Journal of Evolutionary Biology

Published by Wiley and European Society of Evolutionary Biology
Online ISSN: 1420-9101
Publications
Article
In many katydids, the male feeds his mate with a large gelatinous spermatophore. Males of most species also produce elaborate calling songs. We predicted a negative relationship between spermatophore size and call frequency because of trade-offs between these two costly traits. Our comparative analysis controlling phylogeny and body size supported this prediction. Although call frequency is expected to decrease with increasing body size, after controlling for phylogeny, both variables were not related. Finally, given that song frequency and spermatophore size are likely targets of sexual selection, we examined the relationship between these variables and sexual size dimorphism (SSD) which can be influenced by sexual selection on body size. We found that only female body size was positively related to SSD, suggesting that natural and/or sexual selection on female body size may be stronger than sexual selection on male and spermatophore size.
 
Association between family mean and variance component genetic correlation estimates. Each point represents the estimate of genetic correlation for a trait in different environments by family mean correlation and by correlation based on variance components. Solid line shows 1 : 1 relationship. (a) Circles and long dashed line are broad sense estimates and X and short dashed line are narrow sense estimates. Both regression lines are model II. (b) Open circles are estimates based on separate variance components (methods 2 and 4) and closed circles are estimates based of common variance components (methods 3 and 5). Dashed line is model II regression of all data. 
Relationship between magnitude of discrepancy in genetic correlation estimates and average family size. Solid line shows slope of best fit. 
a) Means and full-sib heritabilities for H. axyridis traits calculated from variance components obtained from REML and LS ANOVA ANOVA estimates. (b) Means and half-sib heritabilities for N. cinerea traits calculated from variance components obtained from REML and LS ANOVA ANOVA estimates. 
Article
Advanced techniques for quantitative genetic parameter estimation may not always be necessary to answer broad genetic questions. However, simpler methods are often biased, and the extent of this determines their usefulness. In this study we compare family mean correlations to least squares and restricted error maximum likelihood (REML) variance component approaches to estimating cross-environment genetic correlations. We analysed empirical data from studies where both types of estimates were made, and from studies in our own laboratories. We found that the agreement between estimates was better when full-sib rather than half-sib estimates of cross-environment genetic correlations were used and when mean family size increased. We also note biases in REML estimation that may be especially important when testing to see if correlations differ from 0 or 1. We conclude that correlations calculated from family means can be used to test for the presence of genetic correlations across environments, which is sufficient for some research questions. Variance component approaches should be used when parameter estimation is the objective, or if the goal is anything other than determining broad patterns.
 
Article
Parasite-mediated selection on major histocompatibility complex (MHC) genes has mainly been explored at the intraspecific level, although many molecular studies have revealed trans-species polymorphism. Interspecific patterns of MHC diversity might reveal factors responsible for the long-term evolution of MHC polymorphism. We hypothesize that host taxa harbouring high parasite diversity should exhibit high levels of MHC genetic diversity. We test this assumption using data on rodent species and their helminth parasites compiled from the literature. Controlling for similarity due to common descent, we present evidence indicating that high helminth species richness in rodent species is associated with increased MHC class II polymorphism. Our results are consistent with the idea that parasites sharing a long-term coevolutionary history with their hosts are the agents of selection explaining MHC polymorphism.
 
Article
Serpentine soils provide a difficult substrate for plant colonization and growth and therefore represent an ideal system for studying the genetics of habitat adaptation and the evolution of plant-ecotypes. Using an F2 mapping population derived from an intraspecific cross between a serpentine and a nonserpentine ecotype of Silene vulgaris, the genetic architecture of seven morphological, physiological and life-history traits was explored. A quantitative trait locus (QTL) analysis identified 23 QTLs, 15 of which were classified as major QTLs. The observed genetic architecture suggests that traits potentially involved in habitat adaptation are controlled by few genes of major effect and have evolved under consistent directional selection. Several linkage groups harboured overlapping QTLs for different traits, which can be due to either pleiotropy or linkage. The potential roles of these factors and of the time available for habitat adaptation and ecological speciation on serpentine are discussed.
 
Article
Fine scale spatial structure (FSSS) of cytoplasmic genes in plants is thought to be generated via founder events and can be amplified when seeds germinate close to their mother. In gynodioecious species these processes are expected to generate FSSS in sex ratio because maternally inherited cytoplasmic male sterility genes partially influence sex expression. Here we document a striking example of FSSS in both mitochondrial genetic markers and sex in roadside populations of Silene vulgaris. We show that in one population FSSS of sexes influences relative fruit production of females compared to hermaphrodites. Furthermore, FSSS in sex ratio is expected to persist into future generations because offspring sex ratios from females are female-biased whereas offspring sex ratios from hermaphrodites are hermaphrodite-biased. Earlier studies indicated that pollen limitation is the most likely mechanism underlying negative frequency dependent fitness of females. Our results support the theoretical predictions that FSSS in sex ratio can reduce female fitness by decreasing the frequency at which females experience hermaphrodites. We argue that the influence of FSSS on female fitness is complementary to the influence of larger scale population structure on female fitness, and that population structure at both scales will act to decrease female frequencies in gynodioecious species. Better comprehension of the spatial structure of genders and genes controlling sex expression at a local scale is required for future progress toward understanding sex ratio evolution in gynodioecious plants.
 
