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A Preliminary Study of Genetic Variation in the Endangered, Chinese Endemic Species Dysosma Versipellis (Berberidaceae)

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A Preliminary Study of Genetic Variation in the Endangered, Chinese Endemic Species Dysosma Versipellis (Berberidaceae) QIUBot. Bull.et al. Acad. — Genetic Sin. (2005) variation 46: 61-69 in Dysosma versipellis 61 A preliminary study of genetic variation in the endangered, Chinese endemic species Dysosma versipellis (Berberidaceae) Ying-Xiong QIU1, Xin-Wen ZHOU1, Cheng-Xin FU1,*, and Yuk-Sing Gilbert CHAN2 1Laboratory of Systematic and Evolutionary Botany, Department of Biology, College of Life Sciences, Zhejiang University, Hangzhou 310029, P. R. China 2Department of Applied Biology and Chemical Technology, The Hong Kong Polytechnic University, Hung Hom, Hong Kong (Received December 23, 2003; Accepted September 30, 2004) Abstract. This study represents a preliminary analysis of allozyme variation in Dysosma versipellis (Berberidaceae), an endangered plant species endemic to China. Five populations of D. versipellis and one population of D. pleiantha were sampled and analyzed using starch gel electrophoresis of nine enzymes that corresponded to nine interpretable loci. Levels of genetic polymorphism within populations (means: P = 15.54%, A = 1.16, He = 0.045) were much smaller than values for seed plants in general (P = 34.2%, A = 1.53, He = 0.113), as well as values for other endemic species (P = 26.3%, A = 1.39, He = 0.063). Mean values for the FST across all D. versipellis populations tended to be high (FST = 0.468). An indirect estimate of the number of migrants per generation (Nm = 0.284) indicated that gene flow is low among populations of D. versipellis. Additionally, analysis of genetic variation revealed a substantial heterozygosity deficiency in all analyzed populations except HB. Genotype frequencies within D. versipellis popu- lations indicate that they may be severely inbred, making inbreeding depression a possible explanation for the low seed set observed in this species. Likewise, the low level of genetic diversity observed within D. versipellis popula- tions suggests that clonal reproduction might be more important than sexual reproduction for D. versipellis. In comparison, genetic variation observed in one population of the closely related species D. pleiantha was much higher than the variation within D. versipellis populations. On the basis of these observations, we suggest that in situ con- servation will be an important and practical measure for maintaining this species. If ex situ conservation is pursued, sampling should cover all populations across the species’ distribution so as to retain as much genetic diversity as possible. Keywords: Allozyme; Asexual reproduction; Dsma versipellis; Endangered species; Genetic variation. Introduction Propagation, therefore, appears to occur mostly by veg- etative means (Ma, 2000; Qiu and Qiu, 2002). In recent Plants in the genus Dysosma (Berberidaceae), with years, natural populations of this species have declined seven species, occur solely in China. Populations of the considerably due to anthropogenic activities like habitat endangered species D. versipellis, an herbaceous peren- destruction and overcollecting for medicinal applications. nial species that grows in the understory of mixed ever- The rhizome of Dysosma has been found to be a source green and deciduous forests (Ying et al., 1993), have a of podophyllotoxin, the active ingredient used as a start- further restricted distribution in the East and South China. ing compound for the chemical synthesis of etoposide Dysosma pleiantha is another rare species in the genus, (VP-16-213) and teniposide (VM-26), effective agents in the but little is known about its biology. However, it is clear treatment of lung cancer, a variety of leukemias, and other that these two species have non-overlapping distributions solid tumor diseases (Jackson and Dewick, 1984, 1985). and differ substantially in both leaf and flower morphol- Nearly all remaining populations of D. versipellis are now ogy (Ying et al., 1993). These observations suggest that located within protected nature reserves, and the species the breeding systems of some Dysosma species may have is classified as endangered in the Chinese Plant Red Book undergone divergent evolution resulting from adaptation (Fu, 1992). Because the species is endangered, and har- to different ecological conditions. Dysosma versipellis re- vesting of rhizomes continues to exceed the rate of natu- produces sexually through cross-pollination and asexually ral regeneration, immediate attention should be given to by spreading rhizomes. Field observations indicates that conserving the species through in situ and/or ex situ although the plants flower almost every year, seed pro- approaches. duction is limited, and seedling establishment is poor. Resource