Chromosome count, male meiotic behaviour and pollen fertility analysis in Agropyron thomsonii Hook.f. and Elymus nutans Griseb. (Triticeae: Poaceae) from Western Himalaya, India
Present cytological study records existing chromosome number diversity, their male meiotic course and pollen fertility analysis in the two wheatgrass species of tribe Triticeae Dumort. (Poaceae) from Western Himalaya, India. Agropyron thomsonii Hook. f. is an endemic grass of alpine zones of Western Himalaya and Elymus nutans Griseb., a widely distributed grass in sub-alpine to glacial regions of Himalaya. The gametophytic chromosome number count of n=21 (Jadh Ganga Valley, Uttarkashi) is a pioneer count for A. thomsonii. During the male meiotic course of A. thomsonii, 14.04-16.29% and 2.97-4.17% pollen mother cells, respectively at prophase-I and metaphase-I, observed to be involved in phenomenon of cytomixis. Seven accessions of E. nutans collected from Bhagirathi Valley and Jadh Ganga Valley of Uttarkashi district, and Pangi Valley of Chamba district, recorded with gametophytic chromosome number count of n=21 and record of 1B-chromosome in PUN61958 is a new record for the species. In three accessions 5.56-9.41% and 2.5% pollen mother cells at prophase-I and metaphase-I, respectively were also noted with phenomenon of cytomixis. In addition to phenomenon of cytomixis, during meiotic course of both species pollen mother cells also depicted associated meiotic course irregularities viz. non-synchronous disjunction of bivalents, chromatin bridges, laggards, micronuclei in sporads and shrivelled microspores. These species are growing in cold climatic condition habitats. So, cold stress seems to be a preferential inductor for cytomixis and associated meiotic abnormalities in the gametic cells of stamens of A. thomsonii and E. nutans that ultimately leads to reduction in pollen fertility.
Basavaiah, Murthy TCS. 1987. Cytomixis in pollen mother cells of Urochloa panicoides P. Beauv. (Poaceae). Cytologia. 52:69–74.
Bauchan GR, Linkous LCW, Tai W. 1987. Cytomixis in Agropyron cristatum. Genome. 29:765–769.
Bedi YS. 1990. Cytomixis in woody species. In: Proc Indian Nat Sci Acad Plant Sci. 100:23–238.
Bellucci M, Roscini C, Mariani A. 2003. Cytomixis in pollen mother cells of Medicago sativa L. J Heredit. 94:512–516.
Bhat TA, Parveen S, Khan AH. 2006. MMS-induced cytomixis in pollen mother cells of broad bean (Vicia faba L.). Turk J Bot. 30:273–279.
Boldrini KR, Pagliarini MS, do Valle CB. 2006. Cell fusion and cytomixis during microsporogenesis in Brachiaria humidicola (Poaceae). S Af J Bot. 72:478–481.
Bor NL. 1960. Grasses of Burma, Ceylon, India and Pakistan (excluding Bambuseae). London: Pergamon Press.
Chen SL, Zhu GH. 2006. Elymus L. Flora of China (Poaceae). Beijing and St. Louis, MO: Science Press and Missouri Botanical Garden. 22:400–429.
Chen SY, Ma X, Zhang XQ, Chen ZH. 2009. Genetic variation and geographical divergence in Elymus nutans Griseb. (Poaceae: Triticeae) from West China. Bioch Syst Ecol. 37:716–722.
Chen S, Zhang X, Ma X, Huang L. 2013. Assessment of genetic diversity and differentiation of Elymus nutans indigenous to Qinghai-Tibet Plateau using simple sequence repeats markers. Can J Pl Sci. 93:1089–1096.
Consolaro MEL, Pagliarini MS. 1995. Cytomixis in pollen mother cells of Centella asiatica L. Nucleus. 38:80–85.
Cope TA. 1982. Poaceae. In: Nasir E, Ali SI. (Eds.). Flora of Pakistan. No. 143. Pakistan: Department of Botany, University of Karachi.
Cuñado N, Santos JL. 1999. On the diploidization mechanism of the genus Aegilops: Meiotic behaviour of interspecific hybrids. Theor. Appl. Genet. 99:1080–1086.
Cuñado N, Callejas S, García MJ, Fernández A, Santos JL. 1996a. Chromosome pairing in the allotetraploid Aegilops biuncialis and a triploid intergeneric hybrid. Genome 39:664–670.
