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Biosystematic Studies on the Genus Ixeris (Compositae-Lactuceae) II

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Biosystematic Studies on the Genus Ixeris (Compositae-Lactuceae) II _??_1990 by Cytologia, Tokyo Cytologia 55: 553-570 , 1990 Biosystematic Studies on the Genus Ixeris (Compositae-Lactuceae) II. Karyological analyses Jae-Hong Pak1 and Shoichi Kawano2 1 Department of Botany, Faculty of Science , Kyoto University, Kyoto 606, Japan 2Department of Botany , Faculty of Science, Kyoto University, Kyoto 606, and Institute of Genetic Ecology, Tohoku University, Sendai 980, Japan Accepted April 20, 1990 Earlier karyological studies of the genus Ixeris and related groups are represented by those of Ishikawa (1921) and Babcock et al. (1937) (also cf. Takemoto 1954, 1956, 1962, 1970, Nisioka 1956, 1960, 1963, Fujishima 1984). Ishikawa (1921) studied the chromosome numbers of 20 Lactuca species from Japan, and showed that Lactuca (sensu Bentham and Hooker 1873, cum Ixeris) comprises some heterogeneous groups, with four different basic chromosome numbers, x=5, 7, 8 and 9. He further pointed out that the basic chromosome numbers and size are important criteria for the delimitation of the genera and assessment of the species relationships in Lactuca s. lat., demonstrating that those with x=5, 7, 8 and 9 base numbers are in good agreement with Nakai's generic concept (1920): Crepidiastrum (x=5), Paraixeris (x=5), Ixeris (x=7, 8), and Lactuca (x=9). However, later on Babcock et al. (1937), on the basis of an ex tensive karyological study of Crepidinae, claimed that the genus Ixeris (sensu Stebbins 1937) consists of the species with three different basic chromosome numbers, x=5, 7 and 8, which contradicts Ishikawa's viewpoint (1921). The genus Ixeris (Compositae-Lactuceae) is at present known to comprise about 20 species (Nakai 1920, Kitamura 1956, 1981) or 50 species, including some species of Crepidiastrum, Paraixeris and even Lactuca (Stebbins 1937, Tomb 1977). However, as stated above, the overall systematic status of the species referred to the genus Ixeris is still uncertain, and thus the de limitations of Ixeris and allied genera are still quite controversial. The objective of the present series of biosystematic studies is, first, to investigate more thoroughly karyological features of the plants referred to Ixeris based upon modern techniques, and second, to contribute to a better understanding of the species relationships and generic delimitations of Ixeris and allied genera. In this paper, we report the chromosome numbers and karyotypes of 1 I species and 5 subspecies, representing all 5 sections of Ixeris (sensu Kita mura 1956), collected from various localities in Japan, Korea and Taiwan; six of these species have not been karyologically well known previously (Table 1). Furthermore, in order to evaluate more precisely the data on chromosome morphology, resting nuclei were critically ex amined on diploid species, and compared with one another. Materials and methods The taxa studied, sampling sites, and habitat conditions are all given in Table 1. Materials used for karyological studies were cultivated in pots in the greenhouse of the Department of Botany, Kyoto University. The somatic chromosomes were observed in the root tip cells of more than 5 individual plants per population. The root tips were pretreated with a mixture of 3 parts of 0.002 mol 8-hydroxyquinoline and 1 part of 0.005% colchicine at 18-20•Ž for 2 hours and 40 minutes, and fixed in 45% acetic acid for 10-15 minutes at 4-8•Ž. Subsequently, 1 Present address: Department of Biology, College of Natural Science, Kyung-Pook National Univer sity, Taegu 702-701, Korea. Table 1. Localities, habitats of sampling sites, and the chromosome numbers of the genus Ixeris species examined in this study 1) Sectional classification followed Kitamura (1956). 2) The somatic chromosome numbers were examined for the first time, 3) The karyotypes were first described. 4) Voucher specimens certified by the authors. 556 Jae-Hong Pak and Shoichi Kawano Cytologia 55 the root tips were macerated in a 1:2 mixture of 45% acetic acid and 1 N HCl, stained with 1 aceto-orcein for 15 minutes, and then squashed. The karyotypes were arranged according to Levan et al. (1964) based on the positions of the centromere and their arm ratios. The de scriptions of resting nuclei followed Tanaka (1971). The voucher specimens are all preserved in the Herbarium of the Department of Botany, Kyoto University (KYO). Results 1. Morphology of resting nuclei and prophase chromosomes Two types of resting nuclei and prophase chromosomes were found in Ixeris. Fig. 1. Photomicrographs of resting nuclei and prometaphase chromosomes. A, B, Ixeris repens. C, D, I. longirostrata. E, F, I. chinensis. G, H, I. stolonifera. I, J, I. tamagawaensis. K, L, I. polycephala. Scale equals 3ƒÊm. 1) X=8 group The basic chromsome number of Ixeris repens, I. stolonifera, I. longirostrata, I. polycephala, I. tamagawaensis and I. chinensis was x=8. Their resting nuclei consisted of many dark stained, round or rod-shaped condensed bodies (Fig. 1-A, C, E, G, I, K). The regions sur rounding these bodies were smooth and stained evenly. Thus, this feature corresponds to the rod or round prochromosome type (Tanaka 1971). Prophase somatic chromosomes were differentiated clearly into early condensing and late condensing parts. Their condensation begins at the proximal part of both arms (Fig. 1-B, D, F, H, J, L). 