Structure of allozyme variation in Nordic Silene nutans (Caryophyllaceae): population size, geographical position and immigration history

We investigated allozyme variation in 34 populations of the perennial herb Silene nutans from Sweden and northern Finland, areas that were ice-covered during the last (Weichselian) glaciation. The present geographical structure of genetic variation in S. nutans in Sweden and northern Finland appears to have been mainly shaped by ancient historical processes. Patterns of variation in allele frequencies suggest two major postglacial immigration routes into Sweden, with populations entering the area from both the south and the east and forming a contact zone with admixed populations in central Sweden. While estimates of within-population genetic diversity and allelic richness are significantly correlated with present population size and geographical position (latitude), population size is not correlated with latitude. Low genetic diversity in the northern populations is more likely to have resulted from ancient stochastic events during the process of immigration than from recent population fragmentation. F _IS values are high and increase with latitude. Evidence of recent bottlenecks was detected in several southern Swedish populations: these can be interpreted in terms of population fragmentation as a result of anthropogenic disturbance. Soil pH is uncorrelated with population size and position.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 357–371.     

Fine-scale spatial genetic structure and gene dispersal in Silene latifolia

Plants are sessile organisms, often characterized by limited dispersal. Seeds and pollen are the critical stages for gene flow. Here we investigate spatial genetic structure, gene dispersal and the relative contribution of pollen vs seed in the movement of genes in a stable metapopulation of the white campion Silene latifolia within its native range. This short-lived perennial plant is dioecious, has gravity-dispersed seeds and moth-mediated pollination. Direct measures of pollen dispersal suggested that large populations receive more pollen than small isolated populations and that most gene flow occurs within tens of meters. However, these studies were performed in the newly colonized range (North America) where the specialist pollinator is absent. In the native range (Europe), gene dispersal could fall on a different spatial scale. We genotyped 258 individuals from large and small (15) subpopulations along a 60 km, elongated metapopulation in Europe using six highly variable microsatellite markers, two X-linked and four autosomal. We found substantial genetic differentiation among subpopulations (global F_ST=0.11) and a general pattern of isolation by distance over the whole sampled area. Spatial autocorrelation revealed high relatedness among neighboring individuals over hundreds of meters. Estimates of gene dispersal revealed gene flow at the scale of tens of meters (5–30 m), similar to the newly colonized range. Contrary to expectations, estimates of dispersal based on X and autosomal markers showed very similar ranges, suggesting similar levels of pollen and seed dispersal. This may be explained by stochastic events of extensive seed dispersal in this area and limited pollen dispersal.     

Climate and clinal variation in seed morphology of the white campion, Silene latifolia (Caryophyllaceae)

Silene latifolia (white campion) is widespread in W Eurasia and shows clinal variation in seed morphology. Two extensive parapatric races, within which seed morphology changes gradually with geographic distance, are separated by a transition zone where the morphological gradient is steeper. A sample of seeds from across the species' native range was scored for morphological characters and analysed using detrended correspondence analysis ordination and multiple regression, in order to simplify mapping of the variation pattern and to investigate statistical relationships with climatic variables. The transition zone, as defined by ordination scores, follows the convolutions of the 0C isotherm for January mean temperature along its entire length from N Europe to the Middle East. January temperature proved to be a highly significant predictor of seed morphology even after the removal of simple spatial trends. Although the origins of comparable large-scale clines have often been interpreted in historical terms, the result reported here supports a role for climatic selection in maintaining the existing variation pattern.     

Numerical analysis of infraspecific variation in European Silene alba and S. dioica(Caryophyllaceae)

Infraspecific variation among European populations of Silene alba and S. dioica was analysed by non-metric multidimensional scaling, which was applied to K-dissimilarity matrices based either on 16 seed characters or on 34 seed, flower and capsule characters. The material used for analysis consisted of 167 collections of wild seed, largely obtained from European botanic gardens, and 97 sets of mature greenhouse-grown progeny. S. alba was found to show topoclinal variation, primarily along an axis from the Iberian peninsula through central Europe to Romania. This pattern is not reflected in current taxonomic treatments. N. and N.W. European S. alba is morphologically heterogeneous and has probably been introduced from various southern sources. S. dioica shows less total variation between populations than S. alba , and consists of local ecogeographic races that intergrade at their margins. The two species are completely discriminated by seed, flower and capsule characters when used in combination, but S. alba populations from the eastern end of the cline closely approach S. dioica in seed morphology. This convergence is unlikely to be an effect of recent introgression.     

