, better fibre quality Non-dehiscent on maturity, blackish grey Non-dehiscent on maturity, blackishgrey Non-dehiscent on maturity, blackish-grey Dehiscent on maturity, brownish-grey Stem and Pod shattering foliage and seed coat colour colour Table 1 Details of released varieties of tossa jute (Corchorus olitorius L.) in India August 2009] 577 CRIJAF, Barrackpore, West Bengal ‘JRO 204’ 2007 2007 2005 CRIJAF, 2008 Barrackpore, West Bengal CRIJAF, Barrackpore, West Bengal ‘S 19’ (‘Subala’) 2002 ‘JBO 2003 H’ (‘Ira’) CRIJAF, Barrackpore, West Bengal ‘JRO 128’ (‘Surya’) 1999 AAU, Jhorhat, Assam CRIJAF, Barrackpore, West Bengal ‘JRO 8432’ (‘Shakti’) Year of release ‘AAU OJ 1’ (‘Tarun’) Developing institute Variety (Table 1 concluded...) 6 (‘JRO 632’ × Full ‘Sudan green Green’) × ‘Tanganyika 1’ Pure line Full selection green between ‘Tanganyika 1’× ‘JRO 640’ Pure line Full selection green between ‘IDN/SU/053’ × ‘KEN/DS/060’ (’JRO620’ × Light ‘Sudan red Green’) × ‘Tanganyika 1’ ‘TJ 6’ × Full ‘Tanganyika I’ green Non-shattering pod Non-shattering pod Non-shattering pod Steel-grey, non-dehiscent on maturity Steel-grey, non-dehiscent on maturity Steel-grey, non-dehiscent on maturity Stem and Pod shattering foliage and seed coat colour colour ‘IC 15901’ × Full ‘Tanganyika I’ green Parentage Resistant to premature flowering, better biotic resistance to stem rot, root rot, anthracnose yellow mite Resistant to premature flowering, better biotic resistance to stem rot, root rot, anthracnose yellow mite Resistant to premature flowering, non-lodging tall cylindrical stem Resistant to premature less flowering on mid-March sowing, finer fibre quality with lignin content Resistant to premature flowering on mid-March sowing, good fibre quality Resistant to premature flowering on mid-March sowing Salient features 120 120 120 170–180 170–180 170–180 230–250 230–250 240–250 260–270 260–270 260–270 Fibre yield (tonnes /ha) Last week of March to end April Mid March to early May 1st week of March to end April 3.6 3.6 3.6–3.8 Mid March to 3.5–4.0 end April Mid March to 3.5–4.0 end April Mid March to 3.5–4.0 late April Days to Seed to seed Optimum 50% flower maturity sowing (days) (days) time 578 KAR ET AL. [Indian Journal of Agricultural Sciences 79 (8) 7 CRIJAF, Barrackpore, West Bengal Jute Research Station, Kendrapara, Orissa ‘JRC 698’ CRIJAF, (‘Shrabanti’) Barrackpore, West Bengal ‘Bidhan B C KV V, Pat 3’ Kalyani, West Bengal ‘Bidhan B C KV V, Pat l’ Kalyani, West Bengal ‘Bidhan B C K V V, Pat 2’ Kalyani, West Bengal ‘JRC 80’ CRIJAF, (‘Metali Barrackpore, White’) West Bengal ‘KC I’ (‘Jaydev’) ‘Padma’ ‘UPC 94’ (‘Reshma’) ‘JRC 4444’ (‘Baldev’) CRIJAF, 1954 Barrackpore, West Bengal CRIJAF, 1971 Barrackpore, West Bengal CRIJAF, 1980 Barrackpore, West Bengal Jute Research 1983 Station Bahraich, Uttar Pradesh ‘JRC 212’ (‘Sabuj Sona’) ‘JRC 7447’ (‘Shyamali’) Multiple crosses involving13 parents ‘D 154’× ‘D 18’(mutant) Gamma-ray derivatives of ‘D 154’ ‘D 154’× ‘D 18’ (mutant) ‘CIN 114’× ‘JRC 321’ 1999 2001 2005 2001 2000 Gammaray derivative of ‘JRC 4444’ ‘JRC 6165’× ‘JRC 412’ ‘JRC 321’× ‘JRC 212’ ‘JRC 212’× ‘D 154’ X-ray derivative of ‘JRC 212’ Selection from indigenous type Selection from indigenous type ‘Hewti’ Parentage 1992 1983 1954 CRIJAF, Barrackpore, West Bengal ‘JRC 321’ (‘Sonali’) Year of release Developing institute Variety Full green Full green Full green FuII green Full green Stem and upper surface of leave foliage coppery red Stem and upper surface of leave foliage coppery red. Full green Full green Full green Stem and upper surface of leaf petiole coppery red. Full green Stem and leaf colour Chocolate Chocolate Chocolate Chocolate Shining chocolate Shining chocolate Shining chocolate Shining chocolate Shining chocolate Shining chocolate Shining chocolate Shining chocolate Seed coat Table 2 Details of released varieties of white jute (Corchorus capsularis L.) in India Non-dehiscent Non-dehiscent Non-dehiscent Non-dehiscent Non-dehiscent Non-dehiscent Non-dehiscent Non-dehiscent Non-dehiscent Non-dehiscent Non-dehiscent Non-dehiscent Pod shattering habit 130–140 150–160 150–160 150–160 120–130 90–95 65–70 110–115 120—130 Early-March sowing does 110–120 days not induce premature flowering Photo-period