Cytology of Some Grasses from Haryana and Shiwalik Hills

Akshita Dhaliwal1, Navjot Kaur2* and Raghbir Chand Gupta2

1 Department of Agriculture, Khalsa College, Patiala 147001, India 2 Department of Botany, Punjabi University, Patiala 147002, India

Received July 9, 2017; accepted July 21, 2017

Summary The present paper deals with the male meiosis of 17 species covered under 11 genera of the family Poaceae. Three species: Cynodon dactylon (n=8), Sporobolus diander (n=9) and S. helvolus (n=9) are the new cytotypes from world. Further, ﬁve species i.e., Avena fatua, A. sativa, C. dactylon, Lolium temulentum var. temulentum and Phalaris minor show anomalous meiotic behavior with the presence of cytomixis, chromatin bridges, laggards, secondary associations and interbivalent connections which lead to low pollen viability.

Key words Male meiosis, Poaceae, Meiotic abnormality, Haryana, Shiwalik.

The present study is in continuation of our previous Leica Qwin and a Nikon 80i Eclipse microscope. Pol- work on the cytology of grasses of Haryana and adjoin- len fertility was estimated by their stainability in 1% ing Shiwalik hills (Gupta et al. 2017). Poaceae is the glycero-acetocarmine. Well stained pollen grains were largest family of Monocots. It includes 700 genera and considered as fertile and shriveled or with unstained 11000 species from world (Chen et al. 2006), of which nuclei as sterile. Voucher specimens are deposited to 250 genera and 1275 species have been reported from Herbarium, Department of Botany, Punjabi University, India (Bor 1960). Family has immense economic impor- Patiala (PUN). tance and also exhibits lot of cytological variations due to hybridization, chromosome repatterning, intraspecific Results and discussion and interspecific polyploidy and aneuploidy. The research on the cytology of monocots including Presently, 17 species belonging to 11 genera of Poace- grasses had been undertaken by the various workers of ae have been worked out cytologically. The information the Department of Botany, Punjabi University, Patiala regarding the meiotic chromosome numbers (n), locality (see Gupta et al. 2014, Kaur and Gupta 2016a, 2016b, with accession numbers (PUN), pollen fertility, ploidy Gupta et al. 2017). The present study covers the cytol- level, previous chromosome reports from India and ogy of 17 species falling into 11 genera of the Poaceae. remarks on meiotic behavior are given in Table 1. The The chromosomal data is compiled and analyzed on results on species with new meiotic chromosome report population basis which could be further used for cyto- and those showing abnormal meiosis only been further taxonomic and phylogenetic research. It is attempt on discussed in detail. cytological investigation of grasses from Haryana and In Avena fatua L., during the present study, hexaploid Shiwalik hills. The chromosomal data is compiled and cytotype with abnormal meiosis has been observed. It analyzed on population basis which could be further shows 21 bivalents at diakinesis, 21 : 21 chromosomes used for cytotaxonomic and phylogenetic research. distribution at anaphase I (A-I) (Figs. 2 and 3). Various meiotic anomalies such as un-oriented bivalent at meta- Materials and methods phase I (M-I) (Fig. 4); cytomixis at prophase-I and M-I (Figs. 5 and 6); chromatin bridges at A-I (Fig. 7) and During the present meiotic analysis, young inflores- laggards at early and late telophase I (T-I) (Figs. 8 and cences were collected from different localities of Hary- 9) have been observed. The frequency of 0–5 unoriented ana and Shiwalik hills (Table 1), and fixed in Carnoy’s bivalents at M-I varies from 5.6–24.1% PMCs; cyto- fixative (alcohol : chloroform : acetic acid in 6 : 3 : 1 v/v) mixis 21.7–31.5% PMCs; chromatin bridges 5.6–45.9% for 24 h. After that these were transferred to 70% alcohol PMCs and laggards 10.7–16.6% PMCs (Table 2). These for further preservation. Meiotic studies were carried meiotic abnormalities lead to the low pollen fertility and out by preparing smears of pollen mother cells (PMCs) heterogenous sized pollen grains (Fig. 10). The species in 1% acetocarmine. Photomicrographs of chromo- confirms the previous reports from India and outside. some were obtained from freshly prepared slides using Besides this hexaploid cytotype, the species is also reported with diploid and tetraploid cytotypes (Table 1). * Corresponding author, e-mail: [email protected] A. sativa L.: Presently, the species shows 21 bivalents DOI: 10.1508/cytologia.83.23 at diakinesis (Fig. 11). The meiotic behavior is found to 24 A. Dhaliwal et al. Cytologia 83(1)

Table 1. Information with voucher data, chromosome number (n), pollen fertility, ploidy level, previous chromosome reports and meiotic behavior of the species studied presently.

