Cytological Studies in Some Members of Tribe Senecioneae (Asteraceae) from North and Central India
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© 2010 The Japan Mendel Society Cytologia 75(4): 369–378, 2010 Cytological Studies in Some Members of Tribe Senecioneae (Asteraceae) from North and Central India Raghbir C. Gupta, Santosh Bala*, Henna Goyal, Reyaz A. Malik and Santosh Kumari Department of Botany, Punjabi University, Patiala-147002, Punjab, India Received July 24, 2010; accepted August 24, 2010 Summary Cytological studies have been made of 12 species of tribe Senecioneae (Asteraceae) from north and central India. Of these, 3 species, namely S. grahami Hook. f. (nϭ10), S. kunthianus Wall ex DC. (nϭ20) and S. sexatilis Sensu Hook. f. (nϭ10), are counted for the first time. Additional and/or variable cytotypes are recorded for 2 species, namely S. nudicaulis Buch.-Ham D. Don. (nϭ5, 10) and S. rufinervis DC. (nϭ10). Existence of B-chromosomes has been recorded in S. krascheninnikovii Schischk. (nϭ10ϩ0Ϫ2B). Two species, S. vulgaris L. (nϭ20) and S. kraschenin- nikovii Schischk. (nϭ10), are explored for the first time from India. Besides these, cytomorphologi- cal variabilities have been observed in S. rufinervis (2nϭ20, 40) and S. nudicaulis Buch-Ham ex D. Don (2nϭ10, 20 and 40). Great variation in morphological characteristic have been noticed in different accessions of S. laetus Edgew, however all the accessions are found to exist at 2nϭ40. Meiotic configurations in the form of multivalents and/or secondary association of bivalents are found in Gynura nepalensis DC., Senecio laetus Edgew, S. kunthianus Wall ex DC., S. kraschenin- nikovii Schischk. and S. nudicaulis Buch-Ham ex D. Don. Key words Senecioneae, Base number, B-chromosome, Cytomorphovariants, Multivalents, Secondary associations. Senecioneae is one of the largest tribes in the family Asteraceae, comprised of approximately 3,100 species and 155 genera, and has a more or less cosmopolitan distribution and show remarkable cytomorphological as well as ecological diversity. Senecio is the largest genus with approximately 1,000 species, with 43 species reported in India (Hajra et al. 1995). Phylogenetically, the tribe Senecioneae is proposed as the most primitive tribe of the family Asteraceae (Small 1919). The proposed base number for the tribe is xϭ5. From an economic point of view, the members of this tribe have a significant importance in the field of medicine and for the perfumery. Materials and methods For meiotic studies, usual acetocarmine smears of appropriate sized young capitula were made after fixing them in Carnoy’s fixative. Pollen fertility was estimated by mounting mature pollen grains in glycerol-acetocarmine. Voucher specimens are available in Herbarium, Department of Botany, Punjabi University, Patiala (PUN) (Table 1). Results and discussion Information on taxa, specific locality, accession number (PUN), chromosome number and meiotic observations of the presently investigated 12 species of the tribe Senecioneae is * Corresponding author, e-mail: [email protected] 370 R. C. Gupta et al. Cytologia 75(4) Table 1. Information on taxa, voucher data, accession number, chromosome number and meiotic configurations of presently investigated 12 species of the tribe Senecioneae. Accession Chromo- S. Voucher data Meiotic Taxa some no. (Locality with altitude) configurations number number (n) 1. Gynura nepalensis DC. Palampur, 1219 m 52812 20 20 II (70%) 1 IVϩ18 II (20%) 2 IVϩ16 II (10%) 2. Emilia sonchifolia (L.) Kangra, 530 m 52813 5 5 II (38.46%) DC. Sangrur, 250 m 24964 5 2 IIϩ6 I (61.53%) Dehra Dun, 1200 m 24502 5 Pachmarhi, 1000 m 24955 5 3. S. laetus Edgew Lohardi; Chhota Banghal, 2200 m 52604 20 20 II (83.33%) Manali, 2050 m 52807 20 1IVϩ18 II (8.33%) Ahrarbal; Kashmir, 2400 m 52814 20 Secondary associations Swar; ChhotaBanghal, 2500 m 52605 20 of 2–8 bivalents (8.33%) Zispa; Lahaul -Spiti, 3142 m 52806 20 Koksar; Lahaul-Spiti, 3160 m 52805 20 Marhi; Kullu, 3320 m 52815 20 Keylong; Lahaul-Spiti, 3350 m 52804 20 Zingzingbar; Lahaul-Spiti, 52803 20 4328 m 4. S. elegans Linn. Patiala, 250 m 24416 10 10II (100%) 5. S. graciliflorus DC. Lohardi; Chhota Banghal, 52606 20 Secondary associations 2200 m of 2–6 bivalents (100%) 6. * S. grahami Hook f. Pachmarhi, 1000 m 24386 10* 10II (100%) 7. * S. kunthianus Chatru; Spiti, 4270 m 52802 20* 20 II (63.63%) Wall ex DC. 2IVϩ16 II (27.27%) 4IVϩ12 II (9.09%) 8. * S. krascheninnikovii Aharbal; Kashmir, 2400 m 52810 10* 10 II (44.44%) Schischk. 10 IIϩ2B (22.22%) 9 IIϩ2 I (11.11%) 1IVϩ8 II (11.11%) 1 IVϩ7 IIϩ2 I (11.11%) 9. * S. nudicaulis Buch.