Male Meiosis in Some Species of Genus Setaria L. from Different Geographical Regions of Rajasthan, India

Navjot Kaur* and Raghbir Chand Gupta

Department of Botany, Punjabi University, Patiala 147002, India

Received September 11, 2017; accepted October 11, 2017

Summary The genus Setaria L. have economically important as a grains and forage. The genus falls under the tribe Paniceae and family Poaceae. The cytological aspects of eight species and 16 accessions have been observed during the present research. S. geniculata (2n=54), S. homonyma (2n=18), and S. italica (2n=36) are the ﬁrst time reported euploid cytotypes reports from world, whereas, Setaria barbata (2n=54), S. glauca (2n=54), S. pumila (2n=54), S. verticillata (2n=108) are the ﬁrst cytological reports from Rajasthan. Almost all the species show normal meiosis with minor/negligible abnormalities like secondary associations and univalent. The data presented in the present research shows that the genus Setaria is the result of the multiple suppressor system due to which the genus shows more adaptability and stability than the other genera of family Poaceae.

Key words Male meiosis, Setaria, Paniceae, Rajasthan, Chromosome number, Polyploidy.

Setaria P. Beauv. is commonly known as ‘Fox tail accessions under eight species of the genus has been car- millets’ or ‘Bristle grass’ belongs to the tribe Paniceae ried out and explained here. in family Poaceae. The genus is cosmopolitan with 140 species throughout the temperate and tropical regions of Materials and methods the world (Cope 1982). Of which, 17 species are reported in India (Bor 1960). From the flora of Rajasthan, eight Floristic surveys were made in the rainy seasons to species are known taxonomically (Shetty and Singh collect different species of the genus Setaria from differ- 1993). The genus is characterized as annual or perennial ent unexplored sites of Rajasthan (Table 1). The validity grass; tufted cane like spikelets, spikelets are replaced of species name, including author citations were checked by one to many bristles like branchlets which remains and identified from Botanical Survey of India, Arid on the plant after the spikelets have fallen; glumes four Zone Regional Centre, Jodhpur. The voucher specimens membranous and styles quite distinct. Economically, the were submitted in Herbarium of the Department of Bot- genus is rich because the species cultivated as grains or any, Punjabi University, Patiala (PUN). The young inflo- forage are used as important food source for wildlife, rescences were collected on population basis from their whereas, some species are noxious weeds (Rominger natural habitats. These were then fixed in Carnoy’s fixa- 1962, Prasada Rao et al. 1987, Dekker 2003). S. verticil- tive (ethyl alcohol : chloroform : acetic acid; 6 : 3 : 1 v/v). lata is used as rodent repellents. S. italica is cultivated The meiotic studies were carried out on young inflores- worldwide that is used to cure chicken pox in small areas cence using standard acetocarmine smear technique. of Pakistan (Ahmad et al. 2010). Seeds of S. virdis have The detailed meiotic analysis on various stages was done nutritive value as cereal grains (Douglas et al. 1985). S. by taking 200–250 pollen mother cells (PMCs) to get ac- italica has been cultivated across southern Europe, some curate reports. Chromosome number in each accession regions of India, China, Korea and Japan (Gao and Chen was confirmed by studying more than one specimen 1988, Austin 2006). Some workers have contributed that from each population. At least 10 PMCs were analyzed the knowledge of the reproductive biology of this genus at different countable stages to confirm the chromosome is principally limited to the cultivated species (Pensiero number. Microsporogenesis is also studied. Tetrads with et al. 2005, Schrauf et al. 1998, Caponio and Pensiero four equal sized cells were taken as normal, and with 2002). different cell numbers varying from four, with cells of Kaur et al. (2011) studied male meiosis of 38 acces- different sizes or with the presence of micronuclei were sions and six species of Setaria from different parts of considered abnormal. Pollen fertility was observed with India. Still the cytology of this species from Rajasthan their stainability with acetocarmine technique. The well was lacking, detailed study on the male meiosis of 16 stained and well filled pollen grains are counted as fer- tile pollen grains, whereas, unstained and shrunken pol- * Corresponding author, e-mail: [email protected] len grains were considered as sterile. Photomicrographs DOI: 10.1508/cytologia.83.63 of the PMCs were taken from freshly prepared slides 64 N. Kaur and R. C. Gupta Cytologia 83(1)

Table 1. Data showing voucher details, chromosome numbers (2n), ploidy level, pollen viability (%) and remarks of 8 species of genus Setaria studied presently.

