Evolutionary aspects of population structure for molecular and quantitative traits in the freshwater snail Radix balthica

Detecting the action of selection in natural populations can be achieved using the QST-FST comparison that relies on the estimation of FST with neutral markers, and QST using quantitative traits potentially under selection. QST higher than FST suggests the action of directional selection and thus potential local adaptation. In this article, we apply the QST-FST comparison to four populations of the hermaphroditic freshwater snail Radix balthica located in a floodplain habitat. In contrast to most studies published so far, we did not detect evidence of directional selection for local optima for any of the traits we measured: QST calculated using three different methods was never higher than FST. A strong inbreeding depression was also detected, indicating that outcrossing is probably predominant over selfing in the studied populations. Our results suggest that in this floodplain habitat, local adaptation of R. balthica populations may be hindered by genetic drift, and possibly altered by uneven gene flow linked to flood frequency.     

Life history traits variation in heterogeneous environment: The case of a freshwater snail resistance to pond drying

Ecologists and population geneticists have long suspected that the diversity of living organisms was connected to the structure of their environment. In heterogeneous environments, diversifying selection combined to restricted gene flow may indeed lead to locally adapted populations. The freshwater snail, Galba truncatula, is a good model to address this question because it is present in a heterogeneous environment composed of temporary and permanent waters. In order to test the selective importance of those environments, we proposed here to measure survival of lineages from both habitats during drought episodes. To this purpose, we experimentally submitted adults and juveniles individuals from both habitats to drought. We found a difference in desiccation resistance between temporary and permanents waters only for adults. Adults from temporary habitats were found more resistant to drought. This divergence in desiccation resistance seems to explain the unexpected life history traits differences between habitats observed.     

Differences among six woody perennials native to Northern Europe in their level of genetic differentiation and adaptive potential at fine local scale

The ability of perennial species to adapt their phenology to present and future temperature conditions is important for their ability to retain high fitness compared to other competing plant species, pests, and pathogens. Many transplanting studies with forest tree species have previously reported substantial genetic differentiation among populations within their native range. However, the question of “how local is local” is still highly debated in conservation biology because studies on genetic patterns of variation within and among populations at the local scale are limited and scattered. In this study, we compare the level of genetic differentiation among populations of six different perennial plant species based on their variation in spring flushing. We assess the level of additive genetic variation present within the local population. For all six species, we find significant differentiation among populations from sites with mean annual temperature ranging between 7.4°C and 8.4°C. The observed variation can only be partly explained by the climate at the site of origin. Most clear relationship between early flushing and higher average spring temperature is observed for the three wind‐pollinated species in the study, while the relations are much less clear for the three insect‐pollinated species. This supports that pollination system can influence the balance between genetic drift and natural selection and thereby influence the level of local adaptation in long‐lived species. On the positive side, we find that the native populations of woody plant species have maintained high levels of additive genetic variation in spring phenology, although this also differs substantially among the six studied species.     

Molecular and quantitative genetic differentiation in European populations of Silene latifolia (Caryophyllaceae)

BACKGROUND AND AIMS: Among-population differentiation in phenotypic traits and allelic variation is expected as a consequence of isolation, drift, founder effects and local selection. Therefore, investigating molecular and quantitative genetic divergence is a pre-requisite for studies of local adaptation in response to selection under variable environmental conditions. METHODS: Among- and within-population variation were investigated in six geographically separated European populations of the white campion, Silene latifolia, both for molecular variation at six newly developed microsatellite loci and for quantitative variation in morphological and life-history traits. To avoid confounding effects of the maternal environment, phenotypic traits were measured on greenhouse-reared F(1) offspring. Tests were made for clinal variation, and the correlations among molecular, geographic and phenotypic distances were compared with Mantel tests. KEY RESULTS: The six populations of Silene latifolia investigated showed significant molecular and quantitative genetic differentiation. Geographic and phenotypic distances were significantly associated. Age at first flowering increased significantly with latitude and exhibited a Q(st) value of 0.17 in females and 0.10 in males, consistent with adaptation to local environmental conditions. By contrast, no evidence of isolation-by-distance and no significant association between molecular and phenotypic distances were found. CONCLUSIONS: Significant molecular genetic divergence among populations of Silene latifolia, from the European native range is consistent with known limited seed and pollen flow distances, while significant quantitative genetic divergence among populations and clinal variation for age at first flowering suggest local adaptation.     

Interaction between the intermediate host of Fascioliasis in Tunisia,Galba truncatula and a possible competitor,Melanoides tuberculata (Muller, 1774): a field study

The snail Melanoides tuberculata has been used successfully in the control of some snails, intermediate host of parasitosis. Melanoides tuberculata has been found in Tunisia, but its effects on populations of native snail have not yet been evaluated in the field. Our objective is to determine whether M. tuberculata competed with Galba truncatula, using a field study. Twelve monthly investigations were carried out along an irrigation system in Ain Soltane's oasis (southwest of Tunisia). Here, we describe the population dynamics of G. truncatula with and without M. tuberculata in two stations: a witness pilot station (S1), in which the snail lives alone, and an experimental station (S2), where two species live together. The abundance of G. truncatula varied according to subsistence or not of M. tuberculata. The number of annual generations is higher in S1 (four generations) than S2 (two generations). In the absence of other molluscs or predators that can influence the density of G. truncatula and assuming that climatic factors are mitigated by the presence of an oasean microclimate, these results show that the mollusc M. tuberculata has a competition with the vector species of fluke.     

