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High Levels of Genetic Variation in Korean Populations of Sasamorpha Borealis (Poaceae)

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High Levels of Genetic Variation in Korean Populations of Sasamorpha Borealis (Poaceae) LeeBot. andBull. Chung Acad. Sin. Genetic (1999) 40:variation 311-317 in Sasamorpha borealis 311 High levels of genetic variation in Korean populations of Sasamorpha borealis (Poaceae) Nam Won Lee and Myong Gi Chung1 Department of Biology, Gyeongsang National University, Chinju 660-701, The Republic of Korea (Received June 2, 1998; Accepted April 1, 1999) Abstract. Genetic and genotypic diversity of Korean populations in Sasamorpha borealis (Hack.) Nakai were inves- tigated using allozymes as genetic markers. In Korea, S. borealis usually grows on hillsides under pine-oak understory. Many species of bamboos have the intermast period of mast seeding. After spreading by extensive rhizomes for a specific period, nearly all adults in one area produce wind-pollinated flowers, set large quantities of seeds, and die. This study was undertaken to infer relationships between levels of genetic diversity and the reproductive biology of the species. Populations of S. borealis maintain high levels of genetic diversity: 51% of the loci examined were polymorphic and of mean genetic diversity (H = 0.219). The mean number of multilocus genotypes per population eP was 18.4, and the mean genotypic diversity index (D ) was 0.720. However, 31% of the total genetic variation was G found among populations (G = 0.310). Allele frequencies for all loci examined differed significantly among popu- ST lations (p < 0.001). Average genetic identity for all pairs of populations was 0.833 (SD = 0.046). Indirect estimates of the number of migrants per generation (Nm) were 0.56 (calculated from G ) and 0.20 (calculated from the mean ST frequency of ten private alleles). Several biological and ecological traits found in S. borealis (widespread geographical distribution, long-lived habit, clonal reproduction with relatively high genotypic diversity, wind-pollinated breeding system, death of adults bearing large quantities of seeds, probable low gene flow among populations, selection for heterozyous genotypes in some of the sampled populations, the patterns of recolonization, effects of genetic drift, and local selective forces) may have played roles in shaping the population genetic structure of the species. Keywords: Allozymes; Clonal diversity; Genetic diversity; Mast seeding; Sasamorpha borealis; Selection. Introduction that clonal reproduction may act as an enhancer of genetic drift by reducing the effective size of local populations (e. Electrophoretic techniques for the study of clonal plants g., Pleasants and Wendel, 1989; Bayer, 1990; Murawski and provide powerful genetic markers for recognizing indi- Hamrick, 1990; Aspinwall and Christian, 1992; Chung, 1994; vidual plant genotypes (Berg and Hamrick, 1997). This Chung and Kang, 1996; Yeeh et al., 1996). Since the rela- approach has made it possible to better understand the tionship between genetic diversity and mode of reproduc- spatial distributions of clones and the genotype diversity tion remains unclear, further study of clonal plants is called maintained within populations. According to a thorough for. review of studies of clonal plants (Ellstrand and Roose, According to a previous review of bamboo biology 1987), species with predominantly vegetative reproduction (Janzen, 1976), many of the more common Asian species generally have lower levels of genetic diversity than spe- spread by rhizomes for a species-specific period (e.g., Sasa cies that successfully produce progeny solely by sexual tessellata has an intermast period of > 115 years). After reproduction. Previous studies have revealed that veg- reaching a specific period, most Asian bamboos produce etative reproduction and spread have a marked effect on wind-pollinated flowers, synchronously set large quanti- the genetic structure of populations. As Eriksson (1993) ties of seed (mast seeding), and die (Janzen, 1976). The pointed out, however, mechanisms for the maintenance of biology of bamboos may be of interest to plant popula- genetic variation in clonal plants are still controversial. For tion biologists. Here we report levels and partitioning of example, several results indicate that clonal reproduction allozyme diversity within and among populations and the could retard the loss of genetic diversity within popula- extent of clonal reproduction within populations of tions because species with independent ramets could re- Sasamorpha borealis (Hack.) Nakai (Poaceae), a small duce the possibility of genet death (e.g., Parker and bamboo. Sasamorpha borealis is widely distributed in the Hamrick, 1992; Berg and Hamrick, 1994; Lokker et al., 1994; Korean peninsula and Japanese archipelago. In Korea, Kim and Chung, 1995a,b; Mayers et al., 1998; Chung and populations of S. borealis are large and commonly found Chung, 1999; Chung and Epperson, 1999). Others showed on hillsides under pine-oak understory. Unfortunately, in- formation on reproductive biology of the study species (e.g., the exact period of masting and the range of syn- 1Corresponding author. Fax: (0591) 54-0086; Tel: (0591) 751- chronization of the masting period in Korea) is not 5953; E-mail: [email protected] available. 