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Cytological Studies in Dioscoreaceae

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Cytological Studies in Dioscoreaceae 1968 401 Cytological Studies in Dioscoreaceae K. Ramachandran Department of Botany, University of Kerala , Trivandrum, India Received November 8, 1967 Introduction The family Dioscoreaceae consists of ten genera and about 650 species (Lawrence 1960), dominated by the genus Dioscorea which accounts for about 620 species. The family is distributed in all tropical regions and a few species are also found in temperate Europe and North America . The genus Dioscorea is of great economic importance. A few species , the true yams, are cultivated for edible tubers in many tropical countries. The genus is also of interest as tubers of some wild species yield precursors of cortisone and related steroid drugs. The dioecious species of Dioscorea received the attention of cytologists studying the sex chromosome mechanism in plants (Meurman 1925, Nakajima 1933, 1934, 1936, 1937, 1942, Smith 1937). Recently, since the discovery of cortisone precursors in tubers of some wild species, a great deal of work on the cytology and experimental breeding of Dioscoreas has been done by Martin and co-workers (Martin 1966, Martin and Cabanillas 1963, 1966, 1967, Martin and Ortiz 1963, 1966, Martinet al. 1963). These studies have shed valuable light on the pattern of evolution in the family. Cytological studies in the family have been summarised by Martin and Ortiz (1963). Previously, Indian species of Dioscorea were studied by Sundara Raghavan (1958, 1959). A list of chromosome numbers of 9 species and a few varieties are given in his earlier paper, but no figures of chromosomes have been published. Chromosome numbers of two more species were reported later (Sundara Raghavan 1959). Sharma and De (1956) and Sharma and Sharma (1957) studied two species of Dioscorea. The present study was undertaken as part of a larger programme on the breeding of the yams. A preliminary report of this work was published earlier (Ramachandran 1962) in which chromosome numbers and sex determi nation mechanism in Dioscorea were discussed. The present paper gives a detailed account of the cytology of all the South Indian species studied. Materials and methods A large collection of cultivated and wild species of Dioscorea and Tri chopus zeylanicus was made from the different parts of Kerala State, South India. Three clones of D. bulbifera were also obtained from Poona, Western 402 K. Ramachandran Cytologia 33 1968 Cytological Studies in Dioscoreaceae 403 India. The plants were grown in the botanical garden of the Kerala Uni versity, Trivandrum. Cytological studies were made from acetocarmine smears . Root tips for somatic chromosome studies were obtained from yam heads , aerial tubers or, in a few cases, leaf-bud cuttings planted in pots . Root tips were kept at 6•K-10•Ž for an hour and fixed in Carney's fluid (alcohol , acetic acid and chloroform in the proportion of 3:1:1) mordanted with a trace of iron-acetate. For meiotic studies male spikes were fixed in alcohol-acetic acid (3:1). Anthers were dissected out from buds and smeared in acetocarmine . Observations Dioscorea Plim. ex L. The genus consists of about 620 species , distributed in the tropics, with a few species in temperate Europe and North America . A few species are cultivated in most tropical countries. The four species commonly cultivated in South India are D. alata, D. esculenta, D. sativa and D. spinosa. All these have been investigated in the present study. 1. D. alata L. Known variously as 'white yam' , 'water yam' or ' winged yam', this is the most important of the yams, being very widely cultivated in Asiatic tropics, Africa, the Pacific Islands and tropical America. There are a large number of varieties of this species under cultivation. Twenty six clones of this species have been assembled in our collections. They differ chiefly in their growth habit, shape and size of leaves, and shape, size and coloration of skin and flesh of the yams. Other differences relate to the presence or absence of spines, colouration of stem and petioles and shape and size of bulbils. Twenty three out of the twenty six clones of D. alata had 40 chromo somes, two had 60 and the other 80 chromosomes in root tip cells. Fig. 1 shows the somatic chromosomes of a male plant having 40 chromosomes. They are small ranging between 0.6ƒÊ and 2.1ƒÊ in length. One of the chro mosomes is appreciably larger than the others (indicated by arrow in figure). Meiosis was normal and 20 bivalents have been observed at metaphase I (Fig. 2). No irregularities have been found in the few cells examined. Two clones (both female plants) of D. alata in our collection had 60 chromosomes in root tip cells. Fig. 3 shows the chromosome complement of one of these. The chromosomes are between 0.6 and 1.7ƒÊ in length. D. alata var. purpurea Roxb. Female plants of this variety collected from different localities showed Figs. 1-9. Chromosomes of Dioseorea species. •~1250. 