Article
In recent years there has been a large body of theoretical work examining how local competition can reduce and even remove selection for altruism between relatives. However, it is less well appreciated that local competition favours selection for spite, the relatively neglected ugly sister of altruism. Here, we use extensions of social evolution theory that were formulated to deal with the consequences for altruism of competition between social partners, to illustrate several points on the evolution of spite. Specifically, we show that: (i) the conditions for the evolution of spite are less restrictive than previously assumed; (ii) previous models which have demonstrated selection for spite often implicitly assumed local competition; (iii) the scale of competition must be allowed for when distinguishing different forms of spite (Hamiltonian vs. Wilsonian); (iv) local competition can enhance the spread of spiteful greenbeards; and (v) the theory makes testable predictions for how the extent of spite should vary dependent upon population structure and average relatedness.
 
Article
When dispersal is limited, crosses between different regions may generate progeny of higher fitness than crosses within regions, due to the fact that individuals from the same region are more likely to share the same recessive deleterious alleles. This phenomenon (termed heterosis) generates a selective force favouring dispersal; however, the importance of heterosis on dispersal evolution has been a subject for debate. In this paper, we use computer simulations representing deleterious mutations occurring over a whole genome (of arbitrary map length R) to explore the magnitude of heterosis, and its effect on the evolution of dispersal. These results show that heterosis may have important effects on dispersal when it is in the upper range of values observed in natural populations, which occurs in our simulations when the genomic deleterious mutation rate U is also in the upper range of observed values. Comparing the results with extrapolations from an analytical two-locus model indicates that the effect of heterosis is mainly driven by pairwise associations between the locus affecting dispersal and selected loci when U is not too high (roughly, U < 0.5), whereas higher order associations become important for higher values of U.
 
Scatterplot of the ratio of relative arm length (arm length/SVL) to relative leg length (leg length/SVL). The means ± SD are displayed for each genus. Pictures show Myobatrachus gouldii, Mixophyes coggeri and Pseudophryne corroboree, representative morphologies according to relative limb length.
Summary of mean PC 1 values of the environmental data set for each species showing the environmental breadth displayed in each genus.
Article
Body shape is predicted to differ among species for functional reasons and in relation to environmental niche and phylogenetic history. We quantified morphological differences in shape and size among 98.5% of the 129 species and all 21 genera of the Australo-Papuan endemic myobatrachid frogs to test the hypothesis that habitat type predicts body shape in this radiation. We tested this hypothesis in a phylogenetic context at two taxonomic levels: across the entire radiation and within the four largest genera. Thirty-four external measurements were taken on 623 museum specimens representing 127 species. Data for seven key environmental variables relevant to anurans were assembled for all Australian-distributed species based on species' distributions and 131,306 locality records. The Australo-Papuan myobatrachid radiation showed high diversity in adult body size, ranging from minute (15 mm snout-vent length) to very large species (92 mm), and shape, particularly sin relative limb length. Five main morphological and environmental summary variables displayed strong phylogenetic signal. There was no clear relationship between body size and environmental niche, and this result persisted following phylogenetic correction. For most species, there was a better match between environment/habitat and body shape, but this relationship did not persist following phylogenetic correction. At a broad level, species fell into three broad groups based on environmental niche and body shape: 1) species in wet habitats with relatively long limbs, 2) species in arid environments with relatively short limbs (many of which are forward or backward burrowers) and 3) habitat generalist species with a conservative body shape. However, these patterns were not repeated within the four largest genera - Crinia, Limnodynastes, Pseudophryne and Uperoleia. Each of these genera displayed a highly conservative anuran body shape, yet individual species were distributed across the full spectrum of Australian environments. Our results suggest that phylogenetic legacy is important in the evolution of body size and shape in Australian anurans, but also that the conservative body plan of many frogs works well in a wide variety of habitats.
 
Article
Sexually selected traits that are costly are predicted to be more condition dependent than nonsexually selected traits. Assuming resource limitation, increased allocation to a sexually selected trait may also come at a cost to other fitness components. To test these predictions, we varied adult food ration to manipulate condition in the colour dimorphic bug, Phymata americana. We compared the degree of condition dependence in a sexually selected trait expressed in males to a nonsexually selected trait expressed in males and females. We also evaluated the effects of condition on longevity of both sexes. We found that the expression of these colour pattern traits was strongly influenced by both diet and age. As expected, the strength of condition dependence was much more pronounced in the sexually selected, male-limited trait but the nonsexual trait also exhibited significant condition dependence in both sexes. The sexually selected male trait also exhibited a higher coefficient of phenotypic variation than the nonsexually selected trait in males and females. Diet had contrasting effects on male and female longevity; increased food availability had positive effects on female lifespan but these effects were not detected in males, suggesting that males allocated limited resources preferentially to sexually selected traits. These results are consistent with the expectation that optimal allocation to various fitness components differs between the sexes.
 
Article
Why sex is so common remains unclear; what is certain is that the predominance of sex despite its profound costs means that it must confer major advantages. Here, we use elemental and nucleic acid assays to evaluate a key element of a novel, integrative hypothesis considering whether sex might be favoured because of differences in body composition between sexuals and asexuals. We found that asexual Potamopyrgus antipodarum, a New Zealand snail, have markedly higher bodily phosphorus and nucleic acid content per unit mass than sexual counterparts. These differences coincide with and are almost certainly linked to the higher ploidy of the asexuals. Our results are the first documented body composition differences between sexual and asexual organisms, and the first detected phenotypic difference between sexual and asexual P. antipodarum, an important natural model system for the study of the maintenance of sex. These findings also verify a central component of our hypothesis that competition between diploid sexuals and polyploid asexuals could be influenced by phosphorus availability.
 
Article
It has often been argued that evolutionary diversification is the result of divergent natural selection for specialization on alternative resources. I provide a comprehensive review of experiments that examine the ecology and genetics of resource specialization and adaptive radiation in microbial microcosms. In these experiments, resource heterogeneity generates divergent selection for specialization on alternative resources. At a molecular level, the evolution of specialization is generally attributable to mutations that de-regulate the expression of existing biosynthetic and catabolic pathways. Trade-offs are associated with the evolution of resource specialization, but these trade-offs are often not the result of antagonistic pleiotropy. Replicate adaptive radiations result in the evolution of a similar assemblage of specialists, but the genetic basis of specialization differs in replicate radiations. The implications of microbial selection experiments for evolutionary theory are discussed and future directions of research are proposed.
 
