Male Meiosis and Chromosome Number in Asteraceae Family from District Kangra of H.P
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International Journal of Botany and Research (IJBR) ISSN 2277-4815 Vol. 3, Issue 1, Mar 2013, 43-58 © TJPRC Pvt. Ltd. MALE MEIOSIS AND CHROMOSOME NUMBER IN ASTERACEAE FAMILY FROM DISTRICT KANGRA OF H.P. (WESTERN HIMALAYAS) SANTOSH BALA & RAGHBIR CHAND GUPTA Department of Botany, Punjabi University, Patiala, Punjab, India ABSTRACT At present 156 populations covering 44 genera and 74 species under 11 tribes of Asteraceae family have been cytologically worked out from district Kangra of Himachal Pradesh in Western Himalayas. The study reported 14 species with new reports which includes varied chromosome count for Aster lanceolatus (2 n = 48), Cotula australis (2 n = 26), Gerbera gossypina (2 n = 18), Gnaphalium pensylvanicum (2 n = 18), Senecio nudicaulis (2 n = 10); first chromosome count for Carpesium abrotanoides (2 n = 36), Conyza stricta var. stricta (2 n = 18), Erigeron acer (2 n = 18), Gnaphalium coarctatum (2 n = 28), Stevia rebaudiana (2 n = 22), Synedrella vialis (2 n = 24); first B-chromosome count for Anaphalis margaritacea (2 n = 28+2B), Anaphalis royleana (2 n = 28+2B) and Cosmos sulphureus (2n=24+0-3B). Population based study in most of the species reveals cytomorphological diversification. Cyto-variants are reported in two species viz. Galinsoga parviflora (2 n = 2x = 16; 2 n = 4x = 32) and Senecio nudicaulis (2 n = 2x = 10; 2 n = 8x = 40). Cytomorpho- variant is observed for Ageratum conyzoides : blue colored ray floret (2 n = 2x = 20), white colored ray floret (2 n = 4x = 40). Morpho-variants are reported for 10 species which covers flower morphotype for Aster peduncularis , Cosmos bipinnatus, Erigeron annuus, Myriactis wallichii, Vernonia cinerea; leaf morphotype for Conyza japonica, Eclipta alba, Taraxacum officinale ; and habit morphotype for Conyza stricta var. pinnatifida and Sonchus oleraceus. Among presently worked out species 2 n = 18 is the most common chromosome number with x = 9 as the commonest base number. Chromosome number data is presently reviewed for presuming probable base numbers for each genus and frequency of polyploidy. Many species observed with meiotic abnormalities which consecutively reduces the pollen fertility. KEYWORDS: Asteraceae, Chromosome Number, Base Number, Meiosis, Polyploidy, Western Himalayas INTRODUCTION With more than 1,600 genera and 23,000 species, Asteraceae forms the largest family of the flowering plants (Funk et al., 2009). Indian Asteraceae is represented by c. 900 species under 167 genera (Hajra et al. , 1995). Family has a worldwide distribution, being found everywhere, especially more common in tropical and subtropical regions. The Asteraceae family is of great economic importance with a special status in floriculture, medicine, source of oil, insecticide, dye, etc. Due to the abundance of morphological as well as ecological diversity and the existence of many evolutionary trends in different parts, the Asteraceae family offers a suitable material for detail cytological studies. The documented evidences are available for the role of polyploidy, aneuploidy, phylogenetic increase and decrease in chromosome number, structural changes, hybridization, apomixis, etc. in the evolution of the species. Cytological studies particularly on Asteraceae were made by many workers (Stebbins et al., 1953; Raven et al., 1960; Solbrig et al. , 1969; Moore and Frankton, 1962; Ornduff et al. , 1963; Strother, 1976). Heywood et al. , (1977) reviewed various tribes of family. The cytology of Indian Asteraceae carried out by Sobti and Singh (1961); on North as well as South Indian taxa by Mehra et al. , (1965), Mehra and Remanandan (1974, 1975, 1976); on North and Central Indian taxa by Gupta and Gill (1983, 1989) and Gupta et al., (1989); on East and West Indian taxa by Sharma and Sarkar 44 Santosh Bala & Raghbir Chand Gupta (1967-1968), and Subramanyam and Kamble (1967); on South Indian taxa by Mathew and Mathew (1978, 1983, 1988) and Shukur et al,. (1977). A significant contribution on cytology of 134 species of North-Indian Compositae was given by Mehra et al. , (1965). The study covered the plants from the areas of Punjab plains, Mussoorie and Shimla hills in Western Himalayas and Darjeeling hills in Eastern Himalayas. Mehra (1977) discussed the phylogeny and interrelationships among different tribes of family. Gupta and Gill (1983) made cytological studies in 71 species of Compositae from Punjab plains. Gupta and Gill (1984) made cytomorphological studies in intraspecific cytotypes of 7 Indian species of Compositae. Gupta and Gill (1989) gave an account of pollen grain characteristics, pollen size, fertility and exine ornamentation along with chromosome numbers for 132 species of Compositae family from North and Central India. Gupta et al., (1989) made cytological analysis on 40 wild species of West Himalayan Compositae and gave notes on presence of B-chromosomes, intra- and inter-specific polyploidy, meiotic abnormalities, incidence