insensitive Photo-period insensitive Early-March sowing does not induce premature flowering Photo-period insensitive Early-March sowing 140–150 does not induce premature flowering 200–210 170–180 140–150 180–190 210–220 210–220 210–220 210–220 210–225 225–235 220–230 200–210 30–35 20–23 13–14 25–27 30–35 26–33 25–28 25–27 30–32 28–30 25–28 20–25 Days to Seed to seed Fibre 50% flower maturity yield (days) (days) (tonnes /ha) Late-February sowing 140–150 does not induce premature flowering Mid March sowing does not induce premature flowering Mid March sowing does not induce premature flowering Early-March sowing does not induce premature flowering Late-February sowing does not induce premature flowering Late February sowing does not induce premature flowering Flowering behaviour August 2009] DIVERSITY IN JUTE AND ITS UTILIZATION 579 580 KAR ET AL. JRO632 [Indian Journal of Agricultural Sciences 79 (8) Indigenous Hewti JRO7835 JRO128 JRO524 Indigenous type Selection Selection JRC6165 Tarun JR0878 CIN114 JRC212 JRC321 S-19 JRC80 UPC94 IC15901 JBO-2003-H Tobacco leaf Multiple cross 13 parents Tanganyika1 Sudan Green JR0632 JR0620 JR0640 TJG IC30730 Long intemode IDN/SU/053 KEN/DS/060 selection JRC6165 JRO8432 D154 D18 (mutant) JRC4444 Bidhan Pat3 X-ray mutation Bidhan Pat2 Y-ray Y-ray JRC7447 mutation mutation JRC412 KC1 Bidhan Pat1 KTC1 JRO3690 Multiple cross 6 parents JRO66 JRO204 HybridC Mutation gamma rays Fig 2 Pedigree relationship among released varieties of white jute (C. capsularis) KOM62 ‘JRC 212’ contributed genes for wide adaptability.’JRC 321’, a fine fibre genotype, was one of the parents of two varieties ‘JRC 80’ and ‘UPC 94’. This pedigree relationship could correctly explain the high genetic similarity among the released capsularis varieties (Fig. 2). Fig 1 Relationship among different released varieties of C. olitorius in India. Direction of arrow indicates the female parent used in crossing for developing recombinant lines two varieties, viz ‘S 19’ and ‘JBO 2003-D’ where both were used as parents for transferring pre-mature floweringresistance in adapted lines. The other four varieties originated from diverse genotypes, indicating that the genetic base of released olitorius varieties is indeed narrow. Similarly, the nature of genetic diversity among 13 capsularis varieties released till date also represented narrow genetic base considering the pedigree relationship among those varieties. ‘D 154’ was one of the contributing parents involved either directly or indirectly in developing five varieties, whereas ‘JRC 212’ served as a parent for four varieties inclusive of two varieties where both ‘D 154’ and 2.00 1.50 1.00 Dissimilarity 0.50 Qualitative and quantitative traits in jute: an insight through DUS testing All the olitorius varieties (12 notified and six common knowledge) and seven capsularis varieties (‘D 154’, ‘UPC 94’, ‘JRC 4444’, ‘KTC 1’, ‘Bidhan Pat 3’, ‘JRC 7447’ and ‘JRC 698’) were distinguished for DUS testing (Table 3). Kumar et al. (2006) could distinguish only few varieties using 16 morphological traits based on Draft National Test Guidelines of Jute (Kumar and Mahapatra 2004). They advocated exploring a combination of morphological characters with biochemical and molecular markers for BidhanR JRO620 S19 JRO878 KOM62 JRO36E JRO2345 Tanganyka1 Sudangreen ChinsuraG D154 JRO7835 JRO8432 JRO128 JRO524 TJ40 JRO66 JRO3690 JRO632 JRC698 Padma UPC94 JRC321 KC1 JRC80 KTC1 Bpat3 JRC4444 Bpat1 JRC7447 Bpat2 JRC212 2.00 0.00 1.50 1.00 Dissimilarity 0.50 0.00 Fig 3 Dendograms showing the relationship among released/ notified and common knowledge varieties of both (a) C. olitorius and (b) C. capsularis based on 17 morphological (qualitative and quantitative) characteristics as revealed by cluster analysis. Clustering was based on Euclidian distance with Un-weighted Pair Group Method of Arithmetic averages (UPGMA) 8 August 2009] DIVERSITY IN JUTE AND ITS UTILIZATION 581 Table 3 Morphological characteristics used in DUS testing in jute Morphism Morphological traits C. capsularis C. olitorius Monomorphic (i) (ii) (iii) (iv) (v) (vi) (vii) Premature flowering resistance leaf lamina colour stipule colour leaf shape basal stem root primordial pod dehiscence seed colour (i) (ii) (iii) (iv) (v) (vi) (vii) (viii) Premature flowering resistance leaf lamina colour leaf shape time of 50% flowering pigmentation of calyx basal stem root primordial seed size pod dehiscence Dimorphic (i) (ii) (iii) (iv) (v) (vi) (vii) Leaf petiole colour stem colour plant height pigmentation of calyx time of 50% flowering pod pigmentation seed size (i) (ii) (iii) (iv) (v) (vi) (vii) (viii) (ix) Leaf petiole colour leaf vein colour stem colour plant height stipule colour pod pigmentation seed colour fibre fineness fibre strength discriminating capsularis varieties. Attempts have recently been made for precise characterization of jute varieties of both species for obtaining protection under Protection of Plant Varieties and Farmers Right Act in India. Kumar et al. (2008) characterized 25 released or notified varieties and seven common knowledge varieties of both species using 17 morphological characters for DUS testing. Out of 17 morphological traits, eight were monomorphic, seven dimorphic and two polymorphic in C. capsularis. In contrast C. olitorius was represented by eight dimorphic and nine polymorphic traits. The revised DUS test guidelines of jute (PPV and FRA 2008) were formulated keeping cvs ‘Bidhan Rupali’ (C. olitorius) and ‘D 154’ (C. capsularis) as dummy candidate varieties. Although, qualitative morphological characters in jute are mostly monomorphic with few belonging to dimorphic and polymorphic types, certain distinguishing features are being utilized in seed certification programme of released/ notified jute varieties (Kumar et al. 2005). But classifying quantitative morphological traits into categorical morphs could be misleading. Therefore, when quantitative morphological traits were treated as interval variables along with qualitative variables (consisting of nominal and ordinal categories), the clustering pattern in both species is emerged as shown in Figs 3a, b. and for the identification of closely related genotypes (Me´tais et al. 2002). Use of molecular techniques could be of help in devising strategies to improve the jute crop. However, there appears to be very little molecular information about jute or its related species as available at the gene bank, and very few efforts have been made in the past to develop molecular markers to study its genetic variability (Hossain et al. 2002, 2003, Roy et al. 2006, Basu et al. 2004, Haque et al. 2007, Mir et al. 2008). Different authors had determined the genetic variability of various jute varieties and accessions collected from diverse locations, using RAPD (Qi et al. 2003, Haque et al. 2007), ISSR (Qi et al. 2003b), RAPD and AFLP (Hossain et al. 2002, 2003), AFLP and SSR (Basu et al. 2004), STMS, ISSR and RAPD (Roy et al. 2006) and SSR (Mir et al. 2008) markers (Table 4). Hossain et al. (2002) investigated nine different varieties and 12 accessions of jute cultivars belonging to both species using RAPDs. Out of 29 primers used, seven and six primers gave polymorphism within the varieties and accessions, respectively. Distinct clustering of the cultivated jute species was obtained and species-specific RAPDs had been generated to distinguish between C. olitorius and C. capsularis. Existing genetic classification was in agreement with molecular marker data. Qi et al. (2003a) generated fingerprints of 10 species including 27 accessions of the genus Corchorus using 25 RAPD primers. The presence of abundant genetic diversities was reported among 15 wild species and 12 cultivated Corchorus spp, with genetic similarity co-efficients ranging from 0.49 to 0.98, and clustering corroborated the Application of molecular markers Molecular markers provide a direct measure of genetic diversity and go beyond indirect diversity measures based on morphological traits or geographical origin. They have been successfully employed to characterize genetic diversity 9 582 KAR ET AL. [Indian Journal of Agricultural Sciences 79 (8) 36.6 35.4 61.7 41.5 70.9 31.6 48.8 34.6 100 50.4 57.7 88.3 98.9 43.9 100 69.3 66.9 84.4 55.6 71.3 36.1 96.9 109 89.1 0.14 0.21 0.28 0.36 0.43 0.50 0.57 0.64 0.71 0.79 0.86 0.93 C. capsularis elite Indian varieties CIJ010 CIJ011 CIJ056 CIJ012 CIJ013 CIJ032 CIJ033 CIJ143 CIJ057 CIJ058 JRC698 Hybrid C JRC321 UPC94 