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Cytogenetical Investigations in Colchicine-Induced Tetraploids of Brassica Fruticulosa: an Important Wild Relative of Cultivated Brassicas

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Cytogenetical Investigations in Colchicine-Induced Tetraploids of Brassica Fruticulosa: an Important Wild Relative of Cultivated Brassicas © 2015 The Japan Mendel Society Cytologia 80(2): 223–230 Cytogenetical Investigations in Colchicine-Induced Tetraploids of Brassica fruticulosa: An Important Wild Relative of Cultivated Brassicas Arun Kumar*, Binay Kumar Singh, Hari Singh Meena, Bhagirath Ram, Vijay Veer Singh and Dhiraj Singh Directorate of Rapeseed-Mustard Research, Bharatpur-321 303, Rajasthan, India Received October 4, 2014; accepted January 12, 2015 Summary Successful induction of autotetraploidy has been achieved in five plants of B. fruticulosa Cyr. subsp. fruticulosa (2n=16, FF), a wild relative of cultivated brassicas. The diploid seedlings of B. fruticulosa were treated with different concentrations of aqueous colchicine using the cotton-swab method for 8–15 h for 2 to 3 d. The highest percentage of success was recorded when the seedlings were treated with 0.2% aqueous colchicine for 12 h within 2 d. The synthesized plants showed remarkable enhancement in several morphological and floral characters making them more robust. Cytologically, among all the associations, bivalent chromosome associations were observed more frequently (7.20 to 8.76 per cell) in various plants. However, quadrivalent frequency ranging from 2.28 to 3.14 and univalent frequency ranging from 3.44 to 4.09 per cell were characteristic of the colchicine-induced tetraploids. The number of chiasmata ranged from 15 to 36 with an average number of 21.28 and 23.60 and their terminalisation coefficient ranged between 0.73 and 0.86. The anaphase I and II disjunction of bivalents/chromosomes was leading more or less regularly and equally to the formation of seeds from the synthesized plants. Significant enhancement in morphological traits as revealed in colchicine-induced plants and normal meiotic behaviour leading to a good seed set may ultimately result in providing the plant breeder with more variability. The synthesized autotetraploids will have great opportunities to be utilized in hybridization with Brassica juncea and related species for introgression of desirable traits, especially tolerance for mustard aphid. Key words Chiasmata, Cytology, Induced polyploidy, Meiotic behaviour, Quadrivalent. Brassica juncea L. Czern & Coss., commonly known as Indian mustard, is the predominantly cultivated Brassica species in India with yield potential of 15–30 q/ha and 38–42% oil content. This crop often encounters severe biotic and abiotic stresses emerging as a result of unprecedented change in environmental conditions at local, regional and global levels (Chopra et al. 1996). Amongst the biotic stresses, mustard aphid, Lipaphis erysimi (L.) Kaltenbach is a perpetual annual threat leading to productivity losses up to 70%, depending upon the severity of outbreak (Bandopadhyay et al. 2013). Present methods of aphid control in mustard are primarily based on synthetic chemical insecticides. These chemicals, besides aggravating environmental pollution, can also be toxic to friendly insects. In this sense, a resistant cultivar is a more sustainable and environment-friendly option. The development of an insect-resistant cultivar requires a heritable and transferable resistance (Stoner and Shelton 1988). However, cultivated Brassica germplasm has failed so far to provide any source of resistance against mustard aphid. Fortunately, Brassica coenospecies has been bestowed with nearly 100 species of wild and weedy relatives, possessing extensive genetic variability for many defence traits. These species can be effectively utilized to * Corresponding author, e-mail: [email protected] DOI: 10.1508/cytologia.80.223 224 A. Kumar et al. Cytologia 80(2) introduce economically important traits to cultivated species as well as development of potential wide hybrids (Prakash 2001, Singh et al. 2011, Singh et al. 2012). Brassica fruticulosa Cyr., a wild relative of crop brassicas, has been reported to possess resistance against cabbage aphid, Brevicoryne brassicae (Cole 1994a, Ellis and Farrell 1995, Ellis et al. 2000) and a higher concentration of lectins was suggested to be the underlying mechanism of resistance in this species (Cole 1994b). Earlier we have reported the synthesis of an inter-specific hybrid between B. rapa (2n=20, AA) and B. fruticulosa (2n=16, FF) through sexual hybridization (Kumar et al. 2013). However, inter-specific hybridization between homoploid B. fruticulosa and heteroploid B. juncea (2n=36, AABB) could not be achieved. It is evident that when heteroploid species with different ploidy levels are crossed, hybridization between them is relatively more difficult than when species with the same ploidy level are mated. Keeping these in view, present investigations are undertaken to induce