A comment on García et al. (2005, 2007) and related papers on mating patterns and gene dispersal in Prunus mahaleb

In two studies on mating patterns and spatial components of pollen and seed dispersal of Prunus mahaleb based on parentage analysis, García et al. (2005, 2007) depicted their 196 focal trees as a spatially isolated population where all reproductive trees had been genotyped. Additional distributional data for P. mahaleb trees in their study area, however, revealed that García and colleagues’ depiction of their study system bears little resemblance to reality. The trees these authors studied did not form a discrete, geographically isolated population. Around 300 ungenotyped reproductive trees occurred within the 1.5‐km distributional gap to the nearest population proclaimed by García and colleagues. Since exhaustive sampling of potential parental genotypes is essential in parentage analyses, the occurrence of a large number of ungenotyped trees in the immediate neighbourhood of focal trees can severely affect the main conclusions of García et al. (2005, 2007) as well as of several related publications on gene dispersal and mating patterns of P. mahaleb conducted on the same trees and relying on the same false premises of spatial isolation and exhaustive sampling.     

Allozyme and shell variation in two marine snails (Littorina, Prosobranchia) with different dispersal abilities

As adults Littorina littorea and L. saxatilis have overlapping distributions, both on the shore and on a geographical scale, this implies that they are affected by similar selective forces to a large extent. The two species have, however, different reproductive strategies. Littorina littorea has a planktonic larval stage lasting 4–6 weeks, whereas L. saxatilis has direct development. This influences dispersal rates of the two species, and gene flow and effective population size are both assumed to be much smaller in L. saxatilis than in L. littorea. Intraspeciflc variation in morphological and allozyme characters differs between the two species. Littorina littorea is homogeneous for both types of characters, both within and between populations, while populations of L. saxatilis may show pronounced differences in shell characters with micro-environment changes, and allozyme variation is markedly higher in this species too. The polymorphism in shell characters of L. saxatilis and the monomorphism of L. littorea, may be expected from theory as a consequence of the different dispersal strategies. Also the lack of divergence between geographically separate populations of L. littorea, but the high degree of between population differentiation in L. saxatilis, could be attributed to the high and low dispersal rates, respectively. However, the larger allozyme variation within subpopulations of L. saxatilis compared to that of L. littorea, is surprising as the former species has a smaller effective population size than the latter. Furthermore, there is no correlation between dispersal ability and overall allozyme heterozygosity, when 10 additional species of Littorina are included in the comparison. It is suggested that different evolutionary backgrounds of different species will possibly explain this, as neither selectionist nor neutralist models alone account for the observation. That is, an earlier passage through a population bottleneck may, for example, influence the genetic variation for a long period of time.     

Mosaic variation in allozyme and plastid DNA markers in the European ranges of Silene vulgaris and its partially sympatric relative S. uniflora (Caryophyllaceae)

The perennial herbs Silene vulgaris and S. uniflora are closely related, partially sympatric and interfertile, yet morphologically distinct. We used nuclear (allozyme) and plastid (polymerase chain reaction–restriction fragment length polymorphism) DNA markers to investigate whether these species have a shared history of postglacial colonization and hybridization in Europe, as inferred from large‐scale patterns of geographic variation. The majority of plastid haplotypes and allozyme alleles were widespread and patchily distributed within both species and there was no geographic structure in the distributions of shared allozymes or haplotypes. The mosaic variation is consistent with a scenario in which repeated episodes of interspecific hybridization pre‐dated the largely allopatric range expansion of the two species during the postglacial period. Our overall results are not consistent with a scenario of extensive hybridization and introgression during the postglacial range expansion of the species or within their current areas of sympatry, but we found some evidence for local, postglacial evolution and hybridization in the Baltic region. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 127–148.     