of polyploidy, apomixis, hybridization and polyploidy and habit correlation. Though there are some contributions on cytological aspect of Asteraceae family throughout India and North India, but there are many areas which are left unexplored and not taken into consideration. So the present cytological attempt made on plants of district Kangra of Himachal Pradesh, Western Himalayas. Most of the chromosome number reports have already published in chromosome data 12 in taxon. MATERIALS AND METHODS Plant materials of Asteraceae family were collected on population basis from different localities of district Kangra (H.P.), Western Himalayas (Table 1). Appropriate sized young capitula were collected during day time (10 am- 3 pm) and fixed in Carnoy’s fixative (6: 3: 1 = absolute alcohol: chloroform: glacial acetic acid v/v/v) for 24 hrs and preserved in 70 per cent alcohol at 4˚C. The plant specimens have been deposited in Herbarium (PUN), Punjabi University, Patiala. The species subjected to detailed meiotic analysis including meiotic chromosome number, meiotic course, microsporogenesis, pollen fertility and pollen grain size. Smears of pollen mother cells (PMCs) are prepared in 1 % acetocarmine using standard technique. Maximum number of PMCs as possible was observed for meiotic studies at different stages of meiosis. Pollen fertility is estimated by mounting them in 50 % glycero-acetocarmine and kept overnight. Well filled pollen grains with fully stained nuclei are taken as fertile and shrivelled or unstained nuclei as sterile. Photomicrographs of chromosome counts are made from freshly prepared slides using Leica Qwin and Nikon digital imaging systems. Data was compiled for previous chromosome numbers available for each species from different sources: Chromosome atlases of flowering plants by Darlington and Wylie (1955), Fedorov (1969), Kumar and Subramaniam (1986), Index to Plant Chromosome Numbers (1970) onwards. RESULTS AND DISCUSSIONS Data on locality with altitude, accession number, present chromosome number and ploidy level in the presently worked out 74 species of family Asteraceae from district Kangra (H.P.), Western Himalayas is given in Table 1. Only some interesting results are discussed here. CHROMOSOME NUMBER (2n) In family Asteraceae the chromosome number in presently investigated species varies from 2 n = 10 ( Emilia sonchifolia and Senecio nudicaulis ) to 2 n = 72 ( Bidens spp.). The most common chromosome number is 2 n = 18 (29.72 %) followed by 2 n = 16, 36 and 40 (8.10 % each) and the least common are 2 n = 22, 26, 27, 30, 42 and 51 (1.35 % each). Within family the species of tribe Anthemideae basically bear 2 n = 18 while 2 n = 36 ( Chrysanthemum leucanthemum ) and Male Meiosis and Chromosome Number in Asteraceae Family from District Kangra of H.P. (Western Himalayas) 45 2n = 26 ( Cotula australis ) chromosome numbers are also reported. In tribe Astereae, the species mostly bear 2 n = 18 while species with higher chromosome number i.e. multiple of 9 are also reported. Single species, namely, Aster lanceolatus is reported with 2 n = 48. In tribe Cardueae species with 2 n = 32 chromosome number are more common and one species with 2n = 34 is also found ( Cirsium wallichii ). In tribe Cichorieae, the species with three chromosome numbers 2 n = 16, 18 and 32 are noticed of which 2 n = 16 is the most prevalent. Tribe Eupatorieae show mixed results of chromosome numbers 2 n = 20, 22, 40 and 51 in 3 species including one species ( Ageratum conyzoides ) with two intraspecific cytotypes (2 n = 20 and 40). In tribe Heliantheae, Bidens spp. are with 2 n = 72, Cosmos spp. are with 2 n = 24 while rest of the species show 2 n = 16, 22, 24, 30, 34 and 36 chromosome numbers. Species of tribe Inuleae show chromosomal diversity as 2 n = 28 and 42 in Anaphalis spp., 2 n = 18 in Blumea and Vicoa spp., 2 n = 14, 18 and 28 in Gnaphalium spp. and 2 n = 36 in Carpesium abrotanoides . Species in tribe Senecioneae are dominated by 10 based chromosome numbers, 2 n = 10 ( Emilia sonchifolia ) and 2 n = 40 ( Gynura crepidioides and Senecio spp.). However new lowest diploid chromosome number 2 n = 10 is reported in Senecio nudicaulis besides 2 n = 40. Tribes Mutiseae with 2 n = 18 ( Gerbera gossypina ) and Tageteae with 2 n = 48 (Tagetes minuta ) and tribe Vernonieae with 2 n = 18 and 20 in two species are also reported. Among these some species are found with new reports. Mathew and Mathew (1988) reported 2 n = 36 in the individuals of Cotula australis from South India. From outside India, 2n = 18, 36 and 40 chromosome numbers are reported for this species. Thus the present report of 2 n = 26 (Fig. 1a) is a varied chromosome count on worldwide basis based on x = 13, the most common basic number in the genus. Rashid (1986) described Aster lanceolatus as desynaptic with 2 n = 18. Other chromosome number counts (2 n = 32, 40, 48, 56 and 64) are from outside India which represent intraspecific polyploid series (4x, 6x and 8x) based on x = 8. Thus the present report of 2 n = 48 (Fig. 1b) is a new varied chromosome count from India.