Experimental demonstration of a causal relationship between heterogeneity of selection and genetic differentiation in quantitative traits

Comparisons of estimates of genetic differentiation at molecular markers (F(ST)) and at quantitative traits (Q(ST)) are a means of inferring the level and heterogeneity of selection in natural populations. However, such comparisons are questionable because they require that the influence of drift and selection on Q(ST) be detectable over possible background influences of environmental or nonadditive genetic effects on Q(ST)-values. Here we test this using an experimental evolution approach in metapopulations of Arabidopsis thaliana experiencing different levels of drift and selection heterogeneity. We estimated the intensity and heterogeneity of selection on morphological and phenological traits via selection differentials. We demonstrate that Q(ST)-values increased with increasing selection heterogeneity when genetic drift was limited. The effect of selection on Q(ST) was thus detectable despite significant genotype-by-environment interactions that most probably biased the estimates of genetic differentiation. Although they cannot be used as a direct validation of the conclusions of prior studies, our results strongly support both the relevance of Q(ST) as an estimator of genetic differentiation and the role of local selection in shaping the genetic differentiation of natural populations.     

Dispersal and population structure of two sympatric species of the mediterranean land snail genus Mastus (Gastropoda, Pulmonata, Enidae)

We studied the population dynamics of Mastus by investigating the effects of population structure, spatial ecology and biology of reproduction on the genetic diversity of two sympatric Mastus species endemic to the island of Crete. Over a period of 27 months, we carried out both mark–recapture and random quadrat sampling techniques in order to assess the dispersal trends, the aggregation patterns, the neighbourhood size and the habitat preferences of these species. There were 154 recorded movements for M. butoti and 114 for M. cretensis . Mean monthly dispersal was estimated at d  = 0.5 m for M. butoti and d  = 1 m for M. cretensis . Both species showed a random dispersal pattern but tended to aggregate in the field. Their populations were found to be highly structured owing to their highly parsimonious dispersal behaviour and the very low population densities, estimated at D  = 2.07 ± 0.16 and D  = 0.73 ± 0.16 individuals m⁻² for M. butoti and M. cretensis , respectively. The neighbourhood size did not exceed 150 individuals for either species. The habitat occupied by each species changed during the active season, but both the immature and the adult individuals of each species seemed to prefer the same habitats throughout the active season. Partial population activation during the active season was observed in both species. We conclude that the population structure, the partial population activation and the species-specific reproductive strategies have a profound effect on maintaining the genetic diversity of the genus' populations.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 131–144.     

Comparison of quantitative and molecular variation in agroforestry populations of the shea tree (Vitellaria paradoxa C.F. Gaertn) in Mali

In this study we investigated the within- and between-population genetic variation using microsatellite markers and quantitative traits of the shea tree, Vitellaria paradoxa, an important agroforestry tree species of the Sudano–Sahelian region in Africa. Eleven populations were sampled across Mali and in northern Côte d’Ivoire. Leaf size and form and growth traits were measured in a progeny test at the nursery stage. Eight microsatellites were used to assess neutral genetic variation. Low levels of heterozygosity were recorded (1.6–3.0 alleles/locus; H_E = 0.25–0.42) and the fixation index (F_IS = −0.227–0.186) was not significantly different from zero suggesting that Hardy–Weinberg equilibrium is encountered in all populations sampled. Quantitative traits exhibited a strong genetic variation between populations and between families within populations. The degree of population differentiation of the quantitative traits (Q_ST = 0.055–0.283, Q_STmean = 0.189) strongly exceeds that in eight microsatellite loci (F_ST = −0.011–0.142, F_STmean = 0.047). Global and pairwise F_ST values were very low and not significantly different from zero suggesting agroforestry practices are amplifying gene flow (Nm = 5.07). The population means for quantitative traits and the rainfall variable were not correlated, showing variation was not linked with this climatic cline. It is suggested that this marked differentiation for quantitative traits, independent of environmental clines and despite a high gene flow, is a result of local adaptation and human selection of shea trees. This process has induced high linkage disequilibrium between underlying loci of polygenic characters.     

