© 2018 The Japan Mendel Society Cytologia 83(2): 215–220

New and Varied Chromosome Reports in Twenty-Six Species of the Family from Cold Deserts of the Western Himalaya

Raghbir Chand Gupta1, Henna Goyal1, Rajesh Kumar Goel2 and Vijay Singh1,3*

1 Department of Botany, Punjabi University, Patiala, Punjab, -147002 2 Department of Pharmaceutical and Drug Research, Punjabi University, Patiala, Punjab, India-147002 3 Department of Botany, Mata Gujri College, Fatehgarh Sahib, Punjab, India-140407

Received December 10, 2017; accepted February 18, 2018

Summary In the present study, 26 species belonging to 17 genera and five tribes of the family Asteraceae collected from natural habitats in cold deserts of the western Himalaya were analyzed on their meiotic events. The chromosome counts are reported for the first time in four species viz. Brachyactis roylei (n=9), Cousinia thomsonii (n=12), Erigeron umbrosus (n=9) and Waldheimia glabra (n=9). The euploid or aneuploid chromo- some numbers are reported in Arctium lappa (n=17), (n=5) and Saussurea heteromalla (n=8). The chromosome counts of n=9 in E. borealis is new to the Indian populations. The chromosome counts in the rest 18 species are new additions from the cold desserts of Lahaul-Spiti district of . The present meiotic study also discloses meiotic abnormalities viz. chromosome stickiness, cytomixis, laggards, etc. leading to the hypo-/or hyperploid meiocytes. These abnormalities further may alter pollen viability by producing unre- duced pollen grains and might be responsible for the disparity in the chromosome numbers.

Key words Asteraceae, Chromosome number, Meiotic study, Cold desert, Western Himalaya.

Asteraceae is the largest and most diverse family Materials and methods among Angiosperms. The family comprises about 22750 species belonging to 1528 genera (Judd et al. 2008) The materials were collected from the unex- and about 900 species in 167 genera in India (Hajra plored sites of Lahaul-Spiti district (32°30′N, 77°50′E) et al. 1995). The family comprises 10 subfamilies and of Himachal Pradesh. The young flower buds were col- 35 tribes (Panero and Funk 2002). For the most evolved lected on population’s basis (three to seven individuals status in dicots, therefore we pay attention to the family of each population) and fixed in Carnoy’s fixative (etha- with an evolutionary interest. nol : chloroform : acetic acid, 6 : 3 : 1). The anthers were The western Himalaya, an integral part the Great crushed in 1% acetocarmine and studied under a mi- Himalayan range is always bequeathed diverse phyto- croscope. The accurate chromosome numbers and other geographic range, not only support huge floristic diver- meiotic abnormalities were confirmed by observing sity, also offers a scope for assessing phytogeographical 200–250 pollen mother cells (PMCs) at various meiotic interest. Previously a significant number of attempts stages for each specimen in each population of species. were made on the floristic diversity of higher altitude Meiotic index in the species with abnormal microsporo- of western Himalaya, but still, there is a lacuna in chro- genesis was calculated. mosomal studies, particularly for this group. The lack of The staining tests using a glycerol–acetocarmine cytological work in this area is due to the tough climatic (1 : 1) mixture and 1% aniline blue were performed and conditions and abrupt topography. Thus, keeping in view up to 500–800, pollen grains were examined for esti- the cumulative reputation of Asteraceae and importance mate pollen fertility and size. The fully stained pollen of cytology in evolutionary studies, the present study grains were taken as apparently fertile, while shriveled covers the species from various localities of Lahaul-Spiti and unstained pollen as sterile. The photomicrographs district, an ecologically fragile cold dessert in the west- of PMCs at various meiotic stages were taken from pre- ern Himalayas. pared slides using a Leica DM 3000 imaging system. The voucher specimens were deposited in the Herbarium of Department of Botany, Punjabi University Patiala (PUN), only after consulting various local floral keys and confirming finally from Botanical Survey of India * Corresponding author, e-mail: [email protected] (BSI), Dehra Dun. DOI: 10.1508/cytologia.83.215 216 R. C. Gupta et al. Cytologia 83(2)

Table 1. The table showing taxa, localities with altitudes, accession number, chromosome number (2n), ploidal level, pollen viability and re- marks about the studied species from cold desserts of Lahaul-Spiti (Himachal Pradesh).