The spatial distribution of chloroplast DNA and allozyme polymorphisms within A population of Silene alba (Caryophyllaceae)

Genetic structure within a population of Silene alba was studied using a chloroplast DNA (cpDNA) and six allozyme polymorphisms. A 20 × 65 m area was sampled by constructing a 5 × 5 m grid and determining the genotype of the plant nearest to each grid node. Analysis of the spatial distribution of genotypes by Moran's I and join-counts showed a significant degree of association of like cpDNA haplotypes (I = 0.52, S.N.D. = 2.55) but a random or slightly repulsed distribution of allozyme genotypes. A second sample was taken by collecting all individuals from within a 1-m wide transect established along the perimeter of the grid. Genotype and allele frequencies were calculated by grouping individuals from arbitrary 3-m intervals along the transect, and analyzed by Wright's F statistics. The F_st value calculated for cpDNA (0.875) differed markedly from that based on all allozymes (0.027). Taken together, the results suggest that in this population gene flow in the maternally inherited cpDNA is limited by restricted seed dispersal, whereas gene flow in the nuclear DNA based allozymes is more pervasive owing to the added effects of pollen dispersal. The utility of cpDNA polymorphism for the study of fine-scale gene flow is discussed.     

Enzyme polymorphism, morphometric variation and population structure in a restricted endemic, Silene diclinis (Caryophyllaceae)

Silene diclinis is a dioecious perennial herb, restricted to a small area of the SE Spanish province of Valencia. Polyacrylamide gel electrophoresis and morphometric analysis were carried out on cultivated material derived from seed collected from the main population of S. diclinis at Játiva. Only two out of 26 inferred enzyme loci showed substantial variation. There was some indication of population subdivision in both. The esterase locus also showed homozygote excess within the population as a whole and within one subpopulation. Morphological characters showed significant differentiation between subpopulations.
Low allozyme variability characterizes populations of most endemic species that have been investigated. Isolation and historical events may have been as important as population size or edaphic specialization in causing this low variability. But S. diclinis resembles other outbreeding plants in showing population structure and homozygote excess, suggesting restricted within-population gene flow. Mechanisms that may restrict gene flow in S. diclinis include environmentally-induced differences in flowering time, poor seed dispersal and possibly short pollinator foraging distances. Morphological variability and the incidence of developmental abnormalities are speculatively linked with homozygosity. Storage of genetic diversity in space can occur in restricted as well as widespread species and sampling for gene banking should then be spatially representative. Silene diclinis is threatened by continued habitat loss and consequent genetic erosion; its long-term future may depend on the gene bank and the botanic garden.     

Genetic differentiation in the Bladder campions, Silene vulgaris and S. uniflora (Caryophyllaceae), in Sweden

Allozyme variation was studied in Swedish populations of Silene vulgaris (a widespread weed), S. uniflora ssp. uniflora (restricted to coastal habitats) and S. uniflora ssp. petraea (endemic to Sweden and confined to open limestone habitats). The taxa are diploid, gynodioecious, perennial herbs and showed high levels of within-taxon and within-population gene diversity at four polymorphic loci. Within-taxon diversity was highest (H_tax=0.52) in S. vulgaris and lowest (H_tax=0.36) in S. uniflora ssp. uniflora. The weedy S. vulgaris has more alleles than either of the other two taxa and 5 out of a total of 27 alleles are unique to S. vulgaris. Most of the gene diversity within each of the taxa is accounted for by within-population diversity. The between-population component of diversity is 10% in S. vulgaris , and 24% and 5%, respectively, in S. uniflora ssp. uniflora and ssp. petraea. Hybrids may occur between S. vulgaris and S. uniflora , but introgression is limited by the species' ecology. Neither allozyme nor distributional data support the suggestion that ssp. petraea is a recent hybrid between S. vulgaris and S. uniflora ssp. uniflora , although an older hybrid origin for ssp. petraea is possible. Patterns of allele frequency variation suggest that there has been some historical gene flow between taxa, outside their present areas of sympatry. It is likely that the two subspecies of S. uniflora , which occur in naturally open habitats, were able to colonize Sweden during the Late Glacial or early post-glacial, whereas S. vulgaris followed the spread of agriculture into Sweden.     