Cuñado N, Callejas S, García MJ, Fernández A, Santos JL. 1996b. The pattern of zygotene and pachytene pairing in allotetraploid Aegilops species sharing the U genome. Theor. Appl. Genet. 93:1152–1155.
Cuñado N, García MJ, Callejas S, Fernández A, Santos JL. 1996c. The pattern of zygotene and pachytene pairing in allotetraploid Aegilops species sharing the D genome. Theor. Appl. Genet. 93:1175–1179.
Dewey DR. 1984. The genomic system of classification as a guide to intergeneric hybridization with the perennial Triticeae. In: Gustafson JP. (Ed.). Gene manipulation in plant improvement, 16th Stadler Genetics Symposium. Boston: Springer. pp. 209–279.
De M, Sharma AK. 1983. Cytomixis in pollen mother cells of an apomictic ornamental Ervatamia divaricata (Linn.) Alston. Cytologia. 48:201–207.
Dou QW, Chen ZG, Liu YA, Tsujimoto H. 2009. High frequency of karyotype variation revealed by sequential FISH and GISH in plateau perennial grass forage Elymus nutans. Breed Sci. 59:651–656.
Dou Q, Yu F, Li Y, Zhao Y, Liu R. 2017. High molecular karyotype variation revealed in indigenous Elymus nutans in the Qinghai Plateau. Pl Divers. 39:117–122.
Dvorsky M, Klimes L, Dolezal J, Wild J, Dickoré BW. 2018. A Field Guide to the Flora of Ladakh. Praha: Academia.
Gates RR. 1911. Pollen formation in Oenothera gigas. Ann Bot. 25:909–940.
Gohil RN, Koul KK. 1985. In: Bir SS. (ed.). SOCGI plant chromosome number reports. J Cytol Genet. 21:155.
Holm PB. 1986. Chromosome pairing and chiasma formation in allohexaploid wheat, Triticum aestivum, analysed by spreading of meiotic nuclei. Carlsberg Res. Commun. 51:239–294.
Holm PB, Wang X. 1988. The effect of chromosome SB on synapsis and chiasma formation in wheat, Triticum aestivum, cv. Chinese Spring. Carlsberg Res. Commun. 53:191–208.
Hooker JD. 1896. The Flora of British India. London. 7:370.
Jones GH, Brumpton RJ. 1971. Sister and non-sister chromatid U-type exchange in rye meiosis. Chromosoma. 33:115–128.
Karthikeyan S, Jain SK, Nayar MP, Sanjappa M. 1989. Florae Indicae Enumeratio Monocotyledonae. Calcutta: Botanical Survey of India.
Kawano S. 2018. Karyotype and chromosome behavior analyses in three regions of the Indomalayan Realm. Cytologia 83:223–228.
Keshavarzi M, Nasrollahi F, Sheidai M. 2017. Cytogenetic study of the genus Anchusa L. (Boraginaceae) in Iran. Caryologia. 70:57–365.
Khan NA, Singhal VK, Gupta RC. 2018. First record of chromosome count and cytomixis in an endemic species of Clematis ladakhiana Grey-Wilson (Ranunculaceae) from cold deserts of Jammu and Kashmir. Caryologia 71: 233–237.
Körnicke M. 1901. About change of location of cell nuclei. Meeting Reports of Society of Lower Rhine for Nature and Medicine (Bonn). pp. 14–25. (in Germen)
Koul KK. 1990. Cytomixis in pollen mother cells of Alopecurus arundinaceus Poir. Cytologia. 55:169–173.
Kravets EA, Yemets AI, Blume YB. 2017. Cytoskeleton and nucleoskeleton involvement in processes of cytomixis in plants. Cell Biol. Int. [doi:10.1002/cbin. 10842]
Kumar P. 2010. Exploration of cytomorphological diversity in the members of Polypetalae from Lahaul-Spiti and adjoining areas. PhD Thesis. India: Punjabi University Patiala, Punjab.
Kumar P, Singhal VK. 2008. Cytology of Caltha palustris L. (Ranunculaceae) from cold regions of Western Himalayas. Cytologia. 73:137–147.
Kumar P, Singhal VK. 2016. Nucleoli migration coupled with cytomixis. Biologia. 71:651–659.