managers responsible for the conservation and recovery of threatened plant species frequently need to *Corresponding author. Tel: +86-571-86971576; Fax: 86-571- make choices regarding which areas are in need of great- 86432273; E-mail: [email protected] est protection, which populations should have priority for 62 Botanical Bulletin of Academia Sinica, Vol. 46, 2005 preservation, what the minimum number of individuals re- mature, plant sends out a shoot in early March, and a ter- quired to avoid inbreeding and to sustain genetic varia- minal cyme emerges on the shoot in April with 5 to 8 droop- tion is, as well as how germplasm should be collected from ing red-purple flowers. Fruits mature in late June (Figure a population to capture representative genetic diversity 1) (Ying et al., 1993; Qiu and Qiu, 2002). for ex situ conservation (Torres et al., 2003). Thus, under- Dysosma versipellis occurs in the undergrowth of sub- standing how genetic variation is distributed among indi- tropical forests between 200 and 3,500 m elevation, and is viduals and populations must be considered together with distributed across Sichuan, Anhui, Hunan, Guanxi, Jianxi evolutionary history, population dynamics, breeding and Hubei Provinces. From 1998 to 2002 several extensive system, and species/population structure in evaluating the collecting trips for D. versipellis were conducted across critical attributes of rare species, and such knowledge is China. The sampling included five fragmented populations of paramount importance for developing recovery plans located across the species entire range. Samples from one that can meaningfully sample and preserve genetic diver- population of D. pleiantha were also collected (Figure 2). sity (Falk and Holsinger, 1991). To our knowledge, this Due to the different sizes of the populations, the number study represents the first attempt to characterize the ge- of samples collected ranged from 12 to 22. The distance netic diversity within and among populations of D. between adjacent samples was at least 10 m to increase versipellis. the likelihood of sampling non-clonal, inter-individual Among the many molecular techniques available for variation within each population. Samples were taken from evaluating plant genetic variation, allozyme electrophore- the edges, as well as the interior of populations. A total of sis is more than adequate to address many conservation 100 young leaf samples were taken from randomly chosen genetics issues. This technique is amenable for conserva- putative individuals. The leaf material was transported in tion genetic surveys because data can be obtained quickly a hand cooler to the laboratory, where it was stored at - for many individuals. The approach involves analysis of 80°C. single-gene codominant molecular markers (loci) that are biparentally inherited and easily assessed by visualizing band patterns and intensities on starch electrophoretic gels. This approach has made it possible to better under- stand the spatial distribution of clones and genetic diver- sity maintained within and among plant populations (e.g., Parker and Hamrick, 1992; Berg and Hamrick, 1994). In the present study, we used enzyme electrophoresis to assess whether patterns of genetic diversity are con- sistent with clonal structure and low levels of sexual re- production observed across D. versipellis populations in the field. Specifically, we addressed the following questions: (1) Are D. versipellis populations typically com- posed of one or multiple genets? (2) What are the levels and distribution of genetic variability within and among populations of D. versipellis? (3) What are the possible factors that might explain the patterns and levels of ge- netic variation observed? Additionally, we present mea- sures of allozyme variation within and among populations of D. versipellis, and compare these to data published for other plant taxa with similar characteristics (Ellstrand and Roose, 1987; Hamrick and Godt, 1990). It is our hope that such information will be used by conservation biologists to formulate effective management strategies based on ge- netic data for the conservation of this endangered species. Materials and Methods Plant Material and Population Sampling Dysosma versipellis is a perennial rhizomatous herb that undergoes both sexual and asexual reproduction. Indi- vidual plants grow from rhizome and typically reach 40 to 150 cm in height, with an unforked stem bearing one or two alternately arranged leaves, which are rounded, cen- trally peltate, and 4-9-lobed with finely dentate margin. Figure 1. Dysosma versipellis. a, rhizome and roots; b, leaf; c, Plants remain in a juvenile phase for 4-5 years. When cyme; d, sepals; e, petal; f, fruit. QIU et al. — Genetic variation in Dysosma versipellis 63 polymorphic loci (P) (95% criterion) per subpopulation, the AH observed heterozygosity (Ho), and the mean expected het- HB erozygosity
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