Locality (altitude in Chromosome Pollen Species Pollen size meters), geographical number (n)/ viability Remarks PUN (*) (µm±S.E.) co-ordinates Ploidy level (x) (%)

Tribe: Anthemideae Anthemis cotula L. 58560 (3) Keylong (3335), 9/2x 89 23.3±0.2×22.8±0.4 New record from the 32°34′19″N, 77°02′07″E district Artemisia macrocephala Jacq Baralacha La (4883), 9/2x 70 23.1±0.5×20.7±1.1 First report from World ex Bess. 52785 (4) 32°45′37″N, 77°25′12″E Waldheimia glabra (Decne.) Baralacha La (4883), 9/2x 100 21.5±1.2×22.3±1.5 First report from World Regel 52791 (4) 32°45′37″N, 77°25′12″E W. tomentosa (Decne.) Regel Baralacha La (4883), 9/2x 100 20.5±1.5×23.1±1.4 New record from the 52792 (8) 32°45′37″N, 77°25′12″E district Tribe: Asterae Aster indamellus Grierson Chandra Lake (4300), 9/2x 92 26.1±1.1×25.4±1.4 New report from the 52762 (3) 32°28′57″N, 77°36′53″E district Bellis perennis L. 52776 (8) Kibber (3080), 5/2x 92 16.8±0.9×17.4±0.6 New varied report from 32°19′51″N, 78°00′34″E World Brachyactis roylei (DC.) Keylong (3335), 9/2x 57 21.2±2.4×20.4±1.6 First chromosome report Wendelbo 52778 (3) 32°34′19″N, 77°02′07″E from World Erigeron alpinus L. 52797 (3) Keylong (3335), 9/2x 60 21.4±1.2×24.3±0.6 New report from the 32°34′19″N, 77°02′07″E district E. borealis (Vierh.) Simm Baralacha La (4883), 9/2x 84 24.3±0.6×24.3±0.6 First chromosome report 52796 (4) 32°45′37″N, 77°25′12″E from India E. umbrosus (Kar. & Kir.) Boiss Koksar (3160), 9/2x 95 14.2±0.9×15.4±1.6 First chromosome report 52777 (7) 32°24′30″N, 77°15′5″E from World Tribe: Cynareae Arctium lappa L. 52794 (3) Koksar (3140), 17/2x 89 22.2±0.8×23.4±1.2 Varied chromosome 32°24′30″N, 77°15′5″E report from the World Cousinia thomsonii C. B. Chhatru (3360), 12/2x 80 27.8±1.3×28.7±0.9 First chromosome report Clarke. 57517 (5) from world Saussurea heteromalla (D. Don) Koksar (3140), 8/2x 90 38.1± 1.5×37.5±1.8 Varied chromosome Hand.-Mazz. 58532 (3) 32°24′30″N, 77°15′5″E report from World S. jacea (Klotzsch) C. B. Tandi, Keylong (3335), 17/2x 88 31.3±0.9×32.1±1.3 Varied chromosome Clarke. 57516 (4) 32°34′40″N, 77°1′36″E report from World Tribe: Cichoreae Hieracium crocatum Fries Trilokinath (2760), 5/2x 88 15.0±0.6×20.0±0.9 First chromosome report 52760 (6) 32°42′0″N, 76°41′0″E from World H. umbellatum L. Keylong (3080), 27/6x 91 16.8±0.7×20.1±1.2 Varied chromosome 52774 (3) 32°34′19″N, 77°02′07″E report from World Lactuca brunoniana (Wallich) Tandi, Keylong (2960), 8/2x 90 22.9±1.6×23.6±1.1 New report from the C. B. Clarke. 52764 (5) 32°34′40″N, 77°1′36″E district La. dolichophylla Kitam , Lahaul (3100), 8/2x 79 22.9±0.9×21.3±0.4 New report from the 57504 (7) 32°29′0″N, 77°7′0″E district La. lessertiana (Wallich ex DC.) Batal, Spiti (4300), 8/2x 70 22.1±0.9×23.8±1.3 First chromosome report C. B. Clarke. 52766 (6) 32°21′28″N, 77°37′10″E from India Tragopogon dubius Scop. Keylong (3080), 14/4x 90 20.0±1.3×22.0±0.8 Varied chromosome 57505 (6) 32°34′19″N, 77°02′07″E reports from World T. gracilis D. Don 52769 (3) Sissu (3100), 7/2x 70 22.7±0.7×20.1±1.4 Varied chromosome 32°29′0″N, 77°7′0″E report from World Youngia glauca Cass. 52773 (4) Zingzingbar (4270), 7/2x 83 18.6±1.1×19.9±1.4 First chromosome report 32°47′30″N, 77°19′28″E from World Y. tenuifolia (Willd.) Babcock & (3200), 5/2x 87 17.9±0.9×20.1±1.2 First chromosome report Stebbins 52771 (4) 32°38′0″N, 77°10′0″E from India Tribe: Inulae A. contorta (D. Don) HooK. f. Keylong (3340), 14/4x 70 26.4±1.3×29.1±1.4 New report from the Willd. 52782 (3) 32°34′19″N, 77°02′07″E district Inula grandiflora 52781 (4) Gondhla (3160), 8/2x 79 23.5±1.2×23.1±0.9 New report from the 32°28′14″N, 77°00′02″E district Leontopodium alpinum Cass. Chandra Tal Lake (4300), 12/2x 68 31.6±1.4×33.2±1.3 New report from the 57494 (5) 32°28′57″N, 77°36′53″E district

