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Download PDF (2126K) _??_1994 The Japan Mendel Society Cytologia 59: 295 -304 , 1994 Cytotypes and Meiotic Behavior in Mexican Populations of Three Species of Echeandia (Liliaceae) Guadalupe Palomino and Javier Martinez Laboratorio de Citogenetica, Jardin Botanico , Instituto de Biologia, Apartado Postal 70-614, Universidad Nacional Autonoma de Mexico, D . F. 04510, Mexico Accepted June 2, 1994 Echeandia Ort. includes herbaceous perennials distributed from the Southwestern United States to South America. More than 60 species have been described from Mexico and Central America, many of which are narrow endemics (Cruden 1986, 1987, 1993, 1994, Cruden and McVaugh 1989). Mexico is considered the center of origin and evolution for this genus (Cruden pers. comm.). They are commonly found in pine and pine/oak forest, grasslands; xerophyte shrublands, and disturbed areas, (Cruden 1981, 1986, 1987, Cruden and McVaugh 1989). Except polyploidy species based on n=8, as E. longipedicellata n=40, (Cruden 1981); E. altipratensis n=24, and 48; E. luteola n=32, and 64; E. venusta n=84, (Cruden 1986, 1994), chromosomes those exist, little information on interspecific or intraspecific variation in karyo types in Echeandia. In 1988, Palomino and Romo described the karyotypes of E. flavescens (Benth.) Cruden (as E. leptophylla) and E. nana (Baker) Cruden, both of which are characterized by 2 pairs of chromosomes with satellites. Cytotype variation has been observed commonly in the families Liliaceae, Iridaceae, Commelinaceae, and some Poaceae, Acanthaceae and Leguminosae. It is usually due to the presence of spontaneous aberrations in number or structure, heterozygous inversions, Roberts onian translocations, exchanges, deletions and duplications (Sen 1975, Araki 1975, Araki et al. 1976, Brighton 1976, 1977a, b, Datta and De 1990, Christopher and Jacob 1990, Sakya and Joshi 1990, Vijayavalli and Mathew 1990). Heteromorphic bivalents and/or bridges with or without fragments are evidence of these structural aberrations (Brandham 1970, Brandham and Johnson 1977, Jones 1978, Kenton 1981, Palomino and Vazquez 1991). Cytotype distribution is established by geographical isolation and natural selection (Kenton 1981, Kenton et al. 1988). There is also evidence of polyploid cytotypes reported by Jones et al. (1981), Araki (1985), Kumar and Gohil (1990), Piovano and Bernardello (1991). Sometimes cytotypes can be transmitted via sexual reproduction. They are usually maintained by asexual reproduction forming a cluster of plants with the same cytotype (Haga and Noda 1976, Araki 1985, Araki et al. 1976). We describe variation in the cytotypes of three species, the behavior of meiotic chromo some, and pollen fertility for populations of Echeandia echeandioides, E. tenuis and E. mexicana. Material and methods The species studied were: E. echeandioides Cruden (=Anthericum echeandioides Baker), which is endemic species to central part of Mexico, E. tenuis (Weatherby) Cruden and E. mexicana Cruden. Plants were collected from wild populations in pine oak forest (Appendix 1) and voucher specimens were deposited at the National Herbarium (MEXU) of the Universidad Nacional Autonoma de Mexico. 296 Guadalupe Palomino and Javier Martinez Cytologia 59 Three to six individuals from two or three populations of each species (Appendix 1) moved to the Jardin Botanico. The plants were transplanted into pots containing a mixture of vermiculite and organic soil and maintained in a greenhouse. Elongating secondary root tips were placed in a saturated solution of paradichlorobenzene for 6 hr at 4•Ž. They were staining following Feulgen technique (Palomino and Vazquez 1991). Preparations from 3 to 6 plants were made, and ten cells at mitotic metaphase were selected from each for examination. Three of the best cells from each plant of each population were photographed using a Zeiss Photomicroscope II. Idiograms were made using a Zeiss Drawing Apparatus. Chromosomes were classified according to Levan et al. (1964) and Naranjo et al. (1986). Index of asymmetry (TF% or TF index) was obtained following Gupta and Gupta (1978). To study meiotic behavior fresh anthers from young buds were squashed in 1.8% aceto-orcein, and 168 pollen mother cells (PMC) in metaphase I (MI) and 913 to 998 PMC in anaphase I (AI) were analyzed. For MI PMCs, from each population the following informa tion was recorded: type of bivalent (IIs) and quadrivalent (IVs), chiasmata frequency and recombination index (RI, White 1973). In AI PMCs recorded data were: single bridge, double bridge, single bridge with fragment and AI with lagging chromosomes. Pollen fertility was estimated by staining samples with cotton blue in lactophenol. Percentage of well-filled stained grains was obtained. This was carried out for 238 