JRC212 JRC4444 BidhanPat2 JRC7447 BidhanPat1 BidhanPat3 C. trilocularis 41 C. capsularis exotic germplasm 77.4 90 C. olitorius elite Indian varieties 48.8 C. olitorius exotic germplasm 77.4 OIJ005 OIJ151 OIJ154 OIJ157 OIJ158 OIJ166 OIJ167 OIJ168 OIJ161 OIJ165 99.9 JRO524 JRO524E JRO3690 JRO7835 KOM62 JRO66 JRO878 JRO8432 JRO632 TJ40 C. aestuans 100 Jaccard’s similarity coefficient Fig 4 Dendrogram showing relationship and diversity of elite Indian varieties and exotic germplasm of both C. capsularis and C. olitorius along with wild species C. trilocularis and C. aestuans using STMS markers (adapted from Roy et al. 2006) that the two species are allopatric sharing certain common alleles. Such distinction provides support to the earlier hypothesis that the two species originated from two different geographical locations: C. capsularis from the Indo-Burma region including south China and C. olitorius from Africa (Kundu 1951). DNA fingerprinting of 20 exotic germplasm lines and 20 commercial varieties of the two cultivated species (C. olitorius and C. capsularis) and two wild relatives of jute (C. aestuans and C. trilocularis) have been done using RAPD, ISSR and SSR techniques (Roy et al. 2006). Six STMS markers developed from the genomic sequence of C. olitorius was not fully transferable to C. capsularis, and revealed a very low level of intra-specific polymorphism. The four ISSR and 22 RAPD primers employed in the study revealed 98.44% and 100% inter-specific polymorphism, respectively, while intra-specific polymorphism was significantly low. Released varieties of C. capsularis had extremely narrow genetic bases requiring immediate diversification programme. Higher levels of genetic diversity morphological classification; close wild species of cultivated varieties were genetically different from wild species. C. capsularis had a close relationship with C. olitorius, and further relationship with C. uriticifolius, C. uritifolius and C. aestuans could distinguish them as primary wild species of jute. The round-fruit cultivars were found to be genetically more similar than long-fruit cultivated species. Qi et al. (2003b) applied ISSR technique on the same set of 27 accessions to reveal higher intra-specific genetic resemblance and abundant inter-specific genetic diversity. Based on morphology and DNA classification, it was found that C. urticifolius could be one of the original wild species and C.trilocularis was a variation of C.trilocularis. Basu et al. (2004) evaluated 49 genotypes collected from Bangladesh, India, Thailand, Nepal, China, Pakistan, Sudan, Tanzania, Kenya and America, for their genetic diversity using SSR and AFLP markers. On the basis of cluster analysis and pair-wise genetic distances, the two jute species were widely separated, but differences in geographical locations did not affect genetic diversity. The results suggested 10 August 2009] DIVERSITY IN JUTE AND ITS UTILIZATION 41 42 99 I 1b OIJ-103 OIJ-102 OIJ-160 43 583 Cmu 010 (soft stem) CIJ-088 CIJ-067 CIJ-013 CIJ-064 CIJ-010 CIJ-057 70 BZ-1-3 (CIJ-001) SOLIMOS (CIJ-007) LISA (CIJ-005) CIJ-090 CIJ-004 CIJ-060 CIJ-016 CIJ-005 (ROXA) CIJ-056 CIJ-007 CIJ-015 CIJ-009 CIJ-052 CIJ-004 (LISA) CIJ-012 CIJ-074 CIJ-085CIJ-008 CIJ-017 CIJ-077 CIJ-065 CIJ-006 (SOLIMOS) CIJ-019 CIJ-055 CIJ-081 (PARC/2443) CIJ-070 CIJ-003 (BRANCA) JRC-321 JRC-212 OIJ-106 OIJ-176 OIJ-171 OIJ-104 JRO-524 0.4 OIJ-162 OIJ-210 OIJ-208 OIJ-175 OIJ-135 OIJ-170 OIJ-159 OIJ-133 OIJ-138 OIJ-134 OIJ-204 46 OIJ-107 OIJ-153 OIJ-161 49 OIJ-140 OIJ-137 OIJ-158 OIJ-155 OIJ-105 OIJ-173 OIJ-172 49 OIJ-132 OIJ-110 OIJ-137 OIJ-167 OIJ-109 OIJ-108 53 S-19 72 NPL/YPY-026C SALYOUT TANGANYKA-1 OIJ-156 57 OIJ-209 OIJ-174 77 OIJ-207 OIJ-206 46 IIa II 54 11b III 0 70 IIIa IIIb 0.1 Fig 5 Clustering pattern of 81 jute genotypes based on genetic distances estimated from SSR polymorphism. IJO, International Jute Organization; OIJ, C. olitorius IJO collection; OIN, C. olitorius indigenous collection (adapted from Mir R R et al. 2008a) observed with germplasm of both the cultivated species advocated their use in broadening the genetic base in jute. Polyphyletic origin of the two cultivated species was apparent from the co-existence of C. olitorius with the wild species