*Numbers in the parenthesis represent the number of individual considered for the present study from a particular population.

Results and discussion done from the cold deserts of the western Himalaya. The data pertaining to the name of species, accession num- The present study comprises the cytological work ber, flowering and fruiting periods, pollen size, pollen 2018 Chromosome Reports in Asteraceae Species from Cold Deserts of the Western Himalaya 217

Fig. 1. Meiotic chromosomes of PMCs in 26 Asteraceae species in cold desert of western Himalaya. (1a) An. cotula, PMC

with 9II at diakinesis; (1b) Art. macrocephala, PMC showing 9 : 9 chromosomal distribution at AI; (1c) W. glabra, PMC showing 9 : 9 chromosomal distribution at AI; (1d) W. tomentosa, PMC with 9II at diakinesis; (1e) As. indamellus, PMC showing 9 : 9 chromosomal distribution at AI; (1f) Be. perennis, PMC with 5II at diakinesis; (1g) Br. roylei, PMC with 9II at diakinesis; (1h) E. alpinus, PMC with 9II at MI; (1i) E. borealis, PMC with 9II at MI; (1j) E. umbrosus, PMC with 9II at diakinesis; (1k) Arc. lappa, PMC with 17II at diakinesis; (1l) C. thomsonii, PMC with 12II at MI; (1m) S. heteromalla, PMC with 8 : 8 chromosomes at each pole; (1n) S. jacea, PMC with 17II at MI; (1o) H. crocatum, PMC with 5II at MI; (1p) H. umbellatum, PMC with 27II at diakinesis; (1q) La. brunoniana, PMC with 8II at MI; (1r) L. dolichophylla, PMC with 8II at MI; (1s) L. lessertiana, PMC with 9II at MI; (1t) T. dubius, PMC with 14II at diakinesis; (1u) T. gracilis, PMC with 7II at MI; (1v) Y. glauca, PMC with 7II at diakinesis; (1w) Y. tenuifolia, PMC with 5II at diakinesis; (1x) An. contorta, PMC with 14II at MI; (1y) I. grandiflora, PMC with 8II at MI; (1z) Le. alpinum, PMC with 12II at MI. Scale bars=10 µm. fertility, and locality details of 30 populations belonging chromosome number of n=9 (Fig. 1d). The chromosome to 26 species of family Asteraceae from cold desserts of count varied from previous report of 2n=24 (Singh et al. western Himalaya is given in Table 1. 2016). Aster indamellus Grierson had a chromosome count Chromosome numbers of n=9 (Fig. 1e), which is in conformity to the previous Anthemis cotula L. revealed n=9 (Fig. 1a), which sup- report of 2n=18 and 54 (Probatova et al. 2011). ports early count from Kashmir Himalaya (Mehra and Bellis perennis L. had a chromosome count of n=5 Remanandan 1974). (Fig. 1f). Previously, the other cytotype (2n=18) has Artemisia macrocephala Jacq ex Bess. had a chromo- been reported (Mehra and Remanandan 1974). some number of n=9 (Fig. 1b). This supports previous Brachyactis roylei (DC.) Wendelbo had a chromo- report (Garcia 2006). some count of n=9 (Fig. 1g). This count supports previ- Waldheimia glabra (Decne.) had a chromosome count ous report (Podlesch and Dieterle 1969). of n=9 (Fig. 1c). W. tomentosa (Decne.) Regel had a Erigeron alpinus L. had a chromosome count of n=9 218 R. C. Gupta et al. Cytologia 83(2)

Table 2. Data on different meiotic abnormalities in 26 species of Asteraceae from cold deserts of the western Himalaya.

