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A Comparison of the Extent of Genetic Variation in the Endangered Sagittaria Natans and Its Widespread Congener S

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A Comparison of the Extent of Genetic Variation in the Endangered Sagittaria Natans and Its Widespread Congener S http://www.paper.edu.cn Aquatic Botany 87 (2007) 1–6 www.elsevier.com/locate/aquabot A comparison of the extent of genetic variation in the endangered Sagittaria natans and its widespread congener S. trifolia Jin-Ming Chen a, Wahiti Robert Gituru b, Qing-Feng Wang a,* a Laboratory of Plant Systematics and Evolutionary Biology, College of Life Sciences, Wuhan University, Wuhan 430072, Hubei, PR China b Botany Department, Jomo Kenyatta University of Agriculture and Technology, P.O. Box 62000-00200, Nairobi, Kenya Received 13 May 2006; received in revised form 28 November 2006; accepted 1 December 2006 Abstract Genetic variation and clonal diversity of 14 populations of the endangered clonal herb Sagittaria natans and its widespread congener S. trifolia were investigated using inter-simple sequence repeat (ISSR) markers. Using nine effective ISSR primers, a total of 92 DNA fragments were generated with 54 (percentage of polymorphic loci, PPL: 58.7%) being polymorphic. A higher level of genetic diversity among populations was found in S. natans (PPL: 48.9%) than S. trifolia (PPL: 32.6%). Analysis of molecular variance (AMOVA) showed that in each species a similar proportion of the total genetic variation resided within and among populations, and that between species there was a moderate genetic differentiation (Gst: 0.601). With the use of 54 polymorphic ISSR markers, we identified 116 genets among 138 samples from five S. natans populations, and 93 genets among 215 samples from nine S. trifolia populations. The proportion of distinguishable genets (PD: mean 0.82) and Simpson’s diversity index (D: mean 0.95) for S. natans exhibited higher degree of clonal diversity compared to S. trifolia (PD: mean 0.42; D: mean 0.89). Sexual reproduction might have been played an important role in maintaining and increasing the clonal diversity in both species. Recent and on-going decimation of S. natans populations in the region appear not to have had a major impact on genetic diversity in this rare plant. # 2007 Published by Elsevier B.V. Keywords: Clonal structure; Genetic diversity; ISSR; Population differentiation; Sagittaria 1. Introduction one potential limitation of these studies is that the included taxa are treated as independent samples, ignoring their phylogenetic As more species dwindle toward extinction, the rare and relatedness (Gitzendanner and Soltis, 2000). As a result, narrowly distributed species have in recent years received more growing interest in population genetic and conservation biology attention especially with regard to their viability (Gitzendanner has directed attention to comparisons of widespread and and Soltis, 2000). The survival of a species depends not only on restricted congeneric species (Karron et al., 1988; Soltis and habitat and geographic factors but is ultimately linked to the Soltis, 1991; Lewis and Crawford, 1995; Ge et al., 1999; He genetic variation available to a species (Dodd and Helenurm, et al., 2000; Dodd and Helenurm, 2002; Park, 2004). Closely 2002), thus knowledge of the genetic aspects of rarity within related species are likely to share many life-history features due rare and endangered species is critical to their conservation and to recent common ancestry, making it easer to identify the management (Schaal et al., 1991; Hamrick and Godt, 1996). causes of differences in genetic diversity (Dodd and Helenurm, Several previous surveys of genetic variation in plants have 2002). Recent comparative studies of genetic variation between shown that rare, endemic, or narrowly distributed plants tend to rare and widespread species have demonstrated that several rare maintain low degree of genetic variability due to the impact of species are as polymorphic as their widespread congeners genetic drift, the founder effect, and directional selection with (Gitzendanner and Soltis, 2000; Dodd and Helenurm, 2002). high levels of inbreeding (Franklin, 1980; Karron, 1987; Thus, it is difficult to make generalizations that species with Hamrick and Godt, 1989; Ellstrand and Elam, 1993). However, limited geographic range always have low genetic diversity. Sagittaria trifolia is a perennial, erect marsh herb that belongs to the aquatic family Alismataceae. Sagittaria trifolia * Corresponding author. Tel.: +86 27 68752869; fax: +86 27 68752560. is one of the most widespread species in the genus Sagittaria E-mail address: [email protected] (Q.-F. Wang). ranging from north Beikal in Russia to Southeast Asia and 0304-3770/$ – see front matter # 2007 Published by Elsevier B.V. doi:10.1016/j.aquabot.2006.12.001 转载 中国科技论文在线 http://www.paper.edu.cn 2 J.