1990 Biosystematic Studies on the Genus Ixeris II 557 2) X=7 group The basic chromosome number of Ixeris dentata, I. laevigata, I. transnokoensis and I. makinoana was x=7. Their resting nuclei consisted of many small condensed bodies in the whole region of the nucleus (Fig. 2-A, C, E, G). This corresponds to the diffuse type (Tanaka Fig. 2. Photomicrographs of resting nuclei and prometaphase chromosomes. A, B, Ixeris dentata. C, D, I. transnokoensis. E, F, I. laevigata. G, H, I. makinoana. Scale equals 3ƒÊm. 558 Jae-Hong Pak and Shoichi Kawano Cytologia 55 1971), as was shown in the Ixeris dentata complex by Fujishima (1984). No clear distinction can, however, be made between early condensing and late condensing parts in prophase. The condensation gradually proceeds from proximal to distal parts (Takemoto 1962) (Fig. 2-B, D, F, H). Fig. 3. Photomicographs of mitotic metaphase chromosome. A, Ixeris repens. B, I. longiro strata. C, L. stolonifera. D, I. polycephala. E, I. tamagawaensis. F, I. debilis. G, I. chinensis ssp. chinensis. H, I, I. chinensis ssp. strigosa. Arrows indicate secondary constrictions. Scale equals 3ƒÊm. 1990 Biosystematic Stidues on the Genus Ixeris II 559 2. Somatic chromosome numbers and karyotype constitutions Differences in basic chromsome numbers, i.e., x=7 and 8, are also reflected in the dif ferences of total genome size and the length of each chromosome complement. The total genomic size of the x=8 group varied from 25.2ƒÊm to 37.3ƒÊm: Ixeris repens (35.5ƒÊm), I. Fig. 4. Photomicographs of mitotic metaphase chromosome. A, Ixeris dentata ssp. nipponica. B, I. dentata ssp. dentata. C, I. dentata ssp. alpicola. D, I. makinoana. E, I. laevigata. F, I. transnokoensis. Arrows indicate secondary constrictions. Scale equals 3ƒÊm. 560 Jae-Hong Pak and Shoichi Kawano Cytologia 55 stolonifera (37.3ƒÊm), I. longirostrtaa (30.3ƒÊm), I. polycephala (29.3ƒÊm), I. tamagawaensis (25.2ƒÊm), I. chinensis (30.1ƒÊm). The chromosome length of the x=8 group ranged from 1.2ƒÊm to 3.7ƒÊm (Figs. 3, 5). The total genome size of the x=7 group was much larger, ranging from 43.2ƒÊm to 70.9ƒÊm: Ixeris dentata complex (61.0-70.9ƒÊm), I. laevigata (61.6ƒÊ m), I. transnokoensis (66.3ƒÊm), I. makinoana (43.2ƒÊm). The chromosome length of the x=7 group ranged from 2.2ƒÊm to 6.5ƒÊm (Fgs. 4, 6, 7). (1) Ixeris repens 2n=16 (Figs. 3A, 5A, 8A) This is a typical species found on coastal sandy beaches. Five different materials collected from Shiraoi in Hokkaido, Toyocho in Kochi, Hamatsume in Kyoto, and Kampo and Dalri Island, Korea, were karyologically examined. All the plants examined proved to possess the somatic chromosome number of 2n=16. This number was in agreement with that reported by Jinno (1953), Asano (1960) and Nishikawa (1984). Ishikawa (1921, as Lactuca repens) also reported the haploid chromosome number of n=8 (Table 1). The karyotype of this species is described for the first time in this study (Figs. 3A, 6A). The chromosome complement consisted of six median (Nos. 1-5, 8) and two submedian pairs (Nos. 6, 7). The first and eighth pairs were heterozygous regarding the satellite at the distal portion of their short arms. (2) Ixeris polycephala 2n=16 (Figs. 3D, 5B, 8B) Ixeris polycephala is an annual species which grows mainly in crop gardens and/or at the edge of fields. Two populations from Kyongju and Chuhulsan in Korea were examined. All materials had the somatic chromosome number of 2n=16. Ishikawa (1921, as Lactuca ma tsumurae) and Shetty (1967) reported that the haploid chromosome number was n=8 in plants from Japan and India (Table 1). The chromosome complement was six median (Nos. 1-6) and two submedian pairs (Nos. 7, 8) (Figs. 5B, 8B). The largest pairs had satellites at the distal portion of their short arms (No. 1). The somatic chromsosome number and the karyo type of this taxon are reported for the first time in this paper. (3) Ixeris stolonifera 2n=16 (Figs. 3C, 5C, 8C) This is a typical species of the open grassy slopes as well as on foothills along roadsides in the lowlands. Materials collected from three different populations, i.e., Hieizan, Takinoike and Garyusan in central Honshu, were examined. All the materials turned out to possess 2n=16 somatic chromosomes. The results are in agreement with those of the previous reports (Babcock et al. 1937, Takemoto 1952, Lee 1971, Nishikawa 1979). Ishikawa (1911, 1916, 1921, as Lactuca stolonifera) reported its haploid number as n=8 (Table 1).
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