Article
Our study addressed reproductive character displacement between two subspecies of the house mouse, Mus musculus musculus and Mus musculus domesticus that hybridize in Europe along a zone where selection against hybridization is known to occur. Based on a multi-population approach, we investigated spatial patterns of divergence of mate preference in the two taxa. Mate preference was significantly higher in the contact zone than in allopatry in both subspecies, suggesting that reproductive character displacement occurs. Moreover, patterns of preference were stronger in M. m. musculus than in M. m. domesticus, indicating an asymmetrical divergence between the two. In the context of selection against hybridization, our results may provide empirical support for the hypothesis of reinforcement in a parapatric hybrid zone. We discuss factors that could explain the asymmetrical pattern of divergence and the possible impact of a unimodal structure on the maintenance of premating divergence between the two subspecies.
 
Article
Spiders have recently emerged as important diversity hot spots for endosymbiotic bacteria, but the consequences of these symbiotic interactions are largely unknown. In this article, we examined the evolutionary history and effect of the intracellular bacterium Cardinium hertigii in the marbled cellar spider Holocnemus pluchei. We showed that Cardinium infection is primarily transmitted in spider populations maternally via egg cytoplasm, with 100% of the progeny from infected mothers being also infected. Examination of a co-inherited marker, mitochondrial DNA (mtDNA), revealed that Cardinium infection is associated with a wide diversity of mtDNA haplotypes, showing that the interaction between Cardinium and H. pluchei has a long-term evolutionary dimension and that horizontal transmission among individuals could also occur. Although Cardinium is well known to exert sex ratio distortion or cytoplasmic incompatibility in various arthropod hosts, we show, however, that Cardinium does not interact with the reproductive biology of H. pluchei. A field survey shows a clear geographical structuring of Cardinium infection, with a marked gradual variation of infection frequencies from ca. 0.80 to 0. We discuss different mechanistic and evolutionary explanations for these results as well as their consequences for spider phenotypes. Notably, we suggest that Cardinium can either behave as a neutral cytoplasmic element within H. pluchei or exhibit a context-dependent effect, depending on the environmental conditions.
 
Article
When facing the challenge of developing an individual that best fits its environment, nature demonstrates an interesting combination of two fundamentally different adaptive mechanisms: genetic evolution and phenotypic plasticity. Following numerous computational models, it has become the accepted wisdom that lifetime acclimation (e.g. via learning) smooths the fitness landscape and consequently accelerates evolution. However, analytical studies, focusing on the effect of phenotypic plasticity on evolution in simple unimodal landscapes, have often found that learning hinders the evolutionary process rather than accelerating it. Here, we provide a general framework for studying the effect of plasticity on evolution in multipeaked landscapes and introduce a rigorous mathematical analysis of these dynamics. We show that the convergence rate of the evolutionary process in a given arbitrary one-dimensional fitness landscape is dominated by the largest descent (drawdown) in the landscape and provide numerical evidence to support an analogous dominance also in multidimensional landscapes. We consider several schemes of phenotypic plasticity and examine their effect on the landscape drawdown, identifying the conditions under which phenotypic plasticity is advantageous. The lack of such a drawdown in unimodal landscapes vs. its dominance in multipeaked landscapes accounts for the seemingly contradictory findings of previous studies.
 
Illustration of the negative genetic correlation between boldness and docility in the Ram Mountain population. Breeding values of boldness and docility were predicted from a bivariate animal model analysis with VA and year as random effects.
Personality and age-dependent reproductive success in bighorn rams. Annual relative reproductive success as a function of the interaction between docility and age (a), boldness and age (b), and boldness and horn length (c). One hundred and five rams and 461 estimates of paternity were used for the analysis. The length of both X and Y axes is limited to the natural range of data.
Selected generalized linear model of longevity as a function of personality and other traits in bighorn rams.
Selected linear mixed model of annual relative reproduc- tive success as a function on personality and other traits in bighorn rams.
Article
Recent theoretical work suggests that personality is a component of life history, but links between personality and either age-dependent reproductive success or life-history strategy are yet to be established. Using quantitative genetic analyses on a long-term pedigree we estimated indices of boldness and docility for 105 bighorn sheep rams (Ovis canadensis), born between 1983 and 1999, and compared these indices to their reproductive history from 2 years of age until death. Docility and boldness were highly heritable and negatively genetically correlated. Docile and bold rams survived longer than indocile and shy rams. Docility and boldness had a weak negative effect on reproductive success early in life, but a strong positive effect on older rams. Our findings highlight an important role of personality on reproductive success and suggest that personality could be an important component of life-history strategy.
 
Article
Understanding the origin, maintenance and significance of phenotypic variation is one of the central issues in evolutionary biology. An ongoing discussion focuses on the relative roles of isolation and selection as being at the heart of genetically based spatial variation. We address this issue in a representative of a taxon group in which isolation is unlikely: a marine broadcast spawning invertebrate. During the free-swimming larval phase, dispersal is potentially very large. For such taxa, small-scale population genetic structuring in neutral molecular markers tends to be limited, conform expectations. Small-scale differentiation of selective traits is expected to be hindered by the putatively high gene flow. We determined the geographical distribution of molecular markers and of variation in a shell shape measure, globosity, for the bivalve Macoma balthica (L.) in the western Dutch Wadden Sea and adjacent North Sea in three subsequent years, and found that shells of this clam are more globose in the Wadden Sea. By rearing clams in a common garden in the laboratory starting from the gamete phase, we show that the ecotypes are genetically different; heritability is estimated at 23%. The proportion of total genetic variation that is between sites is much larger for the morphological additive genetic variation (QST = 0.416) than for allozyme (FST = 0.000-0.022) and mitochondrial DNA cytochrome-c-oxidase-1 sequence variation (phiST = 0.017). Divergent selection must be involved and intraspecific spatial genetic differentiation in marine broadcast spawners is apparently not constrained by low levels of isolation.
 