Name of Chromosome Ploidy Pollen viability Locality/Altitude (m) Remarks the species No. (2n) level (%)

1. Setaria barbata (Lam.) Kunth P-1 60988 Gulab bagh, Udaipur/ 600 54 6x 92.7 New report from Rajasthan

2. S. geniculata (Lam.) P. Beauv. (=S. parviﬂora (Poir.) M. Kerguelen) P-1 60992 Mount Abu/ 1220 54 6x 89.3 New euploid cytotype from world

3. S. glauca (L.) P. Beauv. (=Pennisetum glaucum (L.) R. Br.) P-1 60995 Jhalawar/ 312 54 6x 72.5 New report from Rajasthan

4. S. homonyma (Steud.) Chiov. P-1 60200 On the way to Guru Sikhar, Mount Abu/ 1220 18 2x 96.1 New euploid cytotype from world

5. S. intermedia Roem. & Schult. P-1 60150 Sirohi/ 321 36 4x 87.5 Conﬁrms previous reports P-2 60997 Mount Abu/ 1220 36 4x 99.3

6. S. italica (L.) P. Beauv. P-1 60991 Herbal garden, Jhalawar/ 312 36 4x 100 New euploid cytotype from world P-2 60996 Mount Abu/ 1220 36 4x 87.4

7. S. pumila (Poir) Roem & Schult. P-1 60178 Kali Sind Dam, Jhalawar/ 460 54 4x 91.2 First cytological report from Rajasthan P-2 60191 D.C. Kothi, Jhalawar/ 460 54 4x 99.5 P-3 60990 Pali/ 218 54 4x 89.6 P-4 60994 Collector house, Jhalawar/ 312 54 4x 76.8

8. S. verticillata (L.) P. Beauv. P-1 60172 Monsoon Palace, Udaipur/ 598 108 12x 98.8 New cytological report from Rajasthan P-2 60989 Gulab bagh, Udaipur/ 598 108 12x 86.1 P-3 60993 Ranakpur/ 486 108 12x 82.4 P-4 60105 Gajner Palace, Bikaner/ 242 108 12x 97.2

using 80i Eclipse digital imaging system (Nikon). bivalents (2n=54) at M-I (Fig. 3) and equal distribution of 27 : 27 chromosomes at anaphase-I (Fig. 4). The Results and discussion meiosis is found to be normal. The present chromosome count is a new report from world. Previously, tetraploid The meiotic analysis has been carried out on eight chromosome report with 2n=36 has been reported from species of Setaria in Rajasthan, the details of which are India (Mehra 1982) whereas, both the tetraploid and concise in Table 1 which includes the name of the spe- octaploid reports are also known from other countries cies, accession number, locality with altitude, chromo- of world. Sacchet et al. (1980) also observed aneuploid some numbers, ploidy level, pollen fertility and remarks. cytotype for the same species with 2n=22. During the present research 16 accessions covered under In S. glauca (L.) P. Beauv.: During the present study, eight species belonging to the genus Setaria have been studied population revealed a hexaploid cytotype with reported. The meiotic study in all the studied species is 27 bivalents (2n=54) at diakinesis (Fig. 5). Some PMCs briefly described as follows: show secondary association of bivalents at M-I (Fig. 6). In Setaria barbata (Lam.) Kunth, single popula- The present report is in conformity with the previous re- tion have been worked out, hexaploid with 27 bivalents ports from India and outside India. This is the new cyto- (2n=54) at meiotic metaphase-I (M-I) (Fig. 1). It shows logical report for the species from Rajasthan. Previously, normal 27 : 27 distribution of chromosomes at anaphase- tetraploid chromosome count of 2n=36 was reported I (A-I) (Fig. 2). Kaur et al. (2011) from Kangra reported from the studied area i.e. Rajasthan. The species show tetraploid cytotypes i.e. 2n=36 whereas, Christopher intraspecific polyploids based on x=9, of these cytotypes and Abraham (1976) reported hexaploid (2n=54) from 2n=36 is the most common. South India. Furthermore, Sarkar et al. (1976) observed S. homonyma (Steud.) Chiov.: The present investiga- aneuploid cytotype with 2n=56 from India. tion reveals the presence of nine bivalents (2n=18) at In S. geniculata (Lam.) P. Beauv. (=S. parviflora M-I (Fig. 7). In some of the PMCs it shows univalent for-

