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Saxifraga Oppositifolia) in the High Arctic Archipelago of Svalbard

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Saxifraga Oppositifolia) in the High Arctic Archipelago of Svalbard Research Article Population genetics of purple saxifrage (Saxifraga oppositifolia) in the high Arctic archipelago of Svalbard Maria Pietila¨inen and Helena Korpelainen* Department of Agricultural Sciences, University of Helsinki, PO Box 27 (Latokartanonkaari 5), Helsinki FI-00014, Finland Downloaded from Received: 20 February 2013; Accepted: 18 March 2013; Published: 8 April 2013 Citation: Pietila¨inen M, Korpelainen H. 2013. Population genetics of purple saxifrage (Saxifraga oppositifolia) in the high Arctic archipelago of Svalbard. AoB PLANTS 5: plt024; doi:10.1093/aobpla/plt024 Abstract. We investigated patterns of genetic variability in Saxifraga oppositifolia in the isolated Arctic Svalbard http://aobpla.oxfordjournals.org/ archipelago. The genetic analysis included genotyping using nine polymorphic microsatellite markers and sequen- cing of the nuclear internal transcribed spacer region. Among populations, mean allele numbers per microsatellite locus ranged from 2.0 to 2.6, and 9 % of alleles were unique. Observed (HO) and expected (HE) heterozygosities aver- aged 0.522 and 0.445, respectively. Typically negative but non-significant FIS values (mean 20.173) were found in S. oppositifolia populations. FST values were relatively low (mean 0.123). The Bayesian structure analysis provided additional information on population genetic structures. Seven out of 11 studied populations, including populations located both near each other and far apart (distances 5–210 km), showed relatively homogeneous clustering pat- terns, while one population located on a slope in the main settlement of Longyearbyen possessed a unique genetic structure. The Mantel test proved that there is no significant correlation between genetic and geographical dis- at Helsinki University Library on May 17, 2016 tances. Different growth habits (compact, trailing and intermediate) did not possess distinct genetic compositions based on microsatellite variation. Internal transcribed spacer sequencing revealed 12 polymorphic sites. Among 24 sequenced Svalbard samples, eight haplotypes were detected, none shared by the mainland samples. Population genetic structures of S. oppositifolia in Svalbard show that both genetic variation and differentiation levels are modest, outcrossing is the main mating system, and dispersal and gene flow are important, probably attributable to strong winds and human and animal vectors. Keywords: Arctic; ITS sequencing; microsatellites; population genetic structure; Saxifraga oppositifolia. ecological amplitude. It is well established that biological Introduction systems are dynamic: genetic variation enables adapta- In natural environments, organisms are typically exposed tion through selection, while in small and isolated popula- to several stress factors simultaneously, and the stresses tions random evolutionary processes, such as genetic are often most pronounced in extreme habitats, such as drift, may become strong. The consequences of evolution- those at high latitudes and altitudes. For instance, Arctic ary actions will be visible in the pattern of genetic diversity plants may encounter many unique environmental and differentiation of populations. factors, such as drought, permafrost, nutrient leaching, Purple saxifrage, Saxifraga oppositifolia, is an Arctic– cryoturbation, extreme temperatures and short growing Alpine early flowering perennial herb. It is a circumpolar, seasons (Ko¨rner 2003). The extent to which an organism ecologically and morphologically variable species with a is able to deal with stress determines the limits of its wide range of habitats. It probably has the widest global * Corresponding author’s e-mail address: helena.korpelainen@helsinki.fi Published by Oxford University Press on behalf of the Annals of Botany Company. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. AoB PLANTS www.aobplants.oxfordjournals.org & The Authors 2013 1 Pietila¨inen and Korpelainen — Population genetics of purple saxifrage distribution in the family Saxifragaceae (Webb and In addition, we compared nuclear ITS sequence variation Gornall 1989). Saxifraga oppositifolia grows in dry to of plants from Svalbard and more southern mainland moist soil and from sea level up to 4500 m in the Alps, regions of distribution in order to reveal relationships including the coldest known places with angiosperm among plants across a wider geographic area. plant life (Ko¨rner 2011). It is also mentioned as the northernmost vascular plant species, having been Methods found up in northern Greenland at 83815′ (Gjaervoll and Ro¨nning 