C. aestuans and of C. capsularis with C. trilocularis with distinct clusters. Haque et al. (2007) determined genetic diversity and species relationship among 18 jute genotypes of both C. capsularis (6 varieties and 5 land races) and C. olitorius (4 varieties and 3 land races) using 40 RAPD primers. However, only 12 primers were able to generate intra-and inter-specific polymorphism among the jute genotypes. This study revealed that the two jute species are distantly related, with a high level of similarity within the species. Genomic SSRs were first developed by Mir R R et al. (2008a) in C. olitorius cv O4. Out of 67 genomic SSRs, 60 were found to be simple, and the remaining 7 were compound. In both species, using a subset of 45 genomic SSRs derived from C. olitorius, genetic diversity and DNA polymorphism analyses showed that SSRs of C. olitorius have high transferability (70.43%) to C. capsularis; and this can further be improved by the use of EST-SSR. EST-SSRs represent the transcribed region of genome compared to lowcopy sequences. Higher level of genetic as well as allelic diversities were found in olitorius genotypes than in capsularis genotypes using cpSSRs and genomic SSRs, ISSR and RAPD markers, which could be attributed to higher level of natural out-crossing in C. olitorius (8–12%) than in C. capsularis (3–4%). Mir et al. (2008a) advocated that divergent genotypes of both species should be used in intraspecific crosses for developing improved cultivars of white and tossa jute. It was also shown that many SSRs examined were redundant because 15 polymorphic SSRs resulted in the same level of polymorphism as realized with 41 11 584 KAR ET AL. polymorphic SSRs. Mir et al. (2008b) carried out DNA fingerprinting of 16 selected jute accessions, 8 each from C. olitorius and C. capsularis, using SSR and RAPD markers to develop mapping populations on fibre fineness. High level of polymorphism was detected between fine and coarse fibre-yielding jute accessions differentiating two main clusters between C. olitorius and C. capsularis and each main cluster into two clusters containing fine and coarse fibre jute accessions. RAPD and SSR marker data were highly correlated (r = 0.97). Grouping of jute accessions based on morphological and molecular data was also highly correlated. [Indian Journal of Agricultural Sciences 79 (8) successful inter-specific hybrid between wild species C. pseudo-olitorius (‘WCIJ 34’) × C. capsularis (‘Tripura’) was obtained to introgress resistance to diseases like stem rot, black band, soft rot and anthracnose (Palve et al. 2005). This inter-specific hybrid exhibited partial to full male sterility due to non-homology between parents. The resultant semifertile hybrids, which are now in F4 generation, are expected to have considerable values with respect to breeding for disease resistance and fibre quality. Utilization of wild species of Corchorus Most of the wild species of Corchorus are poor yielder, but potent sources of biotic and abiotic stress tolerance coupled with finest quality of fibre (Palve et al. 2004; Mahapatra and Saha 2008). Corchorus depressus, C. asplenifolius, C. cinerascens, C. erinoceus and C. erodiodes exhibited a higher degree of drought tolerance and C. trilocularis and C. tridens (WCIJ 046) showed a significant level of tolerance to water stagnation. Corchorus hirtus, C. sulcatus and C. siliquosus have the ability to colonize on extremely shallow soil. Corchorus pseudo-olitorius showed immune reaction to fungal diseases like stem rot, root rot, black band, soft rot and anthracnose. Similarly, C. urticifolius and C. pseudo-capsularis exhibited resistant reactions to all diseases except soft rot and anthracnose, respectively. Besides these, C. junodii, C. pinnatipertitus, C. saxatilis and C. gillettii are also potent sources of biotic stress tolerance. C. urticifolius, C. pseudo-capsularis and C. pseudo-olitorius were found to be moderately resistant to root knot nematode, Meloidogyne incognita (Cofoid & Chitwood) Chitwood. Nematode galls were observed only in the lateral roots, but the main root was free of galls. The number of egg masses and galls/plant were 7 and 12 in C. pseudo-capsularis as against 310 and 403 in C. capsularis cv ‘JRC 