(Poir.) M. Kerguelen), presently, single population col- mation (8II+2I) at M-I (Fig. 8). The present diploid count lected from Mount Abu has been investigated cytologi- is a new euploid cytotype for the species from world. cally. The hexaploid cytotype reveals the presence of 27 Majority of the previous reports are of tetraploid cyto- 2018 Male Meiosis in Some Species of Genus Setaria L. from Different Geographical Regions of Rajasthan, India 65

Figs. 1–13. Meiotic ﬁgures in PMCs of eight Setaria species. 1. PMC of S. barbata showing 27 bivalents at M-I. 2. PMC with 27 : 27 distribution of chromosomes at A-I. 3. S. geniculate. PMC with 27 bivalents at M-I. 4. PMC showing 27 : 27 chromosomes at A-I. S. glauca. 5. PMC showing 27 bivalents at diakinesis. 6. Secondary associations. S. hom-

onyma. 7. PMC with nine bivalents at M-I. 8. PMC showing 8II+2I at M-I (arrow). S. intermedia. 9. PMC showing 18 bivalents at M-I. 10. PMC with 18 : 18 chromosomes at A-I. S. italica. 11. PMC showing 18 : 18 chromosomes at A-I. S. pumila. 12. PMC with 27 bivalents at diakinesis. S. verticillata. 13. PMC showing 2n=108 at mixed A-I. Scale bar=10 µm. type with 2n=36 from India and outside India. Other an- cytotype with 2n=38 (Bir and Sahni 1987, Sahni 1989) euploid cytotypes with 2n=20 and 2n=32 have also been from Punjab plains. reported for the species. Thus, the species shows intra- S. italica (L.) P. Beauv.: During the present study, two specific polyploidy based on x=8, 9 and 10. Of which, populations have been collected from different locali- x=9 is the most common base number. ties of Rajasthan. Both the populations show the similar S. intermedia Roem. & Schult.: Presently, two popu- meiotic chromosome number of 2n=36 with 18 : 18 equal lations have been analyzed meiotically. Both the popu- distribution of chromosomes at A-I (Fig. 11). All the lations show same chromosome count of 2n=36, 18 workers had reported diploid report with 2n=18 from bivalents at M-I (Fig. 9) and equal distribution of 18 : 18 India and outside. This is a new tetraploid report from chromosomes at A-I (Fig. 10). The present chromosome world for the species based on the basic chromosome report is in conformity with previous reports from India number x=9. and outside India and also from the studied area i.e., Ra- S. pumila (Poir) Roem & Schult.(=S. pallide-fusca jasthan (Kaur and Gupta 2016). The species shows intra- (Schumach.) Stapf & C. E. Hubb.): Presently, four specific polyploid races with 2n=18, 36, 54 and 72 based populations of the species have been worked out cyto- on the basic chromosome number of x=9 and aneuploid logically. All the populations show normal meiosis with 66 N. Kaur and R. C. Gupta Cytologia 83(1) chromosome count of 2n=54. It shows the presence of References 27 bivalents (2n=54) at diakinensis (Fig. 12). The present result is in conformity with the previous reports Ahmad, F., Khan, M. A., Ahmad, M., Zafar, M., Mahmood, T., Ja- from India and other countries of the world. This is a been, A. and Marwat, S. K. 2010. Ethnomedicinal uses of grasses in salt range regions of Northern Pakistan. J. Med. Plants Res. 4: first cytological report from Rajasthan. 362–369. S. verticillata (L.) P. Beauv.: The present chromosome Austin, D. F. 2006. Fox-tail millets (Setaria: Poaceae)̶abandoned count of 2n=108 (Fig. 13) forms the first cytological food in two hemispheres. Econ. Bot. 60: 143–158. report for the species from the study area i.e. Rajast- Bir, S. S. and Sahni, M. 1987. Chromosomal and morphological varia- han. It is dodecaploid ploidy level based on x=9. Same tions in grasses of Punjab. Cytol. Genet. 22: 12–22. Bir, S. S. and Sahni, M. 1986. In: Bir, S. S. (ed.). 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