Name of Meiotic Previous reports Locality with altitude, Ploidy Pollen the species chr. no. Other Meiotic course sea level (m) level fertility India (PUN) (n) countries

Phragmites karka (Retz.) Trin. ex Steud. P-1 50191 Indri, Karnal 252 18 (Fig. 1) 4x 92.8 36, 38, 48 24: Bir and Sahni 1987, 36: N P-2 50192 HAU, Hissar 216 18 4x 92.6 Ramanathan 1950, 48: Christo- N P-3 50193 Shahpur, Ambala 272 18 4x 98 pher and Abraham 1971, Kalia N 1978, Mehra 1982 Avena fatua L. P-1 49636 Shazadpur, Ambala 272 21 (Figs. 6x 34.5 42, 28 14, 28: Mehra and Remanandan Unoriented bivalents, 2–10) 1973a, 28: Singhal et al. 2014, cytomixis, chromatin P-2 49637 Pipli, Kurukshetra 350 21 6x 28.7 42: Mehra et al. 1968, Mehra bridges, laggards, P-3 49640 Kalka, Shiwaliks 624 21 6x 57.03 1982, 42+: Bir and Sahni 1987 heterogenous sized P-4 49645 Jabli, Shiwaliks 900 21 6x 42.4 bivalents P-5 49651 Dharampur, Solan 1350 21 6x 15.7 A. sativa L. P-1 49605 Pipli, Kurukshetra 350 21 (Figs. 6x 35.7 20, 21, 42: Mehra et al. 1968, Mehra 1982, Cytomixis, unoriented 11–14) 4.-42, 41, Koul and Gohil 1987, Koul et al. bivalents, multiple P-2 49607 Kalka, Shiwaliks 624 21 6x 49.5 42, 48, 63 2000, 42+: Dherawattana and chromatin bridges P-3 49628 Jabli, Shiwaliks 900 21 6x 42.3 Sadananga 1973 Bromus unioloides Kunth (=B. catharticus Vahl.) P-1 50638 Chambaghat, Solan 1350 7 (Fig. 15) 2x 92.0 28, 42, 14: Sharma and Sharma 1979, N 42+2B, Gupta et al. 2014, 42: Mehra 56 and Sharma 1975, Mehra 1982, Singhal et al. 2014 Chloris barbata Sw. P-1 50576 Morni, Panchkula 365 10 (Fig. 16) 2x 93.4 20, 40, 20: Shanthamma & Narayan 1976, N c50 40: Christopher and Abraham 1974, Sharma et al. 1978 C. virgata Sw. P-1 50725 Mullana, Ambala 272 10 (Fig. 17) 2x 97.6 14, 20, 26, 20: Singh and Godward 1960, N P-2 50278 Sanauli, Panipat 218 10 2x 89.0 40 Malik and Tripathi 1970, Gupta N P-3 50282 Devsar, Bhiwani 224 10 2x 97.3 1971, Rao and Mwasumbi 1981, N Sahni and Bir 1985, Bir et al. 1987, 30: Krishnaswamy 1941 Cynodon dactylon (L.) Pers. Cyt-A (n=8) 18, 30, 36, 18: Gupta and Srivastava 1970, P-1 50475 Dharampur, Solan 1350 8 (Fig. 18) 2x 82.9 36+1-2B, 18, 18+0-3B, 36: Tripathi et al. Cyt-B (n=9) 40 1977, 18, 18+0-3B, 36, 54: Malik P-2 50476 Barara, Ambala 272 9 (Fig. 19) 2x 92.9 1966, 18+1-3B, 36, 54: Malik N P-3 50579 Kalanaur, Yamunanagar 255 9 2x 95.6 and Tripathi 1968, 27: Gupta and N Cyt-C (n=18) Srivastava 1970, Christopher P-4 50477 Cheeka, Kaithal 219 18 (Figs. 4x 43.5 and Abraham 1974, Kuumar and Laggards and 20–22) Sachdeva 1988, Sinha et al. 1990, chromatin bridges P-5 50580 Pinjour, Panchkula 365 18 4x 27.9 27, 36: Sachdeva and Bhatia 1980, 36: Malik 1966, Malik and Tripathi 1968, Mehra et al. 1968, Mehra and Sharma 1975, Kalia 1978, Sharma et al. 1978, Mehra 1982, Bir et al.1987, Koul and Gohil 1987, Sinha et al. 1990, Singhal et al. 2014, 36+0-2B: Tripathi et al. 1977, 36, 40: Mehra et al. 1968, 30, 36, 40: Malik and Tripathi 1968 Lolium temulentum Linn. var. temulentum P-1 50163 Morni, Panchkula, 365 7 (Fig. 2x 59.2 14, 14: Mehra and Sharma 1975, Chromatin transfer, 23–26) 14+0-2B, Parkash 1979, Mehra 1982, laggards, chromatin P-2 50164 Kalka, Panchkula 624 7 2x 44.7 14+1s, 28, Mehra and Remanandan 1973a, bridges 56, 70 14, 28+0-3B: Bir and Sahni 1987, 28+2B: Mehra and Sunder 1969 2018 Cytology of Some Grasses from Haryana and Shiwalik Hills 25