Sublethal effects of a Beauveria bassiana-based mycopesticide on Dutch and Serbian populations of Encarsia formosa (Hymenoptera: Aphelinidae)

ABSTRACT

The effects of a Beauveria bassiana-based product (trade name Naturalis-L; strain ATCC 74040) on the behaviour, life history traits and population growth of one commercial Dutch strain and two local Serbian populations (Bujanovac and Negotin) of Encarsia formosa Gahan (Hymen.: Aphelinidae) were evaluated in laboratory bioassays. In a two-choice test, parasitoids from all tested populations preferred to parasitise untreated whitefly nymphs over those treated with 1, 0.5, 0.25, 0.125 and 0.0625?ml/l of the mycopesticide. The longevity of adults of population Bujanovac exposed for 48?h to residues of the mycoinsecticide applied at the LC₅₀ concentration was 1.4 days shorter than that of control wasps. Total parasitism was significantly reduced (by 17.5, 26.2 and 31.4% for the Negotin, Dutch and Bujanovac populations, respectively) compared to control, as well as adult emergence in the F₁ generation (reductions of 20.3 and 26.7%, for the Dutch and Bujanovac populations, respectively). The instantaneous rate of increase (r_i) of surviving adults was also significantly reduced by mycopesticide treatment by 6.3, 13.8 and 15.1%, for the Negotin, Bujanovac, and Dutch populations, respectively. Adult parasioids emerged from pupae treated with the LC₅₀ concentration of the mycopesticide showed slower juvenile development, longevity, parasitism, adult emergence, and r_i values, but the reduction in r_i was significant only for the Bujanovac and Dutch populations (4.2 and 8.3%, respectively). The implications of these results on integrated control of the greenhouse whitefly Trialeurodes vaporariorum Westwood (Hem.: Aleyrodidae) are discussed in terms of adjusting the timing of Naturalis-L applications and parasitoid release.     

Genetic variation of Mimetes hirtus and Mimetes fimbriifolius (Proteaceae) – a comparative analysis of two closely related fynbos species

We investigated the influence of differing life history traits on the genetic structure of the related species Mimetes fimbriifolius and Mimetes hirtus (Proteaceae), which occur in the South African fynbos. Both species are bird‐pollinated and ant‐dispersed, but differ in rarity, longevity, ecological strategy and the fragmentation of their distribution area. We used AFLPs to study genetic variation within and between 21 populations of these two species across their distribution range. AFLP analysis revealed significantly higher genetic variation within populations of M. fimbriifolius than within M. hirtus. While M. fimbriifolius clearly lacked any significant genetic differentiation between populations, a distinct geographic pattern was observed for M. hirtus. Differentiation was, however, stronger at the regional (Φ_PT = 0.57) than at the local scale (Φ_PT = 0.08). Our results clearly indicate that even closely related species that share the same mode of pollination and seed dispersal can differ in their genetic structure, depending on the magnitude of fragmentation, longevity of individuals and ecological strategy.     

Contrasting patterns of diversity and population differentiation at the innate immunity gene toll-like receptor 2 (TLR2) in two sympatric rodent species

Comparing patterns of diversity and divergence between populations at immune genes and neutral markers can give insights into the nature and geographic scale of parasite-mediated selection. To date, studies investigating such patterns of selection in vertebrates have primarily focused on the acquired branch of the immune system, whereas it remains largely unknown how parasite-mediated selection shapes innate immune genes both within and across vertebrate populations. Here, we present a study on the diversity and population differentiation at the innate immune gene Toll-like receptor 2 (TLR2) across nine populations of yellow-necked mice (Apodemus flavicollis) and bank voles (Myodes glareolus) in southern Sweden. In yellow-necked mice, TLR2 diversity was very low, as was TLR2 population differentiation compared to neutral loci. In contrast, several TLR2 haplotypes co-occurred at intermediate frequencies within and across bank vole populations, and pronounced isolation by distance between populations was observed. The diversity and differentiation at neutral loci was similar in the two species. These results indicate that parasite-mediated selection has been acting in dramatically different ways on a given immune gene in ecologically similar and sympatric species. Furthermore, the finding of TLR2 population differentiation at a small geographical scale in bank voles highlights that vertebrate innate immune defense may be evolutionarily more dynamic than has previously been appreciated.     

Natural selection and the genetic differentiation of coastal and Arctic populations of the Atlantic cod in northern Norway: a test involving nucleotide sequence variation at the pantophysin (PanI) locus

To examine the role of contemporary selection in maintaining significant allele frequency differences at the pantophysin (PanI) locus among populations of the Atlantic cod, Gadus morhua, in northern Norway, we sequenced 127 PanIA alleles sampled from six coastal and two Barents Sea populations. The distributions of variable sites segregating within the PanIA allelic class were then compared among the populations. Significant differences were detected in the overall frequencies of PanIA alleles among populations within coastal and Arctic regions that was similar in magnitude to heterogeneity in the distributions of polymorphic sites segregating within the PanIA allelic class. The differentiation observed at silent sites in the PanIA allelic class contradicts the predicted effects of widescale gene flow and suggests that postsettlement selection acting on cohorts cannot be responsible for the genetic differences described between coastal and Arctic populations. Our results suggest that the marked differences observed between coastal and Arctic populations of G. morhua in northern Norway at the PanI locus reflect the action of recent diversifying selection and that populations throughout the region may be more independent than suggested by previous studies.