Cytomixis Meiotic course

PMCs with PMCs with Species PMCs involved No. of PMCs PMC with bridges PMCs with chromosomal unoriented laggards (%) involved (%) (at AI, II/TI, II) (%) (at AI, II/TI, II) (%) stickiness at MI (%) bivalents at MI (%)

An. cotula 24.0 (30/125) 2–6 3.1 (4/129) 7.2 (9/125) 10.1 (12/114) 10.6 (13/123) Art. macrocephala 24.6 (31/126) 2–6 5.5 (6/109) 2.3 (3/130) 4.3 (6/139) 1.5 (2/129) W. glabra 22.5 (22/111) 2–4 11.6 (14/121) 13.8 (16/118) 26.9 (38/141) 10.4 (14/134) W. tomentosa 1.6 (27/124) 1–2 0 -/- -/- -/- As. indamellus 5.5 (6/109) 1–2 22.1 (29/131) 0.1 (1/101) 5.4 (6/110) -/- Be. perennis 0.9 (1/113) 1–2 3.5 (4/113) 1.8 (2/113) -/- 1.8 (2/113) Br. roylei 22.2 (30/135) 2–6 14.5 (19/131) 5.5 (7/128) 33.7 (42/127) 3.2 (4/123) E. alpinus 1.7 (2/160) 1–2 1.4 (2/145) 2.4 (3/126) 1.2 (2/125) 23.1 (34/147) E. borealis 2.4 (3/123) 1–2 0 -/- -/- -/- E. umbrosus 12.8 (15/117) 2–3 13.9 (18/129) 13.8 (16/116) 12.5 (17/136) 10.4 (14/134) Arc. lappa 5.4 (6/111) 1–2 3.2 (4/126) 2.7 (3/111) -/- -/- C. thomsonii 1.8 (2/112) 1–2 7.2 (9/125) -/- 39.5 (51/129) 3.8 (8/130) S. heteromalla 4.4 (5/144) 2–3 5.5 (6/109) 8.4 (11/131) 3.8 (8/130) 10.1 (12/114) S. jacea 1.0 (1/97) 1–2 11.3 (11/97) 6.1 (6/97) 7.21 (7/97) 2.06 (2/97) H. crocatum -/- 0 0 -/- -/- -/- H. umbellatum L. 2.4 (3/125) 1–2 4.0 (5/125) -/- -/- -/- La. brunoniana -/- 0 0 1.8 (2/111) 4.5 (5/111) 0.8 (1/111) La. dolichophylla -/- 0 0 -/- -/- -/- La. lessertiana 2.8 (4/144) 1–2 0 2.0 (3/144) -/- -/- T. dubius 8.4 (9/107) 1–4 4.1 (5/123) 11.4 (15/132) 8.7 (9/103) 10.2 (13/128) T. gracilis 0 0 2.8 (3/105) 4.1 (5/123) 9.3 (11/118) 16.5 (21/130) Y. glauca 2.0 (2/100) 1–2 0 0 2.8 (3/107) 2.7 (3/109) Y. tenuifolia 14.5 (19/131) 2–4 11.1 (12/108) 0 0 -/- An. contorta 2.5 (3/119) 1–2 15.1 (18/119) 4.7 (5/107) 9.8 (1/103) 7.9 (6/76) I. grandiflora 15.5 (18/116) 2–6 0 9.8 (11/112) 12.6 (15/119) 8.8 (10/120) Le. alpinum 0 0 21.9 (23/105) 4.6 (5/123) 9.3 (11/118) 16.15% (21/130)