Table 1. Continued.

Name of Meiotic Previous reports Locality with altitude, Ploidy Pollen the species chr. no. Other Meiotic course sea level (m) level fertility India (PUN) (n) countries

Poa annua L. P-1 50165 Dharampur, Solan 1350 14 (Fig. 27) 4x 47.6 14, 14: Mehra and Sunder 1969, 28: Quadrivalents P-2 50166 Jabli, Shiwaliks 900 14 4x 59.6 21+0-1B, Christopher and Abraham 1971, P-3 50168 Mullana, Ambala 272 14 4x 100.0 24, 26, 28, Mehra and Sharma 1975, Parkash P-4 50171 Guhla, Kaithal 219 14 4x 85.2 42, 52 1979, Mehra 1982, Bir and Sahni 1987, Koul and Gohil 1987, Bir et al. 1987, Kaur et al. 2010a, Gupta et al. 2014, 28+1B: Mehra et al. 1968, 28+0-1B:Bir and Sahni 1987, 28+0-3B: Kaur et al. 2011b, 52: Mehra and Remanandan 1973a Perotis indica (L.) Kuntze P-1 50561 Mirpur, Rewari 241 20 (Fig. 28) 4x 82.4 20 20: Mehra et al. 1968, Malik and N P-2 50563 Jagadhri, Yamunanagar 255 20 4x 87.6 Mary 1970, 40: Christopher and N P-3 Namaul, Mahendragarh 261 20 4x 87.3 Abraham 1974 N Phalaris minor Retz. P-1 49677 Pipli, Kurukshetra 350 14 4x 32.5 ̶ 28: Bir and Sahni 1987 Multivalents, interbiva- (Fig. 29–37) lent connection, unoriented bivalents, chroma- P-2 49673 Jabli, Shiwaliks 900 14 4x 45.2 tin transfer, secondary P-3 49687 Shahzadpur, Ambala 272 14 4x 46.3 associations, laggards, P-4 49707 Mandi Dabwali, Sirsa 204 14 4x 27.6 chromatin bridges Sporobolus coromandelianus (Retz.) Kunth P-1 50763 Mullana, Ambala 272 18 (Fig. 38) 4x 92.9 36 24: Christopher and Abraham N P-2 50765 Indri, Karnal 252 18 4x 98.3 1974, 24, 36: Bir et al. 1988, 36: N Bir and Sahni 1985, Gupta et al. 2014 S. diander (Retz.) P. Beauv. Cytotype A (n=9) 24: Christopher and Abraham P-1 50480 Devsar, Bhiwani, 224 9 (Fig. 39) 2x 17.4 24, 36 1974, Mehra and Sharma 1975, P-2 50481 Sanauli, Panipat, 218 9 2x 29.9 Mehra 1982, 24, 36: Bir and Cytotype B (n=18) Sahni 1987, Bir et al. 1988, 36: P-1 50482 KUK, Kurukshetra 350 18 (Fig. 40) 4x 94.8 Gupta et al. 2014, 54: Mehra N P-2 50773 Agroha, Hissar 216 18 4x 93.2 et al. 1968 N S. helvolus (Trin.) T. Durand and Schinz P-1 50864 Lalru, Ambala 272 9 (Fig. 41) 2x 85.7 16, 20, 28, 30, 32: Bir et al. 1987, P-2 50870 Radaur, Yamunanagar 255 9 2x 90.0 36: Bir and Sahni 1987, Bir et al. P-3 50871 Gohana, Sonipat 219 9 2x 91.7 1988 S. indicus (L.) R. Br. P-1 50769 Parwanoo, Solan 1350 18 (Fig. 42) 4x 92.4 18, 24, 36, 18: Mehra et al. 1968, 30: N P-2 50770 Barog, Solan 1350 18 4x 84.9 40–45, Christopher and Abraham 1974, N ≈48, ≈54 36: Bir and Sahni 1987, Bir et al. 1988, 48: Christopher and Abraham 1974 S. marginatus Hochst. ex A. Rich. P-1 50771 HAU, Hissar 216 18 (Fig. 43) 4x 30.2 18, 20 18+: Bir and Sahni 1985, 18, 36: N Bir et al. 1988, 36: Mehra et al. 1968, Bir et al. 1988 Tragus roxburghii Panigrahi P-1 50616 Indri, Karnal 252 10 (Fig. 44) 2x 100.0 20 20: Sahni and Bir 1985 N P-2 50619 Jyotisar, Kurukshetra 350 10 2x 100.0 N P-3 50621 Ballabgarh, Faridabad 197 10 2x 100.0 N

