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Cytological Investigations on W. Himalayan Pooideae P. N. Mehra

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Cytological Investigations on W. Himalayan Pooideae P. N. Mehra Cytologia 38: 237-258, 1973 Cytological Investigations on W. Himalayan Pooideae P. N. Mehra and P. Remanandan Departmentof Botany,Panjab University, Chandigarh,India ReceivedSeptember 9, 1971 Introduction In recent years the agricultural scientists in India have started realising the severity of shortage of cattle feed and a number of exotics have been introduced and tried for fodder purposes. The results have not been much encouraging . What is perhaps needed is the exploitation of the indigenous material for which the Hima layas are exceptionally rich. The moist hills and valleys of these mountains possess excellent grass-lands which extend up to 4,500m. A very small proportion of these meadows is utilized for grazing by goats, sheep and yaks. With such a wealth of indigenous species which give excellent green stuff and can be made into palatable hay or silage it is worthwhile to make selection and effect systematic improvement of these by cytogenetic methods. The objective of the present study was to investigate the cytology and distribu tional pattern of Indian grasses to enrich the germplasm and also to delineate the evolutionary trends based on the meiotic behaviour of chromosomes. The first paper of this series (Mehra et al. 1968) presented the cytological data on 145 species belonging to 68 genera mostly distributed in Punjab plains and nearby hills. An attempt was made to trace the evolutionary trends and a scheme for the probable cytological evolution of the family was presented. The second paper (Mehra and Shyamsunder 1970) presented the cytological data on 63 species belonging to 23 genera, distributed mainly in the Kashmir hills. The present paper is a continua tion of these earlier works and cytological data of 33 taxa belonging to 28 species comprising 16 genera is presented. Material and methods The material was collected from wild populations in the Kashmir and Kumaon hills of the northwest Himalayas. The grasses were identified in the field with the help of Bor's keys (Bor 1960) and confirmed by comparing them at the herbaria of Forest Research Institute, Dehradun, and Botanical Survey of India zonal head quarters, Dehradun. The voucher specimens and the permanent slides have been deposited in the herbarium of the department of Botany, Panjab University, Chan digarh. Cytological studies were made at meiotic divisions of microsporocytes. Flower buds were fixed in Carnoy's fluid and simple acetocarmine squashes were prepared 238 P. N. Mehra and P. Remanandan Cytologia 38 Table 1 1973 Cytological Investigations on W. Himalayan Pooideae 239 Table 1. (contd.) 240 P. N. Mehra and P. Remanandan Cytologia 38 Table 1. (contd.) 1973 Cytological Investigations on W. Himalayan Pooideae 241 Table 1. (contd.) 242 P. N. Mehra and P. Remanandan Cytologia 38 for the study of microsporogenesis. To detect intraspecific cytological races, same species was screened from a wide range of localities. All the photomicrographs were taken at a magnification of •~1,130. Slides were made permanent following the procedure of Bhaduri and Ghosh (1954). Results Table 1 summarizes the results of investigations on 28 species. Five species, marked * are worked out for the first time, whereas those marked t represent new cytotypes. A+mark on the•eprevious authors'side indicates that this species has been extensively investigated and only the more significant reports are included. In the succeeding pages only the more interesting features, such as the meiotic aberrations and comparative morphology of the intraspecific cytological races are briefly discussed. Meiotic irregularities It has been noticed that most of the grasses of alpine regions, as a rule, do not set seeds, or if they do so, the seeds are mostly sterile. This is ascribed to a number of causes such as environmental factors, extensive vegetative method of multipli cation, browsing by animals, and genomic instability as reflected in the various meiotic aberrations and subsequent production of sterile pollen. Alopecurus Three species of this genus were investigated of which A. arundinaceus is a valu able constituent of the alpine pastures. In Kashmir A. himalaicus has been noticed only on hill tops of snowy ranges which are not easily accessible to cattle. How ever, it has been reported that cattle and yak do relish it (Bor 1960). Meiotic disturbances were observed in all the three species. A. aequalis is a diploid with 2n=14. In this species as many as 8 chromosomes were found to be involved in the formation of a complex structure with chiamata between nonhomologous chromosomes, representing reciprocal translocations. In majority of cases this structure did not form a complete ring and two free ends were noticeable (Fig. 1). It showed delayed separation of chromosomes at A, (Fig. 2). At late AI laggards were common (Fig. 3). Only 75% of pollen grains were stainable. A. arundinaceus is a tetraploid and 14 bivalents were observed in some cells at diakinesis (Fig. 4). However, at MI several pollen mother cells showed a large ring suggestive of translocations, obviously involving several chromosomes (Fig. 5). Further course of meiosis was regular and 90% of the pollen grains were fertile. Figs. 1-9. 