Figs in parenthesis denote observed number of abnormal PMCs in the numerator and total PMCs observed in the denominator. AI: anaphase I: AII anaphase II, MI: metaphase I, MII: metaphase II, TI: telophase I, TII: telophase II. *Numbers in the parenthesis represent the number of individual considered for the present study from a particular population. (Fig. 1h), which supports previous reports (Mehra and been reported (Gupta et al. 2014). Remanandan 1974). E. borealis (Vierh.) Simm. had a Lactuca brunoniana (Wallich) C. B. Clarke. had a chromosome count of n=9 (Fig. 1i). This count supports chromosome count of n=8 (Fig. 1q), which supports previous reports (Lövkvist and Hultgård 1999). E. um- previous report (Gupta et al. 2014). La. dolichophylla brosus (Kar. & Kir.) Boiss. had a chromosome count of Kitam had a chromosome count of n=8 (Fig. 1r) which n=9 (Fig. 1j) which supports previous report (Kaur and supports previous report (Gupta et al. 2014). La. lesser- Singhal 2013). tiana (Wallich ex DC.) C. B. Clarke. had a chromosome Arctium lappa L. had a chromosome count of n=17 count of n=8 (Fig. 1s), this is the first report of chromo- (Fig. 1k). The chromosome count supports previous some count in this species. report (Ge et al. 1989). The other chromosome counts Tragopogon dubius Scop. had chromosomes of n=14 with 2n=18 (Mehra et al. 1965), 2n=32 (Bala and Gupta (Fig. 1t). This is a new count, but earlier known with 2011) and 2n=36 (Ma et al. 1990) has been also re- 2n=12, 24 and 36 (Gupta et al. 2014). T. gracilis D. Don ported. had a chromosome count of n=7 (Fig. 1u), which is a Cousinia thomsonii C. B. Clarke. had a chromosome new chromosome count. The other chromosome count of count of n=12 (Fig. 1l), this is the first report of chromo- 2n=12 (Gupta et al. 2014) has been reported. some count in this species. Youngia glauca Cass. had the chromosome number of Saussurea heteromalla (D. Don) Hand.-Mazz. had n=7 (Fig. 1v). Earlier, the chromosome count of 2n=16 a chromosome count of n=8 (Fig. 1m). Two chromo- was reported (Kaur and Singhal 2013). Y. tenuifolia some counts with 2n=32 (Mehra et al. 1965) and 2n=34 (Willd.) Babcock & Stebbins had a chromosome report (Gupta and Gill 1983) have been also reported. S. jacea of 2n=16 (Fig. 1w), which confirms the earlier reports (Klotzsch) C. B. Clarke. had a chromosome count of (Gupta et al. 2014). n=14 (Fig. 1n) which supports previous report (Singh Anaphalis contorta (D. Don) HooK. f. had a chromo- et al. 2017). some report of n=14 (Fig. 1x) that supports previous Hieracium crocatum Fries had a chromosome count report (Gupta et al. 1989). of n=5 (Fig. 1o). H. umbellatum L. showed n=27 (Fig. Inula grandiflora Willd. had a chromosome count of 1p). Other chromosome counts of 2n=18 and 27 has n=8 (Fig. 1y). This chromosome count supports previ- 2018 Chromosome Reports in Asteraceae Species from Cold Deserts of the Western Himalaya 219

Table 3. Data on the abnormal microsporogenesis on twenty six species of Asteraceae from cold deserts from the western Himalaya.

Taxon/Accession Monads Dyads Triads Tetrads Meiotic index numbers PUN (*) WM (%) WM (%) WMN (%) WM (%) WMN (%) WM (%)