N stands for normal meiosis. be abnormal with the presence of cytomixis (15.5–17.3% Cynodon dactylon (L.) Pers.: During the present PMCs); unoriented bivalents (14.3–33.0% PMCs) and study, three cytological races with n=8, 9 and 18 have multiple chromatin bridges (4.49–21.6% PMCs) (Figs. been detected from the study area (Figs. 18–20). The 12–14). Pollen fertility is found to be very low. From diploid cytotype with 2n=16 is a new chromosome re- India, only hexaploid cytotype is reported, whereas from port for the species with normal meiosis, whereas the outside India besides this cytotype, other polyploid and other cytotypes with 2n=18 and 36 conﬁrms the previ- aneuploid cytotypes area also reported (Table 1). ous reports (Table 1). Meiotic course of the tetraploid 26 A. Dhaliwal et al. Cytologia 83(1)

Table 2. Data on meiotic abnormalities in the studied species.

Interbivalent Secondary Species name Cytomixis Unoriented bivalents Chromatin bridges Laggards connections associations

1 Avena fatua P-1 13/49 (26.5) 4/71 (5.6) 4/55 (7.2) 8/69 (11.6) ̶ ̶ P-2 7/29 (24.1) 9/52 (17.3) 7/48 (14.5) 12/86 (13.9) ̶ ̶ P-3 15/52 (28.8) 14/58 (24.1) 5/88 (5.6) 7/42 (16.6) ̶ ̶ P-4 12/38 (31.5) 13/79 (16.4) 7/44 (15.9) 4/32 (12.5) ̶ ̶ P-5 10/46 (21.7) 10/55 (18.1) 15/96 (15.6) 7/65 (10.7) ̶ ̶ 2 Avena sativa P-1 14/88 (15.9) 12/84 (14.3) 4/89 (4.49) ̶ ̶ ̶ P-2 7/45 (15.5) 27/82 (33.0) 7/78 (8.97) ̶ ̶ ̶ P-3 9/52 (17.3) 18/70 (25.7) 21/97 (21.6) ̶ ̶ ̶ 3 Cynodon dactylon P-4 ̶ ̶ 7/67 (10.4) 11/85 (12.9) ̶ ̶ P-5 ̶ ̶ 8/82 (9.75) 9/92 (9.78) ̶ ̶ 4 Lolium temulentum var. temulentum P-1 13/94 (13.8) ̶ 5/61 (8.19) 8/54 (14.8) ̶ ̶ P-2 15/87 (17.2) ̶ 9/72 (12.5) 3/56 (5.35) ̶ ̶ 5 Phalaris minor P-1 11/57 (19.3) 14/99 (14.1) 4/50 (8.0) 21/77 (27.3) 5/74 (6.7) 12/50 (24.0) P-2 8/45 (17.7) 21/77 (27.3) 5/98 (5.1) 12/47 (25.5) 4/71 (5.63) 11/42 (26.2) P-3 8/55 (14.5) 17/85 (20.0) 7/69 (10.1) 11/65 (16.9) 7/92 (7.6) 16/54 (29.6) P-4 9/62 (14.5) 7/65 (10.8) 6/74 (8.1) 13/72 (18.1) 9/87 (10.3) 16/68 (23.5)