212’ (S K Laha, personal communication). C. pseudo-capsularis produces finest fibre (0.20 tex), followed by C. urticifolius (0.30 tex), C. aestuans (0.51 tex), C. trilocularis (0.77 tex), and C. pseudo-olitorius (0.95 tex). Corchorus angolensis and C. merxmuelleri were found to be high fibre yielder with superior quality. So, introgression of desirable genes from wild germplasm into cultivated Corchorus species is utmost important. Being divergent from the cultivated species, wild species should be used as source of variability and parents for initiating inter-specific hybridization programme through traditional and advanced tools of biotechnological approaches to increase the productivity and quality of jute fibre. Corchorus species are characterized by a high degree of inter-specific variability, but a low level of intra-specific variability. Due to modern method of cultivation, the genetic base of the cultivated varieties has become very narrow, where C. olitorius cultivars are dominating in cultivation as compared to C. capsularis varieties. Most of the adapted areas of the jute growing region are represented by only a few olitorius varieties, particularly ‘JRO 524’. Narrow genetic STEPS FOR WIDENING GENETIC BASE Intra-specific hybridization Varietal development of cultivated Corchorus is confined to intra-specific hybridization. There is limited scope for further improvement of cultivated varieties of jute in the absence of requisite variability and genetic diversity due to narrow genetic base. Therefore, additional genetic variability is required to provide new phenotypes in breeding populations. Jute breeders should utilize vast array of germplasm, specially from African continent in varietal breeding programmes of both species. Inter-specific hybridization Inter-specific crosses may help in combining desirable attributes from the two jute species, as it has been successfully attempted and utilized in the past (Islam and Rasid 1960, Swaminathan et al. 1961, Islam 1964, Sinha et al. 2004). However, strong sexual incompatibility barrier exists between the two cultivated jute species, which do not crossfertilize (Patel and Datta 1960, Swaminathan et al. 1961). But Choudhuri and Mitra (1962) have succeeded in producing inter-specific jute hybrids. However, those putative hybrids exhibited dominance of the female parent in F1 and F2 generations, making it impossible to release varieties. Rout and Naik (1983) followed the hybrid progenies up to F3 generations, but there was no report on exploitation of those hybrids in later generations. This is probably because of the fact that the later generation of the entire population resembled the female parents. Bhaduri and Bairagi (1968) succeeded in obtaining successful inter-specific hybrids between the two cultivated species. However, all these attempts did not lead to the development of elite cultivars through recombination breeding. Recently, Sinha et al. (2004) obtained successful interspecific hybrids using mutant C. capsularis female parent (‘CMU 001’) with C. olitorius cv ‘JRO 524’. The resulting hybrid exhibited better fibre fineness and cellulose content, but low-lignin content and reduced fibre strength. They advocated selection of progenies with quality fibre in advance generations for utilizing and blending in textile industries. A 12 August 2009] DIVERSITY IN JUTE AND ITS UTILIZATION 585 Table 4 Application of molecular markers in genetic diversity analysis in Corchorus species Plant material studied Molecular marker used* Country Reference 27 accessions of 10 species (C. aestuans, C. tridens, C. fascicularis, C. pseudo-olitorius, C. pseudo-capsularis, C. trilocularis, Tian Jute [untitled], C. capsularis, C. olitorius and C. uriticifolius) in Genus Corchorus 9 different varieties and 12 accessions of both C. olitorius and C. capsularis 2 varieties and 4 accessions of C. olitorious RAPD and ISSR China Qi et al. 2003a, 2003b RAPD Bangladesh Hossain et al. 2002 RAPD(30) and AFLP(25) SSR & AFLP Bangladesh Hossain et al. 2002 India Basu et al. 2004 RAPD (22) ISSR(4) STMS (6) RAPD(40) India Roy et al. 2006 Bangladesh Haque et al. 2007 SSR(45) SSR and RAPD India India Mir R R et al. 2008a Mir J I et al. 2008b C. capsularis (1 