to 877 pollen grain cells of three plants in each population studied. The student "t" test, was applied to detect differences among haploid chromatin length in the populations analyzed. Results All the populations of the three species we studied were diploid with 2n=16 and n=8 chromosomes and in which two pairs of chromosomes had satellites. Each species had a different karyotype (Table 1, Figs. 1, 2) and their cytotypes were uniform within populations but varied within the species studied. Cytotypes of E. echeandioides varied in the number of metacentric (m) and submetacentric (sm) chromosomes. Population 359 had 3 pairs of heteromorphic chromosomes (Figs. 1A, 2A) whereas population 360 had a single heteromor phic pair of chromosomes (Figs. 1B, 2B). Population 321 the least variable cytotype. All the chromosomes were homomorphic and one pair had subtelocentric chromosomes (st), (Table 1, Figs. 1C, 2C), which were absent from the other populations. The two populations of E. tenuis had cytotypes that varied in quantity of m, sm and st chromosomes (Table 1). Population 356 had a pair heteromorphic chromosomes (Figs. 1D, Table 1. Cytotypes of three species of Echeandia 1994 Cytotypes and Meiotic Behavior in Mexican Populations of Three Species of Echeandia 297 Fig. 1. Cytotypes for: A. E. echeandioides (359). B. E. echeandioides (360). C. E. echeand ioides (321). D. E. tenuis (356). E. E. tenuis (484). F. E. mexicana (236). G. E. mexicana (284). H. E. mexicana (357). Numbers 1 and 2 indicate chromosomes with satellites. Scale equals 10ƒÊm. 2D) and population 484 2 pairs (Figs. 1E, 2E). Three populations of E. mexicana were examined. The population 236 and 284 had different cytotypes and exhibited variation in number of m, sm and st chromosomes. Popula tion 357 had only m and sm chromosomes (Table 1). None of the cytotypes of E. mexicana had heteromorphic pairs (Figs. 1F, 2F, 1G, 2G, 1H, 2H). Variation in the genome size was confirmed by the significantly different haploid chromatin lengths (P<0.05, Table 1). Meiotic behavior was shown by the average number of chiasmata per cell and the recombination index. Ring and rod bivalent with variable frequencies were found at MI (Table 2). All the populations analyzed had 1 to 3 heteromorphic bivalents (Figs. 3A, B, C, D). E. echeandioides #359 exhibited a ring and heteromorphic IVs with frequencies of 0.47 and 0.23 (9.7%) respectively. The population 360 had only a heteromorphic chain IVs with a frequency of less than 0.14 (1.7%), (Table 2). All the populations we studied exhibited aberrations at AI with variable frequencies (Table 298 Guadalupe Palomino and Javier Martinez Cytologia 59 3, Fig. 4). E. echeandioides #359 and 360 had cells with bridge and fragment, sometimes one or two bridges, and lagging chromo somes. The population 321, did not display bridges with fragment, or lagging chromo somes (Table 3, Fig 4A). Aberrant AI cells occurred in all populations of all three spe cies that had many shrunken or empty pollen grains, (Table 4). E. tenuis #484 displayed more structural chromosomal aberrations (Table 3, Figs. 4A, C), and many non-viable pollen grains (Table 4) compared to population 356. E. mexicana #284 and 357 had higher percent age of AI aberrations than population 236. No lagging chromosomes were observed (Table 3, Fig. 4B). The highest percentage of non-viable pollen found in E. mexicana occurred in collections 236 and 357 (Table 4). Discussion The populations of the three species we examined were diploid with 2n=16 and n=8 (x=8) chromosomes. Each species had a distinctive karyotype that varied among pop Fig. 2. Idiograms for: A. E. echeandioides (359). B. ulations. Intraspecific cytotype variation was E. echeandioides (360). C. E. echeandioides (321). D. manifest as heteromorphic chromosome E. tenuis (356). E. E. tenuis (484). F. E. mexicana (236). G. E. mexicana (284). H. E. mexicana (357). pairs in E. echeandioides and E. tenuis, num Asterisks indicate pairs of heteromorphic chromo bers of metacentric, submetacentric, and sub somes. Scale equals 10ƒÊm. telocentric chromosomes, and chromosomes with satellites. In addition, Cruden (1981) reported tetraploid populations of E. mexicana from Michoacan and Jalisco. Similar intraspecific cytotype variation has been reported in other Liliaceae. Heteromor phic chromosome pairs resulting from asymmetrical exchanges were reported in Scilla (Sato 1942, Gimenez-Martin 1959, Haga and Noda 1958, Noda 1961), Gloriosa superba (Vijayavalli and Mathew 1990), and in tribe Aloineae (Brandham 1974, 1976). In Scilla, the heteromorphic chromosome pairs were attributed to translocations and deletions. In Scilla scilloides, Araki (1975, 1977, 1985), Araki et al. (1976) studied 46 natural populations, and reporting diploid, polyploid and
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