Kumar P, Singhal VK, Kaur D, Kaur S. 2010. Cytomixis and associated meiotic abnormalities affecting pollen fertility in Clematis orientalis. Biol Plant. 54:181–184.
Kumar P, Singhal VK, Rana PK, Kaur S, Kaur D. 2011. Cytology of Ranunculus laetus Wall. ex Royle from cold desert regions and adjoining hills of North-west Himalayas. Caryologia. 64:25–32.
Kumar P, Rana PK, Singhal VK, Gupta RC. 2014. Cytogeography and phenomenon of cytomixis in Silene vulgaris from cold regions of Northwest Himalayas (India). Plant Syst Evol. 300:831–842.
Kumar P, Singhal VK, Srivastava SK. 2017. First detection of cytomixis and its consequences in Thalictrum cultratum Wall. (Ranunculaceae). Cytol Genet. 51:384–390.
Lattoo SK, Khan S, Bamotra S, Dhar AK. 1990. Cytomixis impairs meiosis and influences reproductive success in Chlorophytum comosum (Thunb.) Jacq. – an additional strategy and possible implications. J Biosci. 31:629–637.
Liu Y. 1985. Studies on the karyotypes of 11 species of Elymus from China. J Wuh Bot Res. 3:325–330.
Liu B, Mo WJ, Zhang D, De Storme N, Geelen D. 2019. Cold influences male reproductive development in plants: A hazard to fertility, but a window for evolution. Plant Cell Physiol. 60:7–18.
Lu BR. 1993. Meiotic studies of Elymus nutans and E. jacquemontii (Poaceae, Triticeae) and their hybrids with Pseudoroegneria spicata and seventeen Elymus species. Plant Syst Evol. 186:193–211.
Lu BR. 1994. Morphological identification of Elymus sibiricus, E. nutans, and E. burchan-buddae, and their genomic relationships. Acta Phytotax Sinic. (China). 32:504–513.
Lu B, Bothmer, RV. 1993. Meiotic analysis of Elymus caucasicus, E. longearistatus, and their interspecific hybrids with twenty-three Elymus species (Triticeae, Poaceae). Plant Syst Evol. 185:35–53.
Lu BR, Yan J, Yang JI. 1990. Cytological observation on Triticeae materials from Xinjiang, Qinghai and Sichuan. Acta Bot. Yunn. (China). 12:57–66.
Malallah GA, Attia TA. 2003. Cytomixis and its possible evolutionary role in a Kuwait population of Diplotaxis harra (Boraginaceae). Bot J Linn Soc. 143:169–175.
Mandal GD, Nandi AK, Das AB. 2013. Cytomixis and associated meiotic abnormalities in pollen mother cells of Chlorophytum tuberosum (Roxb.) Baker. Cytologia. 78:157–162.
Mandal GD, Nandi AK. 2017. Cytomixis with associated chromosomal anomalies and the reproduction of Chlorophytum borivilianum Santapau & RR Fern. Taiwania. 62:211–215.
Mason AS, Pires JC. 2015. Unreduced gametes: meiotic mishap or evolutionary mechanism? Trends Genet. 31:5–10.
McGuire PE, Dvořák J. 1982. Genetic regulation of heterogenetic chromosome pairing in polyploid species of the genus Triticum sensu lato. Canad. J. Genet. Cytol. 24:57–82.
Melderis A. 1978. Elymus. In: Hara H, Chater AO, Williams LHJ. (eds.) An enumeration of the flowering plants of Nepal. London: British Museum (Natural History).
Miao J, Zhang X, Chen S, Ma X, Chen Z, Zhong J, Bai S. 2011. Gliadin analysis of Elymus nutans Griseb. from the Qinghai–Tibetan Plateau and Xinjiang, China. Grassl Sci. 57:127–134.
Mursalimov SR, Deineko EV. 2011. An ultrastructural study of cytomixis in tobacco pollen mother cells. Protoplasma. 248:717–724.
Mursalimov S, Deineko E. 2017. Cytomixis in tobacco microsporogenesis: are there any genome parts predisposed to migration? Protoplasma. 254:1379–1384.
Mursalimov S, Zagorskaya A, Deineko E. 2018. Evaluation of DNA damage in tobacco male meiocytes involved in cytomixis using comet assay. Protoplasma. 255:413–417. [https://doi.org/10.1007/s00709-017-1144-6]
Murti SK. 2001. Flora of cold deserts of Western Himalaya. Vol I. Monocotyledons. Kolkata: Botanical Survey of India.