Article
Putative hybrids between Banksia hookeriana and B. prionotes were identified among 12 of 106 populations of B. hookeriana located at or near anthropogenically disturbed sites, mainly roadways, but none in 156 undisturbed populations. Morphometrics and AFLP markers confirmed that a hybrid swarm existed in a selected disturbed habitat, whereas no intermediates were present where the two species co-occurred in undisturbed vegetation. Individuals of both species in disturbed habitats at 12 sites were more vigorous, with greater size and more flower heads than their counterparts in undisturbed vegetation. These more fecund plants also showed a shift in season and duration of flowering. By promoting earlier flowering of B. hookeriana plants and prolonging flowering of B. prionotes, anthropogenic disturbance broke the phenological barrier between these two species. We conclude that anthropogenic disturbance promotes hybridization through increasing opportunities for gene flow by reducing interpopulation separation, increasing gamete production and, especially, promoting coflowering.
 
Article
Sexual conflict has been proposed as a potential selective agent in the evolution of a variety of traits. Here, we present a simple model that investigates the initial conditions under which sex-linked and sex-limited harming alleles can invade a population. In this paper, we expand previous threshold models to study how sex-linkage and sex determination mechanisms affect the spreading conditions of a harming allele. Our models provide new insights into how sexual conflict could originate, showing that in diploid organisms the probability of a new harming allele spreading is independent of both the genetic sex determination system and the dominance relationships. However, the incidence of interlocus sexual conflicts in the initial steps of the invasion critically depends on the inheritance system.
 
Article
Mutations that are beneficial in one environment can have different fitness effects in other environments. In the context of antibiotic resistance, the resulting genotype-by-environment interactions potentially make selection on resistance unpredictable in heterogeneous environments. Furthermore, resistant bacteria frequently fix additional mutations during evolution in the absence of antibiotics. How do these two types of mutations interact to determine the bacterial phenotype across different environments? To address this, I used Escherichia coli as a model system, measuring the effects of nine different rifampicin resistance mutations on bacterial growth in 31 antibiotic-free environments. I did this both before and after approximately 200 generations of experimental evolution in antibiotic-free conditions (LB medium), and did the same for the antibiotic-sensitive wild type after adaptation to the same environment. The following results were observed: (i) bacteria with and without costly resistance mutations adapted to experimental conditions and reached similar levels of competitive fitness; (ii) rifampicin resistance mutations and adaptation to LB both indirectly altered growth in other environments; and (iii) resistant-evolved genotypes were more phenotypically different from the ancestor and from each other than resistant-nonevolved and sensitive-evolved genotypes. This suggests genotype-by-environment interactions generated by antibiotic resistance mutations, observed previously in short-term experiments, are more pronounced after adaptation to other types of environmental variation, making it difficult to predict long-term selection on resistance mutations from fitness effects in a single environment.
 
Article
Most conifer species occur in large continuous populations, but radiata pine, Pinus radiata, occurs only in five disjunctive natural populations in California and Mexico. The Mexican island populations were presumably colonized from the mainland millions of years ago. According to Axelrod (1981), the mainland populations are relicts of an earlier much wider distribution, reduced some 8,000 years ago, whereas according to Millar (1997, 2000), the patchy metapopulation-like structure is typical of the long-term population demography of the species. We used 19 highly polymorphic microsatellite loci to describe population structure and to search for signs of the dynamics of population demography over space and time. Frequencies of null alleles at microsatellite loci were estimated using an approach based on the probability of identity by descent. Microsatellite genetic diversities were high in all populations [expected heterozygosity (H(e)) = 0.68-0.77], but the island populations had significantly lower estimates. Variation between loci in genetic differentiation (F(ST)) was high, but no locus deviated statistically significantly from the rest at an experiment wide level of 0.05. Thus, all loci were included in subsequent analysis. The average differentiation was measured as F(ST) = 0.14 (SD 0.012), comparable with earlier allozyme results. The island populations were more diverged from the other populations and from an inferred common ancestral gene pool than the mainland ones. All populations showed a deficiency of expected heterozygosity given the number of alleles, the mainland populations more so than the island ones. The results thus do not support a recent important contraction in the mainland range of radiata pine.
 
Article
It is widely assumed that microsatellites are generated by replication slippage, a mutation process specific to repetitive DNA. Consistent with their high mutation rate, microsatellites are highly abundant in most eukaryotic genomes. In Escherichia coli, however, microsatellites are rare and short despite the fact that a high microsatellite mutation rate was described. We show that this high microsatellite instability depends on the presence of the F-plasmid. E. coli cells lacking the F-plasmid have extremely low microsatellite mutation rates. This result provides a possible explanation for the genome-wide low density of microsatellites in E. coli. Furthermore, we show that the F-plasmid induced microsatellite instability is independent of the mismatch repair pathway.
 