1, D. alata L., 2n=40. Note one larger chromosome (possibly Y) indicated by arrow. 2, D. alata L. Diakinesis in a pollen mother cell of a tetraploid plant showing 20 bivalents. 3, D. alata L., 2n=60. 4, D. alata var. purpurea Roxb., 2n=80. 5, D. bulbifera L., clone l from Maharashtra, 2n=80. 6, D. bulbifera L., clone from Ponmudi (Kerala), 2n=80. 7, D. bulbifera L., clone 2 from Maharashtra, 2n=80. 8, D. bulbifera L., clone 3 from Maharashtra, 2n=80. 9, D. bulbifera L., clone from Karthicappally (Kerala), 2n=80. 404 K. Ramachandran Cytologia 33 1968 Cytological Studies in Dioscoreaceae 405 2n=80 chromosomes (Fig. 4). Male plants have not been collected so far . 2. D. bulbifera L. Five different clones of D . bulbifera, 2 from Kerala State and 3 from Maharashtra State possessed 80 somatic chromosomes . Another plant from Kozhencherry (Kerala) showed 2n=100 chromosomes . The somatic chromosomes of the six clones of D . bulbifera are shown in Figs. 5-10. Fig. 11 shows a group of pollen mother cells from a male plant of clone 2. Meiotic chromosomes at metaphase I are seen clumped together . No irregularities have been found in subsequent stages of meiosis . Ninety per cent of pollen grains appeared to be normal , as shown by staining with acetocarmine. 3. D. esculenta Burkill. This is a widely cultivated species , charac terised by slender round stems with small prickles , cordate leaves and several underground tubers in a cluster. There are two easily recognisable cultivated varieties of this species. One variety ('Nanakizhengu') has larger leaves and tubers than the other ('Cherukizhengu'). The former variety had 90 somatic chromosomes (Fig. 12), while the later showed 100 chromosomes (Fig. 13). 4. D. hispida Dennst. This is a common wild species of the Western Ghats. Plants of this species showed 40 chromosomes in root tip cells (Fig. 14). Chromosomes are between 0.9 and 2.7ƒÊ in length. 5. D. oppositifolia L. var. linnaei Pr. and Burk. The chromosome complement of this variety (2n=40) is illustrated in Fig. 15. Chromosomes range between 1ƒÊ and 2ƒÊ in length. Another clone of D. oppositifolia collected from Kannikatty (Madras State) also showed 40 somatic chromosomes (Fig. 16). 6. D. pentaphylla L. Two varieties of D. pentaphylla have been ex amined cytologically in the present study. D. pentaphylla var. linnaei Pr. and Burk. The chromosome number in male plants of this variety collected from Kallar (Kerala State) is 2n=40 (Fig. 17). One chromosome of the com plement is larger than the others being 2ƒÊ long. The others are between 1.2ƒÊ and 1.8ƒÊ in length. D. pentaphylla var. rheedei Pr. and Burk. This is found cultivated in some places in Kerala and produces smooth white cylindrical tubers and elongated bulbils. A female plant of this variety has been examined cytologically. It possessed 40 chromosomes (Fig. 18). Chromosome length ranges between 1.2 and 1.8ƒÊ. Figs. 10-16. Chromosomes of Dioscorea species. All figures except Fig. 11. •~1250. 10, D. bulbifera L., clone from Kozhencherry (Kerala), 2n=100. 11, a group of pollen mother cells from the clone of D. bulbifera from Ponmudi. •~750. 12, D. esculenta Burkill, 2n=90. 13, D. esculenta Burkill, 2n=100. 14, D. hispida Dennst., 2n=40. 15, D. op positifolia L. var. linnaei Pr. and Burk., 2n=40. 16, D. oppositifolia L. from Kannikatty (Madras State), 2n=40. 406 K. Ramachandran Cytologia 33 1968 Cytological Studies in Dioscoreaceae 407 7. D. sativa Thun. (The Potato yam). This species is cultivated widely in tropical Asia, West Indies and Africa for its edible aerial tubers. Female plants of this species were cytologically studied. There are clearly 80 chromosomes in root tip cells (Fig . 19). 8. D. spinosa Roxb. Santapau (1951) considers this as synonymous with D. esculenta. But it is quite distinct with very large tubers (30-40cm long and 15-25cm in diameter) and numerous large spines at the base. The tubers of this species are also edible. It is cultivated in many parts of Kerala and has also been reported as occurring wild in some parts of India. Male plants of this species showed 2n=90 chromosomes (Fig. 20). One chromosome, 2.4ƒÊ long, is appreciably larger than the others, which range between 0.75ƒÊ and 1.8ƒÊ in length. 9. D. tomentosa Koenig. Male plants of this species have been cyto logically examined in the present study. They showed 40 chromosomes in root tip cells (Fig. 21). One chromosome is remarkably larger than the others of the complement. At metaphase I of meiosis twenty bivalents have been observed in pollen mother cells (Fig. 22). The larger chromosome found in somatic plates is found paired with one of the medium sized chromosomes to form an asym metrical bivalent.
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