-M. Chen et al. / Aquatic Botany 87 (2007) 1–6 Southeast Europe (Chen, 1989). Sagittaria natans is a float- Table 1 leaved plant species and is confined to bogs, rivers, and streams Locations of the populations of S. natans and S. trifolia sampled in this study in Northeast Europe and Northeast Asia (Chen, 1989). Liaoning Populations Location Latitude (N)/Longitude (E) Province of China is the southern-most location of S. natans.In S. natans China S. natans has been reported from Liaoning, Jilin, and EKH Erkahe, Inner Mongolia 49830009.700/117845010.000 Helongjiang provinces and also in Inner Mongolia (Chen, SQ Shiquan, Helongjiang 4983201400/120840028.900 1989). However, in our recent field investigation, only five DH Dunhua, Jilin 43820003.900/128813022.800 0 00 0 00 natural populations including one population in Jilin Province, WSL Wusuli, Helongjiang 46848 32.0 /134800 54.0 LHP Lianhuapao, Helongjiang 46844040.800/134801023.000 one population in Inner Mongolia and three populations in Helongjiang Province were found in China. Sagittaria natans is S. trifolia LDS Liudingshan, Jilin 43819026.900/128813050.600 now considered to be rare and threatened or endangered in MS Mishan, Helongjiang 45830040.000/131851020.000 China and is listed among the second category of the key HL Hulin, Jilin 45847035.000/133835045.300 protected wild plants (Yu, 1999). Both S. natans and S. trifolia MJG Majiagang, Helongjiang 45827004.000/132816053.400 are self-compatible species, which can reproduce both sexually XKH Xingkaihu, Helongjiang 45827004.000/13281604000 0 00 0 00 by selfed and out-crossed seeds and vegetatively through corms BC Bacha, Helongjiang 47825 47.0 /133818 32.0 TJ Tongjiang, Helongjiang 48820056.400/134824059.400 (Chen, 1989). Phylogenetic studies of Sagittaria species using RH Raohe, Helongjiang 46844038.800/134801022.000 morphological and molecular data found that S. trifolia and S. DJC Dongjingcheng, Helongjiang 44806054.000/129811024.000 natans are sister species (Chen, 1989; Chen and Wang, unpublished data). In recent years, inter-simple sequence repeats (ISSRs) have powdered in liquid nitrogen, mixed with 2 mL extraction buffer been successfully employed to reveal genetic diversity and [1.4 M NaCl, 100 mM Tris–HCl (pH 8.0), 20 mM EDTA, 2% identify the different clones in populations of clonal plants (V/V) CTAB, and 2% 2-mercaptoethanol] at 65 8C, and (Esselman et al., 1999; Li and Ge, 2001; Chen et al., 2006). The incubated at 65 8C for 30 min with gentle shaking every 5 min. principal goal of this study was to (1) investigate and compare the Proteins were extracted twice with 2 mL of chloroform: amount and distribution of genetic diversity in the endangered isoamylalcohol (24:1), then centrifuged at 10,000 Â g for species S. natans with its widespread congener S. trifolia using 2 min. RNase (10 mgmLÀ1) was added to the supernatant and ISSR markers, (2) to genetically identify S. trifolia and S. natans incubated for 2 h at 37 8C. The mixture was centrifuged at clones as well as to estimate the genetic diversity of these clones. 10,000 Â g for 2 min. The sediment was washed twice in 70% This information was intended to contribute to a better ethanol, air-dried, and resuspended in 100 mL 0.1 Â TE, and understanding of the causes of rarity of S. natans. then stored at À20 8C. 2. Materials and methods 2.3. ISSR PCR amplification 2.1. Plant materials Reactions were carried out in a volume of 25 mL containing 0.25 mM of each dNTP, 2.5 mLof10Â Taq buffer [10 mM The plant specimens of S. natans used in this investigation Tris–HCl (pH 8.3), 1.5 mM MgCl2 and 50 mM KCl], 1 mM were obtained from five populations throughout the natural primer, 1 U Taq Polymerase (Tian Yuan Biotech) and 40 ng of range of the species in China. In each of the five populations, a DNA template. Amplification of genomic DNA was made on a sample of 19 or more plants was taken. A total of 138 PTC-100TM thermocycler (MJ Research, Inc.), and commenced individuals of S. natans from the five populations were included with 4 min at 94 8C, followed by 35 cycles of 1 min at 94 8C, in the study. For S. trifolia, a total of 215 individuals from nine 1 min annealing at 55 8C and 2 min extension at 72 8C, and a populations (18 or more individuals were sampled in each final extension cycle of 7 min at 72 8C. Amplification products population) in Northeast China that were in sympatric to S. were resolved electrophoretically on 1.5% (W/V) agarose gels natans populations were sampled. Details on collection site are run at 100 V in 0.5Â Tris–boric acid–EDTA (TBE), visualized given in Table 1. About 5 g of fresh leaves was harvested from by staining with ethidium bromide, and photographed under each plant and immediately dried in a ziplock plastic bag ultraviolet light. Sixty-five ISSR primers (SBS Genetech. Co. containing about 70 g of silica gel. To estimate genetic Ltd., Shanghai, China) were screened on eight randomly variability within populations we only used plants sampled at selected individuals (four samples of each species). The eight least at 3-m distances from each other.
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