Article
Self-incompatibility in Arabidopsis lyrata is sporophytically controlled by the multi-allelic S-locus. Self-incompatibility alleles (S-alleles) are under strong negative frequency dependent selection because pollen carrying common S-alleles have fewer mating opportunities. Population genetics theory predicts that deleterious alleles can accumulate if linked to the S-locus. This was tested by studying segregation of S-alleles in 11 large full sib families in A. lyrata. Significant segregation distortion leading to an up to fourfold difference in transmission rates was found in six families. Differences in transmission rates were not significantly different in reciprocal crosses and the distortions observed were compatible with selection acting at the gametic stage alone. The S-allele with the largest segregation advantage is also the most recessive, and is very common in natural populations concordant with its apparent segregation advantage. These results imply that frequencies of S-alleles in populations of A. lyrata cannot be predicted based on simple models of frequency-dependent selection alone.
 
Article
In the newly emerging field of statistical phylogeography, consideration of the stochastic nature of genetic processes and explicit reference to theoretical expectations under various models has dramatically transformed how historical processes are studied. Rather than being restricted to ad hoc explanations for observed patterns of genetic variation, assessments about the underlying evolutionary processes are now based on statistical tests of various hypotheses, as well as estimates of the parameters specified by the models. A wide range of demographical and biogeographical processes can be accommodated by these new analytical approaches, providing biologically more realistic models. Because of these advances, statistical phylogeography can provide unprecedented insights about a species' history, including decisive information about the factors that shape patterns of genetic variation, species distributions, and speciation. However, to improve our understanding of such processes, a critical examination and appreciation of the inherent difficulties of historical inference and challenges specific to testing phylogeographical hypotheses are essential. As the field of statistical phylogeography continues to take shape many difficulties have been resolved. Nonetheless, careful attention to the complexities of testing historical hypotheses and further theoretical developments are essential to improving the accuracy of our conclusions about a species' history.
 
Article
Comparative studies of quantitative genetic and neutral marker differentiation have provided means for assessing the relative roles of natural selection and random genetic drift in explaining among-population divergence. This information can be useful for our fundamental understanding of population differentiation, as well as for identifying management units in conservation biology. Here, we provide comprehensive review and meta-analysis of the empirical studies that have compared quantitative genetic (Q(ST)) and neutral marker (F(ST)) differentiation among natural populations. Our analyses confirm the conclusion from previous reviews - based on ca. 100% more data - that the Q(ST) values are on average higher than F(ST) values [mean difference 0.12 (SD 0.27)] suggesting a predominant role for natural selection as a cause of differentiation in quantitative traits. However, although the influence of trait (life history, morphological and behavioural) and marker type (e.g. microsatellites and allozymes) on the variance of the difference between Q(ST) and F(ST) is small, there is much heterogeneity in the data attributable to variation between specific studies and traits. The latter is understandable as there is no reason to expect that natural selection would be acting in similar fashion on all populations and traits (except for fitness itself). We also found evidence to suggest that Q(ST) and F(ST) values across studies are positively correlated, but the significance of this finding remains unclear. We discuss these results in the context of utility of the Q(ST)-F(ST) comparisons as a tool for inferring natural selection, as well as associated methodological and interpretational problems involved with individual and meta-analytic studies.
 
Male productivity in the single mating/single mate condition. The points represent the mean numbers of adult progeny in successive vials produced per individual female after mating with a ST male or a SR5 male (n = 40). For each vial, the male genotype effect was tested by anova, the significance is indicated on the figure (*P < 0.05, **P < 0.01, ***P < 0.001). m, mean male productivity.
Mean premating time and copulation duration (in minutes).
Progeny sex ratios in the sperm competition experiment.
Article
The aim of the present study was to determine whether the effects of sex-ratio segregation distorters on the fertility of male Drosophila simulans can explain the contrasting success of these X-linked meiotic drivers in different populations of the species. We compared the fertility of sex-ratio and wild-type males under different mating conditions. Both types were found to be equally fertile when mating was allowed, with two females per male, during the whole period of egg laying. By contrast sex-ratio males suffered a strong fertility disadvantage when they were offered multiple mates for a limited time, or in sperm competition conditions. In the latter case only, the toll on male fertility exceeded the segregation advantage of the distorters. These results indicate that sex-ratio distorters can either spread or disappear from populations, depending on the mating rate. Population density is therefore expected to play a major role in the evolution of sex-ratio distorters in this Drosophila species.
 
Article
The ability to express heat-shock proteins (HSP) under thermal stress is an essential mechanism for ectotherms to cope with unfavourable conditions. In this study, we investigate if Copper butterflies originating from different altitudes and/or being exposed to different rearing and induction temperatures show differences in HSP70 expression. HSP70 expression increased substantially at the higher rearing temperature in low-altitude butterflies, which might represent an adaptation to occasionally occurring heat spells. On the other hand, high-altitude butterflies showed much less plasticity in response to rearing temperatures, and overall seem to rely more on genetically fixed thermal stress resistance. Whether the latter indicates a higher vulnerability of high-altitude populations to global warming needs further investigation. HSP70 expression increased with both colder and warmer induction temperatures.
 
Article
Phenotypic differentiation between populations is thought to occur mainly at spatial scales where gene-flow is restricted and selection regimes differ. However, if gene flow is nonrandom, dispersal may reinforce, rather than counteract, evolutionary differentiation, meaning that differences occurring over small scales might have a genetic basis. The purpose of this study was to determine the cause of differences in mean phenotype between two parts of a population of great tits Parus major, separated by <3 km. We conducted a partial cross-fostering experiment between two contrasting parts of this population to separate genetic and environmental sources of variation, and to test for gene-environment interaction. We found strong environmental effects on nestling size, mass and condition index, with nestlings reared in a low density part of the population being larger, heavier and in better condition, than those in a high density part, irrespective of their origin. In addition, we found smaller, but significant, differences in nestling condition and shape associated with the areas that birds originated from, suggesting the presence of genetic differences between parts of this population. There was no evidence of gene-environment interaction for any character. This experiment is thus consistent with previous analyses suggesting that differences between parts of this population had evolved recently, apparently due to phenotype-dependent dispersal, and indicates that population differentiation can be maintained over small spatial scales despite extensive dispersal.
 