polyploidy, particularly colchi-autotetraploidy in B. fruticulosa using aqueous colchicine, and their subsequent utilization in inter-specific hybridization with B. juncea. To the best of our knowledge, this is the first report of synthesis and utilization of autotetraploid B. fruticulosa in Brassica crop improvement programmes. Materials and methods Seed samples of B. fruticulosa were obtained from the germplasm section of the Directorate. The seeds were directly sown in earthen pots filled with peat, perlite and sand (1 : 1 : 1, v/v/v). Cotton swab method was employed for colchicinization. Sterilized cotton swabs immersed either in 0.1 or 0.2% aqueous colchicine were placed on emerging apical tip between two cotyledonary leaves. To avoid an increase in the concentration, colchicine was added drop by drop at regular intervals on the cotton swabs with the help of sterilized syringe. Such treatment was done for 2 to 3 d for 3 to 4 h per day. The data on various morphological attributes such as plant height, primary and secondary branches, main shoot length, leaf length, siliqua length, length and breadth of petals, etc. of both diploid and colchicine-induced plants were recorded by physical measurements with centimetre scale or micrometre as the case may be. For meiotic studies flower buds of an appropriate size were collected from selected mature plant and fixed on the spot in freshly prepared carnoy’s fluid (ethanol : chloroform : acetic acid̶6 : 3 : 1), supplemented with a drop of ferric chloride solution, for a minimum of 24 h at room temperature and subsequently stored in 70% alcohol at 10°C. Anthers were squashed in 1% acetocarmine solution with ferric chloride solution as mordant. On average 25 Pollen Mother Cells (PMC) were analyzed at diakinesis/metaphase I to estimate the range of chromosome associations and recombinational frequencies by chiasma analysis. At anaphases I/II on average 15–20 cells were analysed to study the distributional pattern of chromosomes. Results Efficiency of colchicine treatment B. fruticulosa seedlings at cotyledonary leaves stage were treated with aqueous colchicine (0.1% and 0.2% v/v) for 2 to 3 d for 3 to 5 h per day. The highest induction percentage (15.79%) of putative polyploidy was recovered with 0.2% aqueous colchicine (counted as the number of colchicine-induced tetraploids recovered from total number of seedlings treated with aqueous colchicine) (Table 1). A total of five colchicine-induced putative polyploids were synthesized. These plants are numbered as P1, P2, P3, P4 and P5, while diploid plants are numbered as P. 2015 Cytogenetical Investigations in Colchicine-Induced Tetraploids of Brassica fruticulosa 225 Table 1. Efficiency of colchicine treatment in inducing polyploidy by cotton swab method in B. fruticulosa. Colchicine Duration of No. of days of No. of seedlings No. of seedlings No. of colchicine- Percentage of concentration (%) treatment (h) treatment treated survived induced tetraploids tetraploids 0.1 12 3 30 22 1 4.54 15 3 20 8 ̶ 0.0 0.2 8 2 25 12 1 8.33 10 2 25 19 3 15.79 Table 2. Morphological features of diploid and colchicine-induced tetraploids of B. fruticulosa. Sample P P1 P2 P3 P4 P5 Character No. 2x 4x 4x 4x 4x 4x 1. Plant height (cm) 58.5 48.5 60 49.2 51.6 36.5 2. Primary branches per plant 5 5 5 4 3 4 3. Secondary branches per plant 6 10 8 12 5 8 4. Main raceme length (cm) 32.7 25 35 25.5 29.2 28.4 5. Length of petiole (cm) 2.3 2.5 2.6 2.4 2.5 2.3 6. Length of odd leaflet (cm) 8.5 12.3 13.6 11.7 12.0 9.4 7. Width of odd leaflet (cm) 6.2 8.5 7.8 8.1 7.6 6.8 8. Corolla length (cm) 1.0 1.3 1.5 1.4 1.3 1.2 9. Corolla width (cm) 0.5 0.6 0.8 0.7 0.6 0.5 10. Siliqua length (cm) 3.0 3.4 3.5 3.2 3.4 3.8 11. Seeds per siliqua (mean) 16 4 10 7 6 7 Figs. 1–3. Comparison of morphological attributes in diploid and colchicine-induced tetraploids of B. fruticulosa, Fig. 1. Leaf, 2. Flower, 3. Siliquae. Morphology All the five colchicine-induced plants were robust from the initial stages of development and it was maintained till maturity, although different responses to colchicine were evident in all the five plants (Figs. 1–3, Table 2). All the colchicine-induced tetraploid plants showed reduction in plant height except one (P2) when compared with diploids. Secondary branches per plant, length of petiole, length and width of leaflet, corolla length and width and siliqua length showed considerable increase in comparison with the corresponding diploids. As for primary branches per plant in tetraploids, three plants (P3, P4 and P5) showed reduction, whereas in two plants (P1 and P2) numbers of primary branches per plant were found to be similar to diploids. The number of seeds per siliqua decreased in all the synthesized colchicine-induced polyploid plants. Among the five colchicine-induced tetraploid plants, P2 showed considerable increase in all the morphological attributes accept seeds per siliqua which were comparability less and primary branches per plant which was equal to the diploid plants (Table 2).
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