variety and 21 wild accessions), C. olitorious (2 varieties and 25 wild accessions) C. capsularis , C. olitorius varieties and germplasm (10 each), C. aestuans, C. trilocularis C. capsularis (6 varieties and 5 accessions), C. olitorious (4 varieties and 3 accessions) C. capsularis (45) and C. olitorius(36) 16 selected jute accessions, eight each of C. olitorius and C. capsularis *Figures in parentheses denote the number of primer used. base in C. capsularis is a matter of great concern in varietal development. Although a number of molecular markers have been employed, refinement in enrichment and development of more microsatellite markers and their utilization will be essential for jute genomic research. The identification of diverse lines with these molecular markers will assist jute breeders in achieving a greater success for both intra-and inter-specific hybridization. Validation of marker diversity should not be contradicted with morphological, biochemical and physiological diversity. Although both kinds of markers have their advantages and disadvantages, their combined utilization would increase the resolving power of genetic diversity analysis. and improved varieties of jute. Technical Bulletin Series No1/ 2004, p 1–11, CRIJAF, Batrrackpore, Kolkata. Das, B B and Maiti S N. 1998. Jute (Corchorus species) and allied fibres research in India. Indian Journal of Agricultural Research 68(8): 484–93. Gepts P. 1993. The use of molecular and biochemical markers in crop evolution studies. Evolutionary Biology 27: 51–94. Ghosh T. 1983. Plant Production and Protection. Paper No.51. 219pp FAO, Rome. Gilliland T J, Coll R, Calsyn E, de Loose M, van Eijk MJT, RoldanRuiz I. 2000. Estimating genetic conformity between related ryegrass (Lolium) varieties. I. Morphology and biochemical characterization. Molecular Breeding 6: 569–80. Haque S, Begum S, Sarker R H and Haseena Khan. 2007. Determining genetic diversity of some jute varieties and accessions using RAPD markers, Plant Tissue Culture and Biotechnology 17(2): 183–91. Hossain M B, Awal A, Rahman M A, Haque S and Khan H. 2003. Distinction between cold sensitive and cold tolerant jute by DNA polymorphisms. Journal of Biochemistry and Molecular Biology 36: 427–32. Hossain M B, Haque S and Khan H. 2002. DNA fingerprinting of jute germplasm by RAPD. Journal of Biochemistry and Molecular Biology 35: 414–9. Islam A S 1964. A rare hybrid combination through application of hormone and embryo culture. Nature 210: 320. Islam A S and Rashid A.1960 First successful hybridization between the two jute yielding species. Nature 185: 258–9. Islam A S and Rashid A.1961. A new jute hybrid. Journal of Heredity 52: 287–91. Karmakar P G, Hazra S K, Sinha M K and Chaudhury S K. 2008. Breeding for quantitative traits and varietal developments in Jute and allied fibre crops. (in) Jute and Allied Fibre Updates: Production and Technology, pp57–75. Karmakar P G, Hazra S K, Ramasubramanian T, Mandal R K, Sinha M K and Sen H S REFERENCES Basak S L. 1993. Review on the genetics and breeding of jute, International Jute Organization, Dhaka, Bangladesh. Basu A, Ghosh M, Mayer R, Powell W, Basak S L, and Sen S K. 2004. Analysis of genetic diversity in cultivated jute determined by means of SSR markers and AFLP profiling. Crop Science 44: 678–85. Bhaduri P N and Bairagi P. 1963. Inter-specific hybridization in jute(Corchorus capsularis and C. olitorius) Science and Culture 34: 355–7. Brandolini A.1969. European races of maize. Annual Corn Sorghum Research Conference Proceedings 24: 36–48. Bretting P K, Wildrlechner M P.1995. Genetic markers and plant genetic resource management. (in) Plant Breeding Reviews, pp 11–71 Janick J (Ed) Vol 13. Wiley, New York. Chaudhuri S D and Mia A J. 1962. Species crosses in the genus Corchorus (jute plants). Euphytica 11: 61–4. Chaudhury S K, Kumar D and Saha A. 2004. Varietal development 13 586 KAR ET AL. (Eds) CRIJAF, Barrackpore, Kolkata. Kumar D, Begum T, Kotal B and Shil S. 2008. Descriptors of Jute Varieties. Technical Bulletin No. 2/2008, CRIJAF, Barrackpore, Kolkata. Kumar D and Mahapatra A K. 2004. Draft National Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability in Jute, pp 1–14,CRIJAF, Barrackpore. Kumar D, Das A, Mahato P, Lakshman S S and Mandi S. 2005. Identification key for notified varieties and varieties of common knowledge of jute (Corchorus olitorius L. and C. capsularis L.) Indian Journal of Genetics and Plant Breeding 65: 236–8. Kumar D, Mahato P, Lakshman S S and Mandi S. 2006. Morphological characterization of jute varieties (Corchorus olitorious L. and C. capsularis L.) and their application in DUS testing. Indian Journal of Genetics and Plant Breeding 66: 319– 23. Kundu B C. 1951. Origin of jute. Indian Journal of Genetics and Plant Breeding 11: 95–9. Kundu B C, Basak K C and Sarkar P B. 1959. Jute in India-A. monograph. 