312 Botanical Bulletin of Academia Sinica, Vol. 40, 1999 Materials and Methods Enzyme Extraction and Electrophoresis Leaf samples were cut finely, and crushed with a mor- Population Samples tar and pestle. A phosphate-polyvinylpyrrolidone extrac- A total of 446 leaf samples were collected from ten popu- tion buffer (Mitton et al., 1979) was added to the leaf lations of S. borealis in Korea (Figure 1). Population samples to facilitate crushing and to aid enzyme samples of 18 to 73 were employed. Population codes and stabilization. The crushed extract was absorbed onto 4× sample sizes are given in Table 1. Because the species 6-mm wicks cut from Whatman 3 MM chromatography pa- exhibits extensive clonal growth, samples were collected per and were stored at -70°C until needed for analysis. randomly at intervals of > 5m within patch or population Electrophoresis was performed using 10.5% starch gels. to avoid biasing samples toward certain clones. Leaf Fourteen putative loci for S. borealis from nine enzyme samples were placed in plastic bags wrapped with a wet systems were resolved using a Poulik buffer system, a paper towel and stored on ice and transported to the modification (Haufler, 1985) of Soltis et al. (1983) system laboratory. Samples were then stored at 4°C until protein 6. The nine enzyme systems were acid phosphatase extraction. (ACPH), diaphorase (DIA), fluorescent esterase (FE), glutamate dehydrogenase (GDH), leucine aminopeptidase (LAP), peroxidase (PER), phosphoglucoisomerase (PGI), phosphoglucomutase (PGM), and triosephosphate isomerase (TPI). Enzyme staining followed protocols of Cheliak and Pitel (1984) for DIA and Soltis et al. (1983) for all others. Putative loci were designated sequentially, with the most anodally migrating isozyme designated 1, the next 2, etc. Likewise, alleles were designated sequentially with the most anodally migrating alleles designated a. Although the genetic bases of the loci were not documented by con- trolled crosses, the isozymes expressed phenotypes that were consistent in subunit structure and genetic interpre- tation with other isozyme studies in plants, as documented by Weeden and Wendel (1989). Data Analyses A locus was considered polymorphic if two or more al- leles were detected, regardless of their frequencies. Five standard genetic parameters were estimated using a com- puter program developed by M. D. Loveless and A. Schnabel: percent polymorphic loci (P), mean number of Figure 1. The location of the ten populations in Korea exam- alleles per polymorphic locus (AP), mean number of alle- les per locus (A), effective number of alleles per locus (A ), ined in this study (alphabetic codes as in Table 1). e Table 1. Summary of allozyme variation and clonal diversity for 14 loci within ten populations of Sasamorpha borealisa. Popb ALc Nd PAPAA H (SD) H (SD) G G/N D PG e o e G A 950 27 42.86 2.00 1.43 1.39 0.378 (0.025) 0.200 (0.065) 4 0.148 0.556 0.593 B 480 57 64.29 2.56 2.00 1.39 0.262 (0.016) 0.227 (0.056) 38 0.667 0.978 0.123 C 20 44 35.71 2.00 1.36 1.24 0.227 (0.017) 0.125 (0.056) 3 0.068 0.342 0.795 D 350 73 57.14 2.00 1.57 1.55 0.500 (0.016) 0.281 (0.068) 2 0.027 0.106 0.862 E 450 32 50.00 2.29 1.64 1.42 0.362 (0.023) 0.222 (0.064) 15 0.469 0.938 0.156 F 430 65 57.14 2.50 1.86 1.35 0.237 (0.014) 0.209 (0.055) 32 0.492 0.514 0.185 G 320 57 64.29 2.33 1.86 1.49 0.251 (0.015) 0.270 (0.058) 42 0.737 0.983 0.105 H 300 32 57.14 2.13 1.64 1.47 0.333 (0.022) 0.245 (0.065) 24 0.750 0.976 0.125 I 750 18 35.71 2.40 1.50 1.38 0.294 (0.029) 0.183 (0.068) 8 0.444 0.883 0.222 J 1000 41 50.00 2.14 1.57 1.44 0.331 (0.020) 0.231 (0.066) 16 0.390 0.920 0.220 Mean 44.6 51.43 2.23 1.64 1.41 0.317 (0.006) 0.219 (0.020) 18.4 0.419 0.720 0.339 aAbbreviations: P, percentage of polymorphic loci; AP, mean number of alleles per polymorphic locus; A, mean number of alleles per locus; A , effective number of alleles per locus; H , observed heterozygosity; H , Hardy-Weinberg expected heterozygosity or genetic e o e diversity; G, number of multilocus genotypes; D , multilocus genotypic diversity indices; G/N, number of genotypes per population G samples (proportion of distinguishable genotypes per population); PG, probability of the most common multilocus genotype.
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