An. cotula -/- -/- 1.0 (1/103) -/- 2.9 (3/103) -/- 96.1 (99/103) Art. macrocephala 2.8 (3/109) 0.9 (1/109) 2.8 (3/109) -/- 2.8 (3/103) -/- 90.8 (99/109) W. glabra -/- -/- 1.8 (2/111) 2.7 (3/111) 1.8 (2/111) 3.6 (4/111) 90.1 (100/111) W. tomentosa 1.5 (2/130) 0.8 (1/130) 2.3 (3/130) 3.1 (4/130) 2.3 (3/130) 3.8 (5/130) 86.2 (112/130) As. indamellus -/- -/- 1.6 (2/120) -/- -/- -/- 98.3 (118/120) Be. perennis -/- -/- 3.1 (4/129) 0.8 (1/129) 7.0 (9/129) 8.5 (11/129) 80.6 (104/117) Br. roylei 0.9 (1/115) 2.6 (3/115) -/- 2.6 (3/115) 1.7 (2/115) 92.2 (106/115) E. alpinus -/- 0.9 (1/115) 2.6 (3/115) -/- 2.6 (3/115) 1.7 (2/115) 92.2 (106/115) E. borealis 1.7 (2/121) 4.1 (5/121) 0.8 (1/121) 4.1 (5/121) 0.8 (1/121) 0.8 (1/121) 87.6 (106/121) E. umbrosus -/- -/- 0.8 (1/125) -/- 1.6 (2/125) 0.8 (125) 96.8 (121/125) Arc. lappa -/- -/- -/- -/- -/- -/- 100 (121/121) C. thomsonii -/- -/- 1.7 (2/121) -/- 4.1 (5/121) -/- 95.0 (115/121) S. heteromalla -/- 1.0 (1/105) -/- 1.9 (2/105) 2.8 (3/105) 1.1 (2/105) 92.4 (97/105) S. jacea -/- -/- -/- -/- -/- -/- 100 (123/123) H. crocatum -/- 0.8 (1/124) 1.6 (2/124) -/- 4.0 (5/124) 1.6 (2/124) 91.9 (114/124) H. umbellatum L. -/- 1.1 (1/95) -/- -/- -/- -/- 99.2 (139/140) La. brunoniana -/- 1.7 (2/116) 4.3 (5/116) 2.6 (3/116) 4.3 (5/116) 5.2 (6/116) 81.9 (95/116) La. dolichophylla -/- 1.5 (2/130) 0.8 (1/130) 0.8 (1/130) 2.3 (3/130) 1.5 (2/130) 93.1 (121/130) La. lessertiana -/- -/- 0.8 (1/125) 2.4 (3/125) 0.8 (1/125) 1.6 (2/125) 94.4 (118/125) T. dubius 2.3 (2/87) 4.6 (4/87) 1.1 (1/87) 2.3 (2/87) 3.4 (3/87) 11.5 (10/87) 72.4 (63/87) T. gracilis 0 1.8 (2/117) 7.7 (9/117) 0 11.9 (14/117) 5.1 (6/117) 73.5 (86/117) Y. glauca 2.9 (4/139) 3.6 (5/139) 5.0 (7/139) 2.9 (4/139) 5.8 (8/139) 7.9 (11/139) 71.9 (100/139) Y. tenuifolia 0 1.7 (2/116) 4.3 (5/116) 2.6 (3/116) 4.3 (5/116) 5.2 (6/116) 81.9 (95/116) A. contorta 2.2 (3/133) 1.5 (2/133) 3.8 (5/133) 2.2 (3/133) 8.3 (11/133) 6.8 (9/133) 75.2 (100/133) I. grandiflora 2.5 (3/121) 0.8 (1/121) 4.9 (6/121) 0 6.6 (8/121) 2.5 (3/121) 82.6 (100/121) Le. alpinum 2.3 (3/133) 1.5 (2/133) 3.8 (5/133) 2.3 (3/133) 8.3 (11/133) 6.8 (9/133) 75.2 (100/133)