395 pp. Indian Central Jute Committee. Mahapatra A K and Saha A. 2008. Genetics resources of jute and allied fibre crops. (in) Jute and Allied Fibre Updates: Production and Technology, pp18–37, Karmakar P G, Hazra S K, Ramasubramanian T, Mandal R K, Sinha M K and Sen H S (Eds) CRIJAF, Barrackpore, Kolkata. Me´tais I, Hamon B, Jalouzot R, Peltier D. 2002. Structure and level of genetic diversity in various bean types evidenced with microsatellite markers isolated from a genomic enriched library. Theoretical and Applied Genetics 104: 1346–52. Mir R R, Rustgi S, Sharma S, Singh R, Goyal A, Kumar I, Gaur A, Tyagi A K, Khan H, Sinha M K, Balyan H S, and Gupta P K. 2008a. A preliminary genetic analysis of fibre traits and the use of new genomic SSRs for genetic dliversity in jute. Euphytica 161: 413–27. Mir J I, Karmakar P G,Chattopadhyay S, Chaudhury S K, Ghosh S K, Roy A. 2008b. SSR and RAPD profile based grouping of selected jute germplasm with respect to fibre fineness trait. Journal of Plant Biochemistry and Biotechnology 17: 29–35. Palit P, Sasmal B C, Bhattacharryya A C.1996. Germplasm diversity and estimate of genetic advance of four morpho-physiological traits in a world collection of jute. Euphytica 90: 89–110. Palve S M, Sinha M K, Chattopahdyay S. 2004. Genetic variability for fiber strength and fineness in wild relatives of genus Corchorus. (in) Proceedings National Seminar on Diversified Uses of Jute and Allied Fiber Crops, pp 99–103, held during 8th–9th Jan-uary, Barrackpore, Kolkata. Palve S M and Sinha M K. 2005. Morphological studies in interspecific hybrid between Corchorus pseudo-olitorious Islam and Zaid and Corchorus capsularis Linn. Science and Culture 71 (1–2): 35–7 Patel J S and Ghosh R L M. 1940. A review of agricultural investigations on jute in India. Agriculture Research Bulletin No. 1. Patel G I and Datta R M. 1960. Inter-specific hybridization between Corchorus olitorius and C. capsularis and the cytogenetical basis of incompatibility between them. Euphytica 9: 89–110. PPV and FRA. 2008. Specific DUS test guidelines for six notified species of cotton and jute. Plant Variety Journal of India. 2(1&2): 75–86. Qi J, Zhou D, Wu W, Lin L, Wu J, and Fang P.2003a. Application of ISSR technology in genetic diversity detection of jute. Ying Yong Sheng Tai Xue Bao 14: 1473–7. Qi J, Zhou D, Wu W, Lin L, Fang P, and Wu J. 2003b. The application of RAPD technology in genetic diversity detection of Jute. Ying Yong Sheng Tai Xue Bao 30: 926–32. Raut R N and Naik G.1983. Inter-specific hybridization in cultivated jute. Proceedings of FAO Expert Consultation on Jute and Kenaf improvement, pp 1–11. held during 5–9 September, Calcutta. Roy A, Bandyopadhyay A, Mahapatra A K, Ghosh S K, Singh N K, Bansal K E, Koundal K R, and Mohapatra T. 2006. Evaluation of genetic diversity in jute (Corchorus species) using STMS, ISSR and RAPD markers, Plant Breeding 125: 292–7. Saha A and Hazra S K. 2008. Jute – the Heritage of India. (in) Jute and Allied Fibre Updates: Production and Technology, pp1– 17. Karmakar P G, Hazra S K, Ramasubramanian T, Mandal R K, Sinha M K and Sen H S(Eds) CRIJAF, Barrackpore, Kolkata. Sinha M K, Mitra S, Nandy S, Sengupta D , Dutta P, Das F and Chakrabarty S C. 2004. Inter-specific hybrid between two jute (Corchorus) species for textile quality fibre. Indian Journal of Genetics and Plant Breeding 64(4): 310–4. Swaminathan M S, Iyer R D and Sulbha K.1961. Morphology, cytology and breeding behaviour of hybrids between Corchorus olitorius and C. capsularis. Current Science 30: 67–8. 14 View publication stats [Indian Journal of Agricultural Sciences 79 (8)