Identification and properties of UDP-glucose: Cyanidin-3-O-glucosyltransferase isolated from petals of the red campion (Silene dioica)

An enzyme catalyzing the transfer of the glucosyl moiety of UDP-glucose to the 3-hydroxyl group of cyanidin has been demonstrated in petal extracts of Silene dioica mutants with cyanidin-3-O-glucoside in the petals. This transferase activity was also present in young rosette leaves and calyces of these plants. The highest glucosyltransferase activity was found in petals of opening flowers of young plants. The enzyme was purified ninetyfold by PVP and Sephadex chromatography. The glucosyltransferase had a pH optimum of 7.5, had a “true K_m value” of 4.1×10^?4 m for UDP-glucose and 0.4×10^?4 m for cyanidin chloride, and was not stimulated by divalent metal ions. Both p-chloromercuribenzoate and HgCl₂ inhibited the enzyme activity. Pelargonidin chloride and delphinidin chloride at reduced rates also served as substrates. The enzyme did not catalyze the glucosylation of the 3-hydroxyl group of flavonols or the 5-hydroxyl group of anthocyanins. ADP-glucose could not serve as a glucosyl donor. The results of Sephadex G150 chromatography suggest that the glucosyltransferase can exist as dimer of about 125,000 daltons and as active monomers of 60,000 daltons. The genetic control of the glucosyltransferase activity is discussed.     

Demographic and genetic connectivity: the role and consequences of reproduction,dispersal and recruitment in seagrasses

Accurate estimation of connectivity among populations is fundamental for determining the drivers of population resilience, genetic diversity, adaptation and speciation. However the separation and quantification of contemporary versus historical connectivity remains a major challenge. This review focuses on marine angiosperms, seagrasses, that are fundamental to the health and productivity of temperate and tropical coastal marine environments globally. Our objective is to understand better the role of sexual reproduction and recruitment in influencing demographic and genetic connectivity among seagrass populations through an integrated multidisciplinary assessment of our present ecological, genetic, and demographic understanding, with hydrodynamic modelling of transport. We investigate (i) the demographic consequences of sexual reproduction, dispersal and recruitment in seagrasses, (ii) contemporary transport of seagrass pollen, fruits and seed, and vegetative fragments with a focus on hydrodynamic and particle transport models, and (iii) contemporary genetic connectivity among seagrass meadows as inferred through the application of genetic markers. New approaches are reviewed, followed by a summary outlining future directions for research: integrating seascape genetic approaches; incorporating hydrodynamic modelling for dispersal of pollen, seeds and vegetative fragments; integrating studies across broader geographic ranges; and incorporating non‐equilibrium modelling. These approaches will lead to a more integrated understanding of the role of contemporary dispersal and recruitment in the persistence and evolution of seagrasses.     

The structure of morphometric and allozyme variation in relict populations of Gypsophila fastigiata (Caryophyllaceae) in Sweden

Patterns of variation and the structuring of diversity in seed phenotype and allozymes were investigated in Sweden Gypsophila fastigiata, a perennial herb with a disjunct ‘Late Glacial relict’ distribution in eastern and northern Europe. The overall patterns of variation in allozymes and seed morphology are congruent and are significantly correlated with geographic distance. However, most of the congruence between the distance matrices based on allozymes and seed morphology is attributable to regional differentiation between the isolated Öland and Dalarna metapopulations. On a local scale, within the two metapopulations, seed shape variation is partially correlated with geography whereas allozyme variation is unrelated to the geographic disposition of the sampling sites. Despite the fact that regional metapopulations can be discriminated on the basis of seed shape and allozyme frequencies, relatively little of the total diversity in seed morphology and allozymes is due to differentiation between regions. Partitioning of diversity into its within- and among-regional, site and individual components, showed that the majority of diversity (52% for seed shape and 91% for allozymes) is stored within sites. Seventy-three percent of the variation in allozymes is due to variation within individuals (i.e. heterozygosity) whereas phenotypic variation in seed shape within individuals is low (0.4% of the total diversity). The events that led to the large scale disjunction of the Late Glacial distribution of G. fastigiata may have shaped the pattern of differentiation observed among the regional isolates of the species. However, the species' history of local population disjunction during post-glacial and historic times has not had a substantial impact on the spatial organization of allozyme and morphometric diversity within regions. Extensive local gene flow may have allowed the regional metapopulations to have functioned as extended effective populations–on a time scale of tens or hundreds of years–and may have hindered the loss of within-site diversity.     