Fig.s in parenthesis denote observed number of abnormal PMCs in the numerator and total number of PMCs observed in denominator, WMN - without micronuclei, WM with micronuclei. *Numbers in the parenthesis represent the number of individual considered for the present study from a particular population. ous work (Shetty 1967). Another chromosome count of Cytomixis may be the result of physical factors like 2n=32 had also been reported (Gupta et al. 2017). temperature stress (Narain 1976), under even direct Leontopodium alpinum Cass. had a chromosome genetic control (Haroun et al. 2004) or genetic control count of n=12 (Fig. 1z). coupled with stress factors (Baptista-Giacomelli et al. 2000, Ghanima and Talaat 2003), and due to retention Meiotic irregularities of wide plasmodesmata during cell wall formation in Twenty one species out of 26 species were observed pre-meiotic stage (Yu et al. 2004). It plays a vital role in with many or at least one type of meiotic abnormal- chromosomal diversity or even in evolution, as it forms ity, which leads to the abnormal microsporogenesis and hypo-/hyperploid-cells and uneven gametes afterward ultimately reduce pollen viability (Table 1). The most (Levan 1941, Zheng et al. 1987, Kim et al. 2009). common meiotic abnormality (Table 2) observed was Chromosome stickiness is thought to be a consequence cytomixis (21 spp.), followed by chromatin stickiness of mutated sticky gene that disrupts the conformation (18 spp.), bridges and laggards (17 spp. each). The fre- of some histone proteins (Utsunomiya et al. 2005, Pa- quency of cytomixis was found to be maximum in Ar. gliarini et al. 2008). Chromatin bridges are generally crocephala (24.6%), followed by An. cotula (24.0%) formed due to malfunctioning of non-histone proteins or and W. glabra (22.5%). Another unusual meiotic phe- paracentric inversions (Gaulden 1987), or they might be nomenon, chromatin stickiness is reported in eighteen due to telomeres in breakage-fusion-bridge cycles (Jones species (Table 2) with a maximum frequency in As. 2005). The laggards are the result of either spindle ab- indamellus (22.1%), followed by L. alpinum (21.9%) and normalities or late chiasma terminalization (Pagliarini An. contorta (15.1%). Further, the meiotic abnormali- 2000). These abnormalities result in abnormal microspo- ties like laggards and bridges are also observed with a rogenesis (Utsunomiya et al. 2002) with the formation maximum frequency in E. alpinus (23.1%) and S. het- of monads, dyads, triads with or without micronucleus eromalla (39.5%), respectively. Besides, the abnormal (Table 3), or even micro-pollen/uneven gametes, sub- microsporogenesis with the formation of monads, dyads, sequently (Table 3). Thus, the formation of aneuploid triads with or without micronucleus or micronuclei even gametes by cytomixis (Falistocco et al. 1995, Ghaffari in normal tetrads are also observed in case of E. borealis 2006), produce heterogeneity in pollen grains, which and H. umbellatum (Table 3). might be attributed to the genetic (Fadaei et al. 2010) 220 R. C. Gupta et al. Cytologia 83(2) or environmental reasons (Nirmala and Rao 1996), and Jones, R. N. 2005. McClintock’s controlling elements: the full story. even to, the genome-environment interaction (Baptista- Cytogenet. Genome Res. 109: 90–103. Judd, W. S., Campbell, C. S., Kellogg, E. A., Stevens, P. F. and Dono- Giacomelli et al. 2000). ghue, M. J. 2008. Plant Systematics̶A Phylogenetic Approach, 3rd edn. Sinauer Associates, Inc., Sunderland. Acknowledgements Kaur, D. and Singhal, V. K. 2013. In: Marhold, K. and Breitwieser, I. (eds.). IAPT/IOPB chromosome data 13. Taxon 61: 6–7. The authors are thankful for the IPLS-DBT (Refer- Kim, J. S., Oginuma, K. and Tobe, H. 2009. Syncyte formation in the microsporangium of Chrysanthemum (Asteraceae): a pathway to ence number: BT/PR 4548/INF/22/146/2012) and UGC- infraspecific polyploidy. J. Plant Res. 122: 439–444. BSR Fellowship scheme (Ref. No. 6007/04-04-2013) for Levan, A. 1941. Syncyte formation in the pollen mother cells of hap- the financial assistance. The authors are highly thankful loid Phleum pratense. Hereditas 27: 243–253. to Lt. Shri Krishan Lal Ji for sharing his tremendous Lövkvist, B. and Hultgård, U. M. 1999. Chromosome numbers in field knowledge during the collection plant materials. south Swedish vascular . Opera Bot. 137: 1–42. Ma, X. H., Ma, X. Q. and Li, N. 1990. Chromosome observation of some drug plants in Xinjiang. Acta Bot. Boreal. Occid. Sin. 10: References 203–210. Mehra, P. N., Gill, B. S., Mehta, J. K. and Sidhu, S. S. 1965. Cyto- Bala, S. and Gupta, R. C. 2011. In: Marhold, K. (ed.). IAPT/IOPB logical investigations on the Indian Compositae. I. North-Indian chromosome data 12. Taxon 60: 1784–1786. taxa. Caryologia 18: 35–68. Baptista-Giacomelli, F. R., Pagliarini, M. S. and