Article
We investigated genetic variability and genetic correlations in early life-history traits of Crassostrea gigas. Larval survival, larval development rate, size at settlement and metamorphosis success were found to be substantially heritable, whereas larval growth rate and juvenile traits were not. We identified a strong positive genetic correlation between larval development rate and size at settlement, and argue that selection could optimize both age and size at settlement. However, trade-offs, resulting in costs of metamorphosing early and large, were suggested by negative genetic correlations or covariances between larval development rate/size at settlement and both metamorphosis success and juvenile survival. Moreover, size advantage at settlement disappeared with time during the juvenile stage. Finally, we observed no genetic correlations between larval and juvenile stages, implying genetic independence of life-history traits between life-stages. We suggest two possible scenarios for the maintenance of genetic polymorphism in the early life-history strategy of C. gigas.
 
Article
The maternally inherited bacterium, Wolbachia pipientis, manipulates host reproduction by rendering uninfected females reproductively incompatible with infected males (cytoplasmic incompatibility, CI). Hosts may evolve mechanisms, such as mate preferences, to avoid fitness costs of Wolbachia infection. Despite the potential importance of mate choice for Wolbachia population dynamics, this possibility remains largely unexplored. Here we model the spread of an allele encoding female mate preference for uninfected males alongside the spread of CI inducing Wolbachia. Mate preferences can evolve but the spread of the preference allele depends on factors associated with both Wolbachia infection and the preference allele itself. Incomplete maternal transmission of Wolbachia, fitness costs and low CI, improve the spread of the preference allele and impact on the population dynamics of Wolbachia. In addition, mate preferences are found in infected individuals. These results have important consequences for the fate of Wolbachia and studies addressing mate preferences in infected populations.
 
Article
Maternal effects on progeny wing size and shape in a homozygous parthenogenetic strain of Drosophila mercatorum were investigated. The impact of external maternal factors (heat stress) and the impact of internal maternal factors (different maternal and grand maternal age) were studied. The offspring developed under identical environmental conditions, and due to lack of genetic variation any phenotypic difference among offspring could be ascribed to maternal effects. Wing size was estimated by centroid size, shape was analysed with the Procrustes geometric morphometric method and variation in landmark displacement was visualized by principal component analysis. Both kinds of maternal effects had a significant impact on progeny wing size and shape. Maternal heat stress led to the same pattern of response in size and shape among the progeny, with increased difference between the control group and progeny from heat stressed flies in both size and shape with increased maternal heat stress temperature. The effects of maternal age, however, led to different responses in size and shape between the different progeny groups. The observed variation in landmark displacements was similar, and in both cases mainly associated with shape differences of the posterior part of the wing. Finally, our results suggest that maternal effect has some evolutionary implications by altering the genetic correlations among traits, which can affect the response to selective pressures.
 
Article
Extreme environments and adaptation.- The evolution of plants in metal-contaminated environments.- Responses of aquatic organisms to pollutant stress: Theoretical and practical implications.- Conifers from the cold.- Genetic variation and environmental stress.- Phenotypic plasticity and fluctuating asymmetry as responses to environmental stress in the butterflyBicyclus anynana.- Environmental stress and the expression of genetic variation.- Worldwide latitudinal clines for the alcohol dehydrogenase polymorphism in Drosophila melanogaster: What is the unit of selection?.- Stress and metabolic regulation inDrosophila.- Acclimation and response to thermal stress.- Phenotypic and evolutionary adaptation of a model bacterial system to stressful thermal environments.- Ecological and evolutionary physiology of heat shock proteins and the stress response inDrosophila: Complementary insights from genetic engineering and natural variation.- High-temperature stress and the evolution of thermal resistance inDrosophila.- Stress, selection and extinction.- Genetic and environmental stress, and the persistence of populations.- Adaptation and extinction in changing environments.- Environmental stress and evolution: A theoretical study.- Stress, developmental stability and sexual selection.- Evolution and stress.- Genetic variability and adaptation to stress.- Stress-resistance genotypes, metabolic efficiency and interpreting evolutionary change.- The Plus ca change model: Explaining stasis and evolution in response to abiotic stress over geological timescales.
 
Article
Heritable maternal effects have important consequences for the evolutionary dynamics of phenotypic traits under selection, but have only rarely been tested for or quantified in evolutionary studies. Here we estimate maternal effects on early-life traits in a feral population of Soay sheep (Ovis aries) from St Kilda, Scotland. We then partition the maternal effects into genetic and environmental components to obtain the first direct estimates of maternal genetic effects in a free-living population, and furthermore test for covariance between direct and maternal genetic effects. Using an animal model approach, direct heritabilities (h2) were low but maternal genetic effects (m2) represented a relatively large proportion of the total phenotypic variance for each trait (birth weight m2=0.119, birth date m2=0.197, natal litter size m2=0.211). A negative correlation between direct and maternal genetic effects was estimated for each trait, but was only statistically significant for natal litter size (ram= -0.714). Total heritabilities (incorporating variance from heritable maternal effects and the direct-maternal genetic covariance) were significant for birth weight and birth date but not for natal litter size. Inadequately specified models greatly overestimated additive genetic variance and hence direct h2 (by a factor of up to 6.45 in the case of birth date). We conclude that failure to model heritable maternal variance can result in over- or under-estimation of the potential for traits to respond to selection, and advocate an increased effort to explicitly measure maternal genetic effects in evolutionary studies.
 