Figures outside parentheses denote number of abnormal cells in numerator and total number of observed cells in denominator; figures inside parentheses are their percentage. cytotype is found to be abnormal with the presence of diploid cytotype (n=9) is a new euploid cytotype for the laggards (10.4% PMCs in P-4, 9.75% PMCs in P-5) and species from world. Already, the species is known with chromatin bridges (12.9% PMCs in P-4, 9.78% PMCs in 2n=24, 36 and 54 from India. P-5) (Table 2; Figs. 21 and 22). The species shows well S. helvolus (Trin.) T. Durand & Schinz.: Presently, a developed polyploid (2x, 3x, 4x and 6x) and aneuploid large number of PMCs at M-I confirm the presence of cytotypes. n=9 (Fig. 41). Meiosis is found to be normal with high Lolium temulentum Linn. var. temulentum: Presently, pollen fertility (Table 1). The present chromosome re- both the studied populations show diploid chromosome port of diploid cytotype is a new euploid cytotype for the count with seven bivalents at M-I (Fig. 23). Meiosis is species from world. Already, the species is reported with quite abnormal due to the presence of cytomixis (13.8– many other cytotypes (2n=16, 20, 28, 30, 32 and 36) 17.2% PMCs); chromatin bridges (8.19–12.5% PMCs) Different meiotic chromosome numbers (n=7, 8, 9, and laggards (14.8–5.35% PMCs). The pollen fertility 10, 14, 18, 20 and 21) have been observed during the in the two populations is 59.2 and 44.7% PMCs (Table present study. Ploidy level varies from 2x to 6x, out of 2, Figs. 24–26). Besides, the diploid cytotype, tetraploid which 4x is the most common recorded in nine species, cytotype is also reported. followed by 2x in eight species based on basic chromo- Phalaris minor Retz.: During the present study, tetra- some numbers x=7, 8, 9 and 10. Out of the 17 studied ploid chromosome count with 14 bivalents at M-I (Fig. species, five species (one diploid and four polyploids) 29) has been observed. Further, the meiotic course in all showed abnormal meiotic behavior which includes cy- the populations was found to be abnormal. The number tomixis, secondary chromosomal associations, inter- of quadrivalents per PMC varies from 0–3 (Fig. 30). The bivalent connections, chromatin bridges and laggards, overall frequency of the quadrivalents comes out to be resulting into the pollen sterility and heterogenous sized 0.54 per PMC. Various meiotic abnormalities such as pollen grains. interbivalent connections (Fig. 31); unoriented bivalents Cytomixis is the phenomena of chromatin/chro- (10.8–27.3% PMCs); cytomixis (14.5–19.3% PMCs); mosome transfer between the neighboring meiocytes secondary associations (23.5–29.6% PMCs); laggards through cytoplasmic channels and or intercellular bridg- (16.9–27.3% PMCs) and chromatin bridges (5.1–10.1% es (Bione et al. 2000, Li et al. 2009). It is reported in PMCs) are observed (Table 2). Microsporogenesis is number of angiosperm species (Malallah and Attia found to be normal. Already, the species is known with 2003). According to Haroun (1995), it is more prevalent the same chromosome number (Table 1). in apomicts and aneuploids or even among polyploids Sporobolus diander (Retz.) P. Beauv.: During the (Semyarkhina and Kuptsou 1974). Some of the workers present study, two cytotypes with n=9 and n=18 have (Srivastav and Raina 1980, Zheng et al. 1987) consid- been observed (Figs. 39 and 40). The present report of ered it as one of the mechanisms for evolution. Earlier 2018 Cytology of Some Grasses from Haryana and Shiwalik Hills 27

Figs. 1–26. Meiotic chromosomes, abnormalities and pollen grains. 1. In Phragmites karka. PMC showing 18II at diakinesis. 2. In Avena fatua. PMC showing 21II at diakinesis. 3. PMC with 21 : 21 chromosomes at A-I. 4. Unoriented bivalents (arrowed). 5,6. Cytomixis at diakinesis and M-I (arrowed). 7. Chromatin bridges (arrowed). 8,9. Laggards. 10. Het-

erogenous sized pollen grains. 11. In A. sativa. PMC showing 21II at diakinesis. 12. Cytomixis (arrowed). 13. Un- oriented bivalents (arrowed). 14. Multiple chromatin bridges (arrowed). 15. In Bromus unioloides. PMC showing 7II at diakinesis. 16. In Chloris barbata. PMC showing 10II at M-I. 17. In C. virgata. 17 : 17 chromosomes at A-I. 18. In Cynodon dactylon. (Cytotype-A) PMC showing 8II at diakinesis. 19. (Cytotype-B) PMC showing 9 : 9 chromosomes at A-I. 20. (Cytotype-C) PMC showing 18II at M-I. 21. Laggards (arrowed). 22. Chromatin bridges (arrowed). 23. In Lolium temulentum var. temulentum. PMC showing 7II at M-I. 24. Chromatin transfer (arrowed). 25. Laggards (arrowed). 26. Chromatin bridges (arrowed). Scale bar=10 µm. 28 A. Dhaliwal et al. Cytologia 83(1)