Oliver SN, Dennis ES, Dolferus R. 2007. ABA regulates apoplastic sugar transport and is a potential signal for cold-induced pollen sterility 90 in rice. Plant Cell Physiol. 48:1319–1330.
Oliver SN, Van Dongen JT, Alfred SC, Mamun EA, Zhao X, Saini HS, Fernandes SF, Blanchard CL, Sutton BG, Geigenberger P, Dennis ES, Dolferus R. 2005. Cold-induced repression of the rice anther-specific cell wall invertase gene OSINV4 is correlated with sucrose accumulation and pollen sterility. Plant Cell Environ. 28:1534–1551.
Pierozzi NI, Benatti R. 1998. Cytological analysis in the microsporogenesis of ramie Boehmeria nivea Gaud. (Urticaceae) and the effect of colchicine on the chiasma frequency. Cytologia. 63:213–221.
Pusalkar PK, Singh DK. 2012. Flora of Gangotri National Park, Western Himalaya, India. Kolkata: Botanical Survey of India.
Salomon B, Bothmer RV, Yang JL, Lu BR. 1988. Notes on the perennial Triticeae species in Northern Pakistan. Bot Jahrb Syst. 110:7–15.
Sharma KD, Nayyar H. 2014. Cold stress alters transcription in meiotic anthers of cold tolerant chickpea (Cicer arietinum L.). BMC Res. Notes 7:717.
Sheila M. 1993. Male gametophyte develpoment. Plant Cell. 5:1265–1275.
Sidorchuk YV, Novikovskaya AA, Deineko EV. 2016. Cytomixis in the cereal (Gramineae) microsporogenesis. Protoplasma. 253:291–298.
Singh G. 1983. New combinations in Asiatic Elymus (Poaceae). Taxon 32:639–640.
Singhal VK, Kumar P. 2008. Impact of cytomixis on meiosis, pollen viability and pollen size in wild populations of Himalayan poppy (Meconopsis aculeata Royle). J Biosci. 33:371–380.
Singhal VK, Kumar P. 2010. Variable sized pollen grains due to impaired male meiosis in the cold desert plants of North West Himalayas (India). In: Benjamin JK. (Ed.). Pollen: structure, types and effects. New York: Nova Science Publishers, pp 101–126.
Singhal VK, Kumar R, Kumar P. 2018. A new perception about cytomixis: Causes, mechanisms and role. Chromosome Sci. 21:61–66.
Singhal VK, Singh J, Kumari V. 2018. Chromosome counts through male meiosis in seven species of genus Elymus L.(Tribe Triticeae: Poaceae) from North West Himalayas, India. Cytologia 83: 365–368.
Singhal VK, Singh J, Singh H, Kumar P, Kholia BS, Tewari, L. M. 2018. Chromosome count, meiotic abnormalities, pollen fertility and karyotype of Elymus semicostatus (Nees ex Steud.) Meld. from North-west Himalaya. Caryologia.
de Souza A, Pagliarini M. 1997. Cytomixis in Brassica napus var. oleifera and Brassica campestris var. oleifera (Brassicaceae). Cytologia. 62:25–29.
Stebbins GL. 1971. Chromosomal evolution in higher plants. London, UK: Edward Arnold.
Thomas H, Humphreys MO. 1991. Progress and potential of interspecific hybrids of Lolium and Festuca. J. Agri. Sci. 117:1–8.
Thomas HM, Thomas BJ. 1993. Synaptonemal complex formation in two allohexaploid Festuca species and a pentaploid hybrid. Heredity 71:305–311.
Vasek FC. 1962. ‘Multiple spindle’ a meiotic irregularity in Clarkia exalis. Amer J Bot. 49:536–539.
Yan XB, Guo YX, Zhao C, Liu FY, Lu BR. 2009. Intra-population genetic diversity of two wheatgrass species along altitude gradients on the Qinghai-Tibetan Plateau: its implication for conservation and utilization. Conserv Genet. 10:359–367.
Yan XB, Guo YX, Liu FY, Zhao C, Liu QL, Lu BR. 2010. Population structure affected by excess gene flow in self-pollinating Elymus nutans and E. burchan-buddae (Triticeae: Poaceae). Popul Ecol. 52:233–241.
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