Article
Inbreeding is expected to decrease the heritability within populations. However, results from empirical studies are inconclusive. In this study, we investigated the effects of three breeding treatments (fast and slow rate of inbreeding - inbred to the same absolute level - and a control) on heritability, phenotypic, genetic and environmental variances of sternopleural bristle number in Drosophila melanogaster. Heritability, and phenotypic, genetic and environmental variances were estimated in 10 replicate lines within each of the three treatments. Standard least squares regression models and Bayesian methods were used to analyse the data. Heritability and additive genetic variance within lines were higher in the control compared with both inbreeding treatments. Heritabilities and additive genetic variances within lines were higher in slow compared with fast inbred lines, indicating that slow inbred lines retain more evolutionary potential despite the same expected absolute level of inbreeding. The between line variance was larger with inbreeding and more than twice as large in the fast than in the slow inbred lines. The different pattern of redistribution of genetic variance within and between lines in the two inbred treatments cannot be explained invoking the standard model based on selective neutrality and additive gene action. Environmental variances were higher with inbreeding, and more so with fast inbreeding, indicating that inbreeding and the rate of inbreeding affect environmental sensitivity. The phenotypic variance decreased with inbreeding, but was not affected by the rate of inbreeding. No inbreeding depression for mean sternopleural bristle number was observed in this study. Considerable variance between lines in additive genetic variance within lines was observed, illustrating between line variation in evolutionary potential.
 
The error bar (mean ± SE) of lytic activity (the change in optical density of hemolymph) in the low drumming rate treatment (control) and high drumming rate treatment (female proximity).
The error bar (mean ± SE) of encapsulation rate (the amount of melanic pigments in a monofilament expressed as grey values) in the low drumming rate treatment (control) and high drumming rate treatment (female proximity).
Article
The field of ecological immunology is ultimately seeking to address the question 'Why is there variation in immune function?' Here, we provide experimental evidence that costs of ubiquitous sexual signals are a significant source of variation in immune function. In the mating season, males of the wolf spider Hygrolycosa rubrofasciata drum against dry leaves while wandering around the habitat searching for receptive females. According to a previous study, the male metabolic rate during the drumming increases 22-fold compared to the resting metabolic rate. In the present study, we examined whether investment in costly courtship drumming decreases male immune function in a wild population of H. rubrofasciata. We induced males to increase their drumming rate by introducing females in proximity. As estimates of male immune function, we used lytic activity and encapsulation rate. Lytic activity estimates the concentration of antimicrobial peptides in haemolymph, which have been shown to play an important role in defence against bacteria, viruses and fungi. Encapsulation is an important defence mechanism against nematodes and insect parasitoids, but it also plays a role in defence against viruses. Our results show that males with nonarbitrarily increased investment in drumming rate had considerably lower lytic activities than control males. Also, there was a tendency for males with nonarbitrarily increased investment in drumming rate to have lower encapsulation rates than control males. This study provides experimental evidence for the first time, to our knowledge, that there are direct immunological costs of sexual signalling in natural populations. Therefore, immunological costs of sexual signals may provide significant phenotypic variation to parasite-mediated sexual selection.
 
Article
Life history theory predicts tradeoffs among reproductive traits, but the physiological mechanisms underlying such tradeoffs remain unclear. Here we examine reproductive tradeoffs and their association with yolk steroids in an oviparous lizard. Female leopard geckos lay two eggs in a clutch, produce multiple clutches in a breeding season, and reproduce for several years. We detected a significant tradeoff between egg size and the number of clutches laid by females during their first two breeding seasons. Total reproductive effort was strongly condition-dependent in the first season, but much less so in the second season. Although these and other tradeoffs were unmistakable, they were not associated with levels of androstenedione, oestradiol, or testosterone in egg yolk. Female condition and egg size, however, were inversely related to dihydrotestosterone (DHT) levels in egg yolk. Finally, steroid levels in egg yolk were not directly related to steroid levels in the maternal circulation when follicles were developing, indicating that steroid transfer to eggs is regulated. These findings suggest that maternal allocation of DHT could mitigate tradeoffs that lead to poor offspring quality (i.e. poor female condition) and small offspring size (i.e. small egg size).
 
Article
Hybrid zones are natural laboratories offering insights into speciation processes. Narrow hybrid zones are less common in the sea than on land consistent with higher dispersal among marine populations. Acanthochromis polyacanthus is an unusual bony marine fish with philopatric dispersal that exists as allopatric stocks of white, bicoloured and black fish on the Great Barrier Reef (GBR). At two latitudes, different morphs coexist and hybridize at narrow contact zones. Sequence data from mitochondrial Hypervariable Region 1 revealed contrasting patterns of introgression across these zones. At the northern hybrid zone, a single clade of mitochondrial haplotypes was found in all white fish, hybrids and tens of kilometres into pure bicoloured stock. At the southern hybrid zone, there was no introgression of mitochondrial genes into black fish and hybrids shared the bicoloured haplotypes. Based on this asymmetry, we postulate that black fish from the southern GBR have experienced a selective sweep of their mitochondrial genome, which has resulted in almost total reproductive isolation.
 
Trade-offs for Vmax  ∈  [0.1,0.9]: (a) f(Vmax) = Vmax, (b) g(Vmax) = 1−Vmax. The structure of the steady state solution N* = (N*1,…,N*n) of eqns 8–11 as a function of the dilution rate D and the mutation rate ɛ is presented in (c). A numerically computed steady state is presented in (d) with ɛ = 10−7 and D = 0.01, and in (e) with ɛ=10−7 and D = 0.04. The remainder of the parameters have the following values:  with n = 500, S0 = 3, c = 1.
Article
Understanding the evolution of microbial diversity is an important and current problem in evolutionary ecology. In this paper, we investigated the role of two established biochemical trade-offs in microbial diversification using a model that connects ecological and evolutionary processes with fundamental aspects of biochemistry. The trade-offs that we investigated are as follows:(1) a trade-off between the rate and affinity of substrate transport; and (2) a trade-off between the rate and yield of ATP production. Our model shows that these biochemical trade-offs can drive evolutionary diversification under the simplest possible ecological conditions: a homogeneous environment containing a single limiting resource. We argue that the results of a number of microbial selection experiments are consistent with the predictions of our model.
 