Paternity analysis of the breeding structure of strangler fig populations: evidence for substantial long-distance wasp dispersal

Abstract. The mutualistic interaction of figs with their species-specific wasp pollinators and the role of figs as 'keystone' plant resources in tropical communities has received substantial attention from both plant and animal ecologists. Despite this focus on the reproductive biology of figs, the minute size of the wasps has effectively precluded our ability to monitor patterns of wasp dispersal and fig mating relationships in natural forest habitats. In this paper we use genetic markers and genealogy reconstruction techniques to examine the breeding structure of populations of four strangler fig species occurring in central Panama. The natural history of figs facilitates the genetic analysis of full-sib progeny arrays from which the genotypes of successful pollen donors can be reconstructed precisely. Paternity reconstruction in the four study species reveals that individual flowering trees may routinely receive pollen from numerous donors despite characteristically low densities of co-flowering individuals. These data indicate not only that breeding populations of these figs are larger than the minimum critical sizes predicted to be necessary to support populations of their species-specific pollinators, but are more extensive in size and area than has been described for any plant species yet examined. Further, the fig wasps are shown to be efficient agents of long-distance dispersal, routinely moving up to 10 km between flowering trees. In accord with the potential for substantial long-distance gene flow and large effective population sizes, ten of eleven species of Panamanian figs assayed were found to maintain exceptionally high levels of genetic variation within their populations. Combined with other reports of occasional long-distance dispersal, the results of this study suggest that fig wasps may be more effective at colonizing isolated fig populations than previously thought.     

Sex expression and fruit set in Silene littorea (Caryophyllaceae): variation among populations

The flowers of Silene littorea may be female or hermaphrodite, and an individual may show any of three sexual phenotypes: female-flowers-only, hermaphroditeflowers-only, or mixed-type. We estimated the relative frequencies and fruit sets of each flower type and each sexual phenotype in eleven S. littorea populations in northwest Spain. For all plants and sites combined, 18% of flowers were female and 82% hermaphrodite. Individuals either had only female flowers (6%), only hermaphrodite flowers (43%), or had both flower types (51%). The relative frequencies of female and hermaphrodite flowers, and of sexual phenotypes, vaned among populations. There was no significant effect of flower sex on its probability of setting fruit. Differences among individual plants within populations had the most prominent role in explaining variation among flowers in probability of setting fruit, while population and its interaction with flower sex had a negligible and nonsignificant influence. Mean fruit set per plant did not differ significantly between sexual phenotypes. Neither population nor its interaction with sexual phenotype accounted for significant amounts of between-plant variance in fruit set.     

Historical and contemporary gene dispersal in wild carrot (Daucus carota ssp. carota) populations

Background and Aims

Wild carrot is the ancestor of cultivated carrot and is the most important gene pool for carrot breeding. Transgenic carrot may be released into the environment in the future. The aim of the present study was to determine how far a gene can disperse in wild carrot populations, facilitating risk assessment and management of transgene introgression from cultivated to wild carrots and helping to design sampling strategies for germplasm collections.

Methods

Wild carrots were sampled from Meijendel and Alkmaar in The Netherlands and genotyped with 12 microsatellite markers. Spatial autocorrelation analyses were used to detect spatial genetic structures (SGSs). Historical gene dispersal estimates were based on an isolation by distance model. Mating system and contemporary pollen dispersal were estimated using 437 offspring of 20 mothers with different spatial distances and a correlated paternity analysis in the Meijendel population.

Key Results

Significant SGSs are found in both populations and they are not significantly different from each other. Combined SGS analysis indicated significant positive genetic correlations up to 27 m. Historical gene dispersal σ_g and neighbourhood size N_b were estimated to be 4–12 m [95 % confidence interval (CI): 3–25] and 42–73 plants (95 % CI: 28–322) in Meijendel and 10–31 m (95 % CI: 7–∞) and 57–198 plants (95 % CI: 28–∞) in Alkmaar with longer gene dispersal in lower density populations. Contemporary pollen dispersal follows a fat-tailed exponential-power distribution, implying pollen of wild carrots could be dispersed by insects over long distance. The estimated outcrossing rate was 96 %.

Conclusions

SGSs in wild carrots may be the result of high outcrossing, restricted seed dispersal and long-distance pollen dispersal. High outcrossing and long-distance pollen dispersal suggest high frequency of transgene flow might occur from cultivated to wild carrots and that they could easily spread within and between populations.