Almeida, J. L. 2000. Mehra, P. N. and Remanandan, R. 1974. Cytological investigations Meiotic behavior in several Brazilian oat cultivars (Avena sativa on Indian Compositae II. Astereae, Heliantheae, Helenieae and L.). Cytologia 65: 371–378. Anthemideae. Caryologia 27: 255–284. Fadaei, F., Sheidai, M. and Asadi, M. 2010. Cytological study of the Narain, P. 1976. Cytomixis in pollen mother cells of Hemerocallis L. Arenaria L. (Caryophyllaceae). Caryologia 63: 149–156. Curr. Sci. 48: 996–998. Falistocco, E., Tosti, N. and Falcinelli, M. 1995. Cytomixis in pollen Nirmala, A. and Rao, P. N. 1996. Genetics of chromosome numerical mother cells of diploid Dactylis one of the origins of 2n gametes. mosaicism in higher plants. Nucleus 39: 151–175. J. Hered. 86: 448–453. Pagliarini, M. S. 2000. Meiotic behavior of economically important Garcia, S., Garnatje, T., Dariimaa, S., Tsooj, S. and Vallès, J. 2006. plant species: the relationship between fertility and male sterility. New or rarely reported chromosome numbers in taxa of subtribe Genet. Mol. Biol. 23: 997–10. Artemisiinae (Anthemideae, Asteraceae) from Mongolia. Bot. J. Pagliarini, M. S., Risso-Pascotto, C., de Souza-Kaneshima, A. M. and Linn. Soc. 150: 203–210. do Valle, C. B. 2008. Analysis of meiotic behavior in selecting Gaulden, M. E. 1987. Hypothesis: some mutagens directly alter spe- potential genitors among diploid and artificially induced tetra- cific chromosomal proteins (DNA topoisomerase II and periph- ploid accessions of Brachiaria ruziziensis (Poaceae). Euphytica eral proteins) to produce chromosome stickiness, which causes 164: 181–187. chromosome aberrations. Mutagenesis 2: 357–365. Panero, J. L. and Funk, V. A. 2002. Toward a phylogenetic subfamilial Ge, C. J., Li, Y. K., Wan, P. and Hsu, P. S. 1989. Chromosome num- classification for the Compositae (Asteraceae). Proc. Biol. Soc. bers of 31 medicinal plants from Shandong Province. In: Hong, Wash. 115: 909–922. D. (ed.). Plant Chromosome Research 1987. Organizing commit- Probatova, N. S., Kazanovsky, S. G., Rudyka, E. G., Barkalov, V. Y., tee Sino-Japanese symposium on plant chromosomes, Beijing. Seledets, V. P. and Nechaev, V. A. 2011. In: Marhold, K. (ed.). pp. 267–272. IAPT/IOPB chromosome data 12. Taxon 60: 1790–1794. Ghaffari, S. M. 2006. Occurrence of diploid and polyploid micro- Shetty, B. V. 1967. In: Löve, Á. (ed.). IOPB chromosome number re- spores in Sorghum bicolor (Poaceae) is the result of cytomixis. ports XIV. Taxon 16: 552–571. Afr. J. Biotechnol. 5: 1450–1453. Singh, V., Gupta, R. C. and Kaur, K. 2016. In: Marhold, K. and Ghanima, A. M. and Talaat, A. A. 2003. Cytomixis and its possible Kučera, J. (eds.). IAPT/IOPB chromosome data 21. Taxon 65: evolutionary role in a Kuwait population of Diplotaxis harra 673–676. (Boraginaceae). Bot. J. Linn. Soc. 143: 169–175. Singh, V., Gupta, R. C., Malik, R. A., Pradhan, S. K., Goyal, H. and Gupta, H., Gupta, R. C., Kumar, R. and Singhal, V. K. 2017. A profile Kaur, K. 2017. Cytogenetic variations in eight species of Sau- of chromosome counts, male meiosis and pollen fertility in 45 ssurea D.C. (Asteraceae, Cardueae) from Northwest Himalaya. species of Asteraceae from Parvati valley in Kullu district, Him- Plant Biosyst. 2017: 1302018. achal Pradesh. Caryologia 71: 1–13. Utsunomiya, K. S., Bione, N. C. P. and Pagliarini, M. S. 2002. How Gupta, R. C. and Gill, B. S. 1983. Cytology of family Compositae many different kinds of meiotic abnormalities could be found in of Punjab plains. Proc. Indian Natl. Sci. Acad. B Biol. Sci. 49: a unique endogamous maize plant? Cytologia 67: 169–176. 359–370. Utsunomiya, K. S., Pagliarini, M. S. and do Valle, C. B. 2005. Micro- Gupta, R. C., Gill, B. S. and Garg, R. K. 1989. Chromosomal con- sporogenesis in tetraploid accessions of Brachiaria nigropedata spectus of western Himalayan Compositae. Aspects Plant Sci. (Ficalho-Hiern) Stapf (Gramineae). Biocell 29: 295–300. 11: 427–437. Yu, C. H., Guo, G. Q., Nie, X. W. and Zheng, G. C. 2004. Cytochemi- Gupta, R. C., Goyal, H., Singh, V. and Goel, R. K. 2014. Meiotic stud- cal localization of pectinase activity in pollen mother cells of to- ies in some species of tribe Cichorieae (Asteraceae) from western bacco during meiotic prophase I and its relation to the formation Himalayas. Sci. World J. 2014: 673456. of secondary plasmodesmata and cytomictic channels. Acta Bot. Hajra, P. K., Rao, R. R., Singh, D. K. and Uniyal, B. P. 1995. Flora of Sin. 46: 1443–1453. India, Vol. XII and XIII. Botanical Survey of India, Calcutta. Zheng, G. C., Yang, Q. and Zheng, Y. 1987. The relationship between Haroun, S. A., Al Shehri, A. M. and Al Wadie, H. M. 2004. Cytomixis cytomixis, chromosome mutation and karyotype evolution in in the microsporogensis of Vicia faba L. (Fabaceae). Cytologia lily. Caryologia 40: 243–259. 69: 7–11.