- Palampur, 1219m 52607 20 1IVϩ18 II (10%) Ham. ex D.Don 2IVϩ16 II (7.14%) 3IV + 14 II (5.71%) Secondary associations of bivalents (31.4%) Killar; Pangi valley, 2600 m 52808 10* 10 II (100%) Badagaon; Chhota Banghal, 52809 5* 5 II (76.05%) 3300m 1IVϩ3 II (5.63%) 3 IIϩ4I (18.30%) 10. * S. rufinervis Dalhousie, 2200 m 24601 10* 20 II (100%) DC. Shimla, 2400 m 24602 20 Mussoorie, 2000 m 49308 20 Lohardi; Chhota Banghal, 2500 m 52608 20 11. * S. sexatilis Sensu Hook. f. Pachmarhi, 1000 m 25569 10* 10 II (100%) 12. * S. vulgaris L. Pulwama; Kashmir, 2000 m 52811 20* 20 II (100%) * New records 2010 Cytological Studies in Some Senecioneae of North and Central India 371 summarised in Table 1. Features of cytological and/or morphological interest are discussed under each genus. Three species, namely S. grahami Hook. f. (nϭ10), S. kunthianus Wall ex DC. (nϭ20) and S. sexatilis Sensu Hook. f., are worked out cytologically for the first time on a worldwide basis. New cytotypes are recorded for 2 species, namely S. nudicaulis Buch.-Ham D. Don. (nϭ5 and 10) and S. rufinervis DC. (nϭ10). Two species, namely S. vulgaris L. (nϭ20) and S. krascheninnikovii Schischk. (nϭ10), are newly reported in India. The degree of pollen stainability was high in all the studied species, ranging from 82 to 100%. Gynura nepalensis DC. (ϭG. foetens Wallich ex DC.) The species is found distributed throughout India up to 2300 m. Meiotic analysis shows the presence of nϭ20 and is in conformity with the previous reports (Mehra et al. 1965, Shetty 1967). 1–2 quadrivalents are present in some PMCs, besides bivalents (Fig. 1, Table 1). Further meiotic course was normal with 100% pollen fertility. Emilia sonchifolia (L.) DC. (ϭCacalia sonchifolia DC.) The species is found distributed throughout India up to 2000 m. Present studies confirm the previous reports of nϭ5 (Fig. 2) both from south India (Mathew and Mathew 1988) and north India (Mehra and Remanandan 1975), and the rest of the world (Afzelius 1924). Besides this, an aneuploid cytotype with 2nϭ16 is also reported (Gupta et al. 1972). Some of the PMCs show the occurrence of early separation of (4–6) bivalents at metaphase-I (Fig. 3, Table 1). Pollen fertility was cent per cent and pollen size ranges between 25.9–29.2 mm. S. laetus Edgew (ϭS. chrysanthemoides DC.ϭS. spectabilis Wallich ex DC.ϭS. sisymriiformis DC.) It is a common species of Temperate Himalayas and distributed up to 2000–4250 m. Present studies were made on 9 populations of the species from different areas of central and north India (Table 1). All showed nϭ20 (Fig. 4) and confirms the previous report (Mehra et al. 1965). Meiosis is normal but secondary associations of 2–8 bivalents are observed only in the Lohardi population in 8.33% of cells (Fig. 5). Some quadrivalents (1–2) are also found, besides bivalents (Table 1). Pollen fertility varies between 89–97%. Morphologically, all the accessions show lot of variations due to different altitudinal distribution range (2050–4328) m. Comparison of some characters of different accessions of Senecio laetus is given in Table 2. At high altitude (4328 m), the height of the plant gradually decreases to 9.5 cm as compared to plants at 2050–3350 m altitude that attain maximum of 30–50 cm height. Number and size of capitula/plant show lot of variation in different altitudinal accessions (Fig. a–i). Leaves also show lot of diversity in the shape, color and size in the different accessions (Fig. a1–i1; Table 2). S. elegans Linn. It is a cultivated species with nϭ10 (Fig. 6) and confirms the previous report from India (Mehra et al. 1965) and elsewhere (Nordenstam 1967). Pollen fertility was cent per cent and pollen size ranges between 22.7–25.9 mm. S. graciliflorus DC. It is commonly found in open places in Temperate Himalayas between 2000–4100 m. The present meiotic studies showed the presence of nϭ20 at diakinesis with interbivalent connections in some bivalents (Fig. 7, Table 1). This confirms the previous chromosome reports (Mehra et al. 1965, Mehra and Remanandan 1975). Meiotic abnormality in the form of unsynchronized (1–3) bivalents at metaphase I was observed in 41.93% cells. Further segregation of chromosomes was found to be normal but some cells show late disjunction of 2–3 bivalents and chromatin bridges at 372 R. C. Gupta et al. Cytologia 75(4) Figs. 1–12. Meiotic analysis. 1) Gynura nepalensis. Metaphase I, 1 IVϩ18 II. (2–3) Emilia sonchifolia. 2) Metaphase I, 5 II. 3) Metaphase I, 2 IIϩ6 I. (4–5) Senecio chrysanthemoides. 4) Diakinesis, 1 IVϩ19 II. 5) Metaphase I, secondary association of bivalents. 6) S. elegans. Metaphase I, 10 II. 7) S. graciliflorus. Diakinesis, interbivalent connections. 8) S. grahami. Metaphase I, 10 II. 9) S. kunthianus. Diakinesis, 4 IVϩ12 II. (10–11) S. krascheninnikovii. 10) Metaphase I, 10 II. 11) Metaphase I, 10 IIϩ2 B. 12) Metaphase I, 1 IVϩ8 II. Scale barϭ10 mm. anaphase I in 5.12%. S. grahami Hook f. The species is quite common at the Pachmarhi hills, and is counted for the first time with nϭ10 (Fig.