Maximum-likelihood trees of green algae, moss, fern, conifer and angiosperm (a) nuclear 18S rDNA and (b) chloroplast rbcL genes. See text for details.
Article
The evolutionary potential of bryophytes (mosses, liverworts and hornworts) has been debated for decades. Fossil record and biogeographical distribution patterns suggest very slow morphological evolution and the retainment of several ancient traits since the split with vascular plants some 450 million years ago. Many have argued that bryophytes may evolve as rapidly as higher plants on the molecular level, but this hypothesis has not been tested so far. Here, it is shown that mosses have experienced significantly lower rates of molecular evolution than higher plants within 18S rDNA (nuclear), rbcL (chloroplast) and nad5 (mitochondrial) genes. Mosses are on an average evolving 2-3 times slower than ferns, gymnosperms and angiosperms; and also green algae seem to be evolving faster than nonvascular plants. These results support the observation of a general correlation between morphological and molecular evolutionary rates in plants and also show that mosses are 'evolutionary sphinxes' regarding both morphological and molecular evolutionary potential.
 
Article
An allopolyploid complex with high genomic integrity has been studied. Dogroses transmit only seven chromosomes (from seven bivalents) through the pollen, whereas 21, 28 or 35 chromosomes (from seven bivalents and 14, 21 or 28 univalents) come from the egg cells. Seedlings derived from two interspecific crosses were analysed with flow cytometry and molecular markers to determine ploidy level, mode of reproduction and genomic constitution. Evidence was obtained for the formation of unreduced male and female gametes, which can take part in fertilization (producing seedlings with higher ploidy than the parental plants) or in apomictic reproduction. Random amplified polymorphic DNA (RAPD) and microsatellite analyses indicated that three seedlings (5%) were derived through apomixis, whereas the other 49 were hybrids. Bivalent formation appears to involve chromosomes that consistently share the same microsatellite alleles. Allele-sharing between the maternally transmitted and highly conserved univalent-forming chromosomes reflected the taxonomic distance between different genotypes. The frequently recombining bivalent-forming chromosomes were taxonomically less informative.
 
Article
We use individual-based stochastic simulations and analytical deterministic predictions to investigate the interaction between drift, natural selection and gene flow on the patterns of local adaptation across a fragmented species' range under clinally varying selection. Migration between populations follows a stepping-stone pattern and density decreases from the centre to the periphery of the range. Increased migration worsens gene swamping in small marginal populations but mitigates the effect of drift by replenishing genetic variance and helping purge deleterious mutations. Contrary to the deterministic prediction that increased connectivity within the range always inhibits local adaptation, simulations show that low intermediate migration rates improve fitness in marginal populations and attenuate fitness heterogeneity across the range. Such migration rates are optimal in that they maximize the total mean fitness at the scale of the range. Optimal migration rates increase with shallower environmental gradients, smaller marginal populations and higher mutation rates affecting fitness.
 
Article
Cases of coexisting sexual and asexual relatives are puzzling, as evolutionary theory predicts that competition for the same ecological niches should lead to the exclusion of one or the other population. In the cyclically parthenogenetic aphid, Rhopalosiphum padi, sexual and facultative asexual lineages are admixed in space at the time of sexual reproduction. We investigated how the interaction of reproductive mode and environment can lead to temporal niche differentiation. We demonstrated theoretically that differential sensitivity of sexual and facultatively asexual aphids to an environmental parameter (mating host suitability) shapes the two strategies: whereas the sexual lineages switch earlier to the production of sexual forms, the facultative asexual lineages delay and spread out their investment in sexual reproduction. This predicted pattern of niche specialization is in agreement with the temporal structure revealed in natura by demographic and genetic data. We propose that partial loss of sex by one pool of aphids and subsequent reduction in gene flow between lineages may favour temporal specialization through disruptive selection.
 
Article
Using Drosophila melanogaster, we explore the consequences of restricted panmixia (RP) on the genetic load caused by segregating deleterious recessive alleles in a population where females mate a full sib with probability about (1/2) and mate randomly otherwise. We find that this breeding structure purges roughly half the load concealed in heterozygous condition. Furthermore, fitness did not increase after panmixia was restored, implying that, during RP, the excess of expressed load induced by inbreeding had also been efficiently purged. We find evidences for adaptation to laboratory conditions and to specific selective pressures imposed by the RP protocol. We discuss some of the consequences of these results, both for the evolution of population breeding structures and for the design of conservation programmes.
 
Article
Environmental variation in temperature can have dramatic effects on plant morphology, phenology, and fitness, and for this reason it is important to understand the evolutionary dynamics of phenotypic plasticity in response to temperature. We investigated constraints on the evolution of phenotypic plasticity in response to a temperature gradient in the model plant Arabidopsis thaliana by applying modern analytical tools to the classic data of Westerman & Lawrence (1970). We found significant evidence for two types of constraints. First, we detected numerous significant genetic correlations between plastic responses to temperature and the mean value of a trait across all environments, which differed qualitatively in pattern between the set of ecotypes and the set of mutant lines in the original sample. Secondly, we detected significant costs of flowering time plasticity in two of the three experimental environments, and a net pattern of selection against flowering