1999). Saxifraga oppositifolia is not endan- Sampling, morphological observations and DNA gered at the moment, but climate change, the potential analyses warming or drying of northern areas, and increased UV We sampled fresh leaf material from 11 populations in radiation could become a threat in the future (Ko¨rner late June 2005 (except population NYB in late June 2003). It is self-compatible but mainly outcrossing and 2010) in the Arctic Svalbard archipelago in locations depends on its pollinators, which are mainly bumblebees between latitudes 78 and 808N (Table 1; Fig. 1). The Downloaded from (Bombus sp.) (Stenstro¨m and Molau 1992; Stenstro¨m and growth form of the sampled plants was classified as Bergman 1998). However, there are no bumblebees in compact, trailing (creeping) or intermediate. Pairwise some Arctic regions, such as Svalbard, where the pollin- distances between populations varied from 5 to ating insects are probably mainly small insects of the 210 km. The collected leaf samples were desiccated in order Diptera (Coulson et al. 2003). silica gel and stored at 280 8C until DNA extraction, http://aobpla.oxfordjournals.org/ Previous population genetic and phylogeographic ana- which was conducted using commercial kits (DNeasy lyses on S. oppositifolia include random amplification of Plant Mini Kit, Qiagen Inc., and E.Z.N.A. Plant DNA Mini- polymorphic DNA studies by Gabrielsen et al. (1997) and prep Kit, Omega Bio-tek, Inc.) following the manufac- Gugerli et al. (1999), amplified fragment length poly- turers’ instructions. morphism (AFLP) studies by Alsos et al. (2007), Kropf We determined the population genetic characteristics et al. (2008), Mu¨ller et al. (2012) and Winkler et al. of the populations using the following nine polymorphic (2012), restriction fragment length polymorphism investi- microsatellite markers developed for S. oppositifolia (Pie- gations (Abbott et al. 1995; Abbott and Comes 2004) and tila¨inen and Korpelainen 2010): SaxJC, SaxS1C, SaxS2C, studies based on cpDNA and nuclear internal transcribed SaxT9C, SaxT10C, S1_2, S4_2, S8_2 and S21_2. In geno- at Helsinki University Library on May 17, 2016 spacer (ITS) sequences (Holderegger and Abbott 2003; typing, one of the primers in each primer pair was fluor- Winkler et al. 2012). All these DNA studies have used uni- escently (FAM, HEX or TET) labelled. Amplifications were versal binary markers or sequence data. On the other performed in 5-mL reactions, containing 0.5 mL of DNA hand, our study utilizes microsatellite markers that we (about 5 ng), 2.75 mL of ddH2O, 0.5 mLof10× buffer, recently developed for S. oppositifolia (Pietila¨inen and 0.1 mL of dNTP mix (10 mM), 0.15 mL of DyNAzyme II Korpelainen 2010), which, besides allowing precise DNA polymerase (Finnzymes) (2 U mL21) and 0.5 mlof genetic analyses, make it possible to draw conclusions both primers (5 pmol mL21). The polymerase chain reac- on the effects of mating systems on population genetic tions (PCR) were carried out as follows: DNA denatur- structures. ation at 94 8C for 4 min followed by 30 cycles of In this study we aimed to reveal patterns of genetic denaturation at 94 8C for 45 s, annealing at 46–60 8C variability in S. oppositifolia in the isolated Arctic Svalbard (depending on the primer pair) for 45 s, and elongation archipelago. We hypothesized that (i) populations possess at 72 8C for 1 min, with a final elongation at 72 8C for low levels of genetic variation and are genetically differen- 10 min. For genotyping, PCR products were first diluted tiated due to the colonization history and small popula- with ddH2O. Following this, 2 mL of each sample were tion sizes and consequent effects of genetic drift, pipetted into a 96-well plate, and 8 mL of standard (ii) inbreeding and heterozygote deficiency are negligible (MegaBACE ET400-R, GE Healthcare) diluted with due to supposedly prevalent outcrossing, and (iii) ddH2O (1/45) were added. The plates were then run founder dynamics combined with gene flow from with the MegaBACE 1000 DNA Analysis System (Amer- various sources has generated an admixed genetic struc- sham Biosciences). Genotyping results were then ana- ture in populations exposed to considerable human inter- lysed with the MegaBACE Fragment Profiler 1.2 ference when compared with pristine populations. Since (Amersham Biosciences). S. oppositifolia is morphologically variable and presents To complement microsatellite analyses, we sequenced different growth habits (compact, trailing and intermedi- 24 samples from Svalbard (Table 1) and five samples ate; Aiken et al. 2005), even within populations, we also from mainland Norway for the nuclear ITS region using investigated whether such morphological features have primers ITS1 (5′-TCC GTA GGT GAA CCT GCG
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