Figs. 27–44. 27. In Poa annua. PMC showing 1IV+12II at M-I (arrowed). 28. In Perotis indica. PMC showing 20II at diakinensis. 29. In Phalaris minor. PMC showing 14II at M-I. 30. PMC showing 1IV+12II at M-I (arrowed). 31. Interbivalent connection (arrowed). 32. Unoriented bivalents (arrowed). 33. Chromatin transfer (arrowed). 34. Secondary associations (arrowed). 35. Laggards (arrowed). 36. Chromatin bridges (arrowed). 37. Heterogenous sized fertile

and sterile pollen grains. 38. In Sporobolus coromandelianus. PMC showing 18II at diakinesis. 39. In S. diander. (Cytotype-A) PMC showing 9 : 9 (n=9) chromosomes at A-I. 40. (Cytotype-B) PMC showing 18II at diakinesis. 41. S. helvolus. PMC showing 9II at M-I. 42. S. indicus. PMC showing 18II at diakinesis. 43. S. marginatus. PMC showing 18II at diakinesis. 44. Tragus roxburghii. PMC showing 10II at diakinesis. Scale bar=10 µm.

cytomixis has been reported in some of the grass species reported in Avena species (Baptista-Giacomelli et al. (Basavaiah and Murthy 1987, Koul 1990). Presently, 2000, Sheidai et al. 2003). However, lack of correlation cytomixis is reported in four species: A. fatua (n=21), between meiotic abnormalities and pollen viability has A. sativa (n=21), L. temulentum var. temulentum (n=7) been reported in Brachiaria brizantha (Mendes-Bonato and P. minor (n=14). Chromatin bridges and laggards et al. 2009). These meiotic abnormalities gradually lead have been reported in A. fatua, A. sativa, C. dactylon, L. to pollen sterility. temulentum var. temulentum and P. minor. The presence of chromatin bridges with accompanying fragments is Acknowledgements an indirect evidence for the existence of chromosomal inversions (Brown 1972). Further, unoriented bivalents The authors are thankful to DBT-IPLS project have also been observed. These may be formed due to (Project no. BT/PR-4548/INF/22/146/2012) sanctioned the impaired attachment of kinetochore or late terminal- to Punjabi University, Patiala for ﬁnancial assistance. ization of chiasmata (Pagliarini 2000). Secondary asso- The authors are also thankful to the Head of Department ciations are the loose associations of chromosomes with- of Botany, Punjabi University, Patiala for providing us out chiasma formation. Presently, these are observed necessary laboratory facilities. only in P. minor. This type of meiotic abnormality was 2018 Cytology of Some Grasses from Haryana and Shiwalik Hills 29