1-3. Alopecurus aequalis, n=7. 1, diakinesis showing 3 bivalents and a multivalent body consisting of 8 chromosomes indicating structural heterozygosity for reciprocal translocation. 2, late separation of multivalents at AI. 3, late AI showing laggards at the equator. 4-5. A. arundinaceus, n=14. 4, diakinesis showing 14 bivalents. Note the chromatin link between chro mosomes of two bivalents (arrow). 5, Mi showing a multivalent ring. 6-9. A. himalaicus, 2n= 45. Illustrations demonstrating meiotic breakdown. 6, nonorientation of univalents at MI. 7, AI showing 2n=45. 8, laggards at late AI. 9, Two chromatin bridges (arrows) and laggards at late AI. •~l,130. 1973 Cytological Investigations on W. Himalayan Pooideae 243 244 P. N. Mehra and P. Remanandan Cytologia 38 A. himalaicus is a newly investigated species with 2n=45 (Fig. 7). Meiotic studies revealed that the divisions undergo a highly disturbed course resulting in various aberrations which lead to a total breakdown of meiosis, and consequent production of sterile pollen. The following were the chief aberrations noticed: i) Nonhomologous associations, ii) loose pairing of chromosomes at diakinesis, iii) nonorientation of univalents at M, (Fig. 6), iv) nonsynchronised disjunction at AI, v) bridges (Fig. 9), laggards (Figs. 8, 9), vi) fragments and vii) unequal distribu tion of chromosomes at A, and AII. Although spindle apparatus was formed, the congression of chromosomes at metaphase plate was imperfect and a number of univalents were found scattered throughout the cytoplasm. This is the first report of a taxon in Alopecurus with 2n=45, all the other cytologically known species of the genus having been found to possess the basic number 7. Cytological evidence suggests that the species may probably be of hybrid origin and meiotic breakdown is due to its unbalanced genomic constitution. Calamagrostis C. pseudophragmites showed a regular cycle of meiosis with 14 bivalents at diakinesis. Thirteen were the ring bivalents with terminalised chiasmata and one bivalent had interstitial chiasma (Fig. 10). However, in C. emodensis meiosis was found to undergo a highly abnormal course leading to total meiotic breakdown. Well spread metaphase plates could not be obtained due to stickiness, secondary associations, and clumping of chromosomes. The taxon, however, appears to be a triploid with 2n=21, but this needs confirmation from mitotic preparations. Polypogon Two species, P. fugax and P. monspeliensis, which provide a rich feed for graz ing animals, have been investigated. The former is a hexaploid (n=21, Fig. 12) and the latter a tetraploid (n=14). These reports are in line with the previous records of many authors (cf. Table 1). However, Mehra et al, (1968) reported n=14, 21 in P. monspeliensis. Their voucher specimen for the taxon with n=21 probably is that of P. fugax. These two species are closely related and are dis tinguished by the relative length of the awn which is shorter in the latter species compared to the former. Avenafatua Linn. This species closely resembles the cultivated species A. sativa Linn. But it can be readily distinguished by its rhachilla which disarticulates between all the florets at maturity and the presence of a thickened callus at the base of all lemmas. The present studies revealed that the species exists in two cytological races, a diploid (n=7) and a tetraploid (n=14). The diploid race, which is investigated for the first time, was detected in Kashmir, while the tetraploid in Kumaon hills. A comparative morphological analysis of the taxa is presented in Table 2 and they are illustrated in Fig. 42. The diploid is evidently more sturdy than the tetraploid. Figs. 10-19. 10, Calamagrostis pseudophragmites, n=14. Diakinesis displaying chiasmata dis tribution. 11, Phleum alpinum, n=14. Diplotene displaying formation of chiasmata. 12, Polypogon fugax, n=21. 13-14. Avena fatua, n=7. 13, diakinesis showing 7 rings. 14, early 1973 Cytological investigations on W . Himalayan Pooideae 245 disjunction at AI. 15-19. A. fatua, n=14. 15-16. early and late diakinesis displaying chiasmata distribution. 17, MI with 14II. 18, MI with 12II and 1VI (arrow). 19, 14: 14 separation at AI. •~1,130. 246 P. N. Mehra and P. Remanandan Cytologia 38 The diploid taxon showed 7 ring bivalents and a well differentiated nucleolus at diakinesis (Fig. 13). Mild disturbances were noticed at AI. Precocious separa tion of some chromosomes (Fig. 14) was occasionally observed. However, further stages were normal. In most of the cells at late AI 7:7 distribution was noticed. 95% pollen were viable. Figs. 15-16 display the chiasmata distribution in the tetraploid at diakinesis. Table 2. Avena fatua* * Average of 10 measurements in each case .
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