References Koul, K. K. and Gohil, R. N. 1987. SOCGI plant chromosome number reports–IV. J. Cytol. Genet. 22: 161–162. Koul, K. K. 1990. Cytomixis in pollen mother cells of Alopecurus Baptista-Giacomelli, F. R., Pagliarini, M. S. and Almeida, J. L. 2000. arundinaceus Poir. Cytologia 55: 169–173. Meiotic behavior in several Brazilian oat cultivars (Avena sativa Koul, K. K., Nagpal, R. and Sharma, A. 2000. Chromosome behav- L.). Cytologia 65: 371–378. ior in the male and female sex mother cells of wheat (Triticum Basavaiah, D. and Murthy, T. C. S. 1987. Cytomixis in pollen mother aestivum L.), oat (Avena sativa L.) and pearl millet (Pennisetum cells of Urochloa panicoides P. Beauv. (Poaceae). Cytologia 52: americanum (L.) Leeke). Caryologia 53: 175–183. 69–74. Krishnaswamy, N. 1941. Untersuchungen zur Cytologie und System- Bione, N. C. P., Pagliarini, M. S. and De Toledo, J. F. F. 2000. Meiotic atic der Gramineen. Beih. Bot. Zentralbl. 60: 1–56. behavior of several Brazilian soybean varieties. Genet. Mol. Biol. Kumar, P. and Sachdeva, S. K. 1988. Karyomorphological observa- 23: 623–631. tions and biochemical variability in triploids of Cynodon dac- Bir, S. S. and Sahni, M. 1985. Cytological investigations on some tylon (L.) Pers. complex. Proc. Indian Sci. Congr. Assoc. 75: grasses from Punjab Plain, North India. Proc. Indian Nat. Sci. 210–211. Acad. Part B: Biol. Sci. 5: 609–626. Li, X. F., Song, Z. Q., Feng, D. S. and Wang, H. G. 2009. Cytomixis in Bir, S. S. and Sahni, M. 1987. Chromosomal and morphological varia- Thinopyrum intermedium, Thinopyrum ponticum and its hybrids tions in grasses of Punjab. J. Cytol. Genet. 22: 12–22. with wheat. Cereal Res. Commun. 37: 353–361. Bir, S. S., Sahni, M., Kanta, S. and Gill, B. S. 1987. Cytological in- Malallah, G. A. and Attia, T. A. 2003. Cytomixis and its possible evo- vestigations on some grasses from Punjab plain. North India. J. lutionary role in a Kuwaiti population of Diplotaxis harra (Bras- Cytol. Genet. 22: 23–47. sicaceae). Bot. J. Linn. Soc. 143: 169–175. Bir, S. S., Sahni, M. and Singh, C. P. 1988. Cytology of genus Spo- Malik, C. P. and Mary, T. N. 1970. In: Löve, A. (ed.). IOBP chromo- robolus R. Br. from north India (Punjab Plain). Cytologia 53: some number reports XXVII. Taxon 19: 437–442. 53–57. Malik, C. P. and Tripathi, R. C. 1968. Cytological evolution within the Bor, N. L. 1960. Grasses of Burma, Ceylon, India. and Pakistan. Per- Cynodon dactylon complex. Biol. Zent. 87: 625–627. gamon Press, Oxford. Malik, C. P. 1966. Polyploidy in Cynodon dactylon (Linn.). Pers. Brown, W. V. 1972. Inversions. In: Brown, W. V. (ed.). Textbook of Chromosome. Inf. Ser. 7: 6–7. Cytogenetics. Mosby Company, Saint Louis. p. 196. Mehra, P. N. and Remanandan, P. 1973. Cytological investigations on Chen, S. et al. 2006. Poaceae. In: Flora of China, Vol. 22 (Poaceae). W. Himalayan Pooideae. Cytologia 38: 237–258. Wu, Z. Y., Raven, P. H. and Hong, D. Y. (eds.). Science Press and Mehra, P. N. and Sharma, M. L. 1975. Cytological studies in some Missouri Botanical Garden Press, Beijing and St. Louis. pp. 1–2. Central and Eastern Himalayan grasses. IV. The Arundinelleae, Christopher, J. and Abraham, A. 1971. Studies on the cytology and Eragrosteae, Isachneae, Chlorideae, Sporoboleae, Meliceae, Sti- phylogeny of South Indian grasses I. Subfamilies Bambusoideae, peae, Arundineae and Garnotieae. Cytologia 40: 453–462. Oryzoideae. Arundinoideae and Festucoideae. Cytologia 36: Mehra, P. N. and Sunder, S. 1969. Cytological studies in the North 579–594. Indian grasses. Part II. Res. Bull. Panjab Uni. Sci. 20: 503–539. Christopher, J. and Abraham, A. 1974. Studies on the cytology and Mehra, P. N., Khosla, P. K., Kohli, B. L. and Koonar, J. S. 1968. Cyto- phylogeny of South Indian grasses. II. Subfamily Eragrostoideae. logical studies in North Indian grasses. I. Res. Bull. Panjab Uni. Cytologia 39: 561–571. Sci. 19: 157–230. Dherawattana, A. and Sadanaga, K. 1973. Cytogenetics of a crown Mehra, P. N. 1982. Cytology of East Indian Grasses. New Delhi. rust-resistant hexaploid oat with 42+2 fragment chromosomes. Mendes-Bonato, A. B., Ferrari Felismino, M., Souza Kaneshima, A. Crop Sci. 13: 591–594. M., Pessim, C., Calisto, V., Pagliarini, M. S. and Borges do Valle, Gupta, P. K. and Srivastava, A. K. 1970. Natural triploidy in Cynodon C. 2009. Abnormal meiosis in tetraploid genotypes of Brachiaria dactylon (L.). Pers. Caryologia 23: 29–35. brizantha (Poaceae) induced by colchicine: its implications for Gupta, P. K. 1971. Cytological investigations in some North Indian breeding. J. Appl. Genet. 50: 83–87. grasses. Genet. Iber. 23: 183–198. Pagliarini, M. S. 2000. Meiotic behavior of economically important Gupta, R. C., Chauhan, H. S., Saggoo, M. I. S. and Kaur, N. 2014. plant species: the relationship between fertility and male sterility. Cytomorphology of some grasses (Poaceae) from Lahaul-Spiti Genet. Mol. Res. 23: 997–1002. (Himachal Pradesh), India. Biolife 2: 1234–1247. Parkash, O. 1979. Cytological investigations in some grasses of north- Gupta, R. C., Gupta, A. and Kaur, N. 2017. Meiotic studies in some eastern India (Festucoids, Ischaemum L., Eulalia Kunth and Sac- members of tribe Andropogoneae (Poaceae) from semi desert charum spontaneum L.). Ph.D. Thesis, Chandigarh. area of North India. Cytologia 82: 105–113. Ramanathan, K. 1950. Addendum to list of chromosomes numbers in Haroun, S. A. 1995. Cytomixis in pollen mother cells of Polygonum economic plants. Curr. Sci. 19: 155. tomentosum Schrank. Cytologia 60: 257–260. Rao, P. N. and Mwasumbi, L. B. 1981. In: Löve, A. (ed.). IOBP chro- Kalia, V. 1978. Cytological investigations in some grasses of north- mosome number reports LXXII. Taxon 30: 701. eastern India. Tribes: Andropogoneae, Arundineae, Oryzeae, Sachdeva, S. K. and Bhatia, M. S. 1980. Chemotaxonomic studies Arundinelleae, Chlorideae, Eragrosteae and Sporoboleae. Dis- in Cynodon dactylon (L.) Pers. complex II. Flavonoid patterns sertation, Panjab University, Chandigarh. and ascorbic acid content. Proc. Indian Acad. Sci. Plant Sci. 89: Kaur, H., Gupta, A., Kumari, S. and Gupta, R. C. 2010a. Meiotic stud- 193–196. ies in Poa annua L. from different altitudinal ranges of North Sahni, M. and Bir, S. S. 1985. SOCGI plant chromosome number re- India. Cytologia 75: 313–318. ports–III. J. Cytol. Genet. 20: 205–206. Kaur, H., Gupta, R. C. and Kumari, S. 2011. In: Marhold, K. et. al. Semyarkhina, S. Y. A. and Kuptsou, M. S. 1974. Cytomixis in various (eds.). IAPT/IOPB chromosome data 12. Taxon 60: 1789. forms of Sugarbeet. Vests. I. ANBSSE Ser. Biyal. 4: 43–47. Kaur, N. and Gupta, R. C. 2016a. Cytological study in some members Shanthamma, LI. and Narayan, K. N. 1976. In: Löve, A. (ed.). IOBP of tribe Paniceae (Poaceae) from Rajasthan. Cytologia 81: 13–17. chromosome number reports. Taxon 25: 155–164. Kaur, N. and Gupta, R. C. 2016b. Anomalous meiotic behavior in Sharma, C. B., Behera, B. N. and Dash, S. K. 1978. A cytological different populations of octaploid Sorghum bicolor (L.) Moench study of some grasses from Orissa, an Eastern coastal belt of from Sirohi District (Rajasthan), India. Chromosome Bot. 11: India. Proc. Indian Natl. Sci. Acad. 87 B. Biol. Sci. 11: 355–360. 21–26. 30 A. Dhaliwal et al. Cytologia 83(1)

Sharma, M. L. and Sharma, K. 1979. Cytological studies in the North Srivastav, P. K. and Raina, S. N. 1980. Cytomixis in Clitoria ternan- Indian grasses. Cytologia 44: 861–872. tea L. var. pleniﬂora Fants. Curr. Sci. 49: 824–835. Sheidai, M., Koobaz, P. and Zehzad, B. 2003. Meiotic studies of some Tripathi, R. C., Sachdeva, S. K. and Malik, C. P. 1977. Cytogenetic Avena species and populations in Iran. J. Sci. I. R. Iran. 14: studies and evolutionary patterns in some Cynodon species. Biol 121–131. Zentr. 96: 423–435. Singh, D. N. and Godward, M. B. E. 1960. Cytological studies in the Zheng, G. C., Yang, Q. and Zheng, Y. 1987. The relationship between Gramineae. Heredity 15: 193–197. cytomixis, chromosome mutation and karyotype evolution in Sinha, R. R. P., Bhardwaj, A. K. and Singh, R. K. 1990. SOCGI plant lily. Caryologia 40: 243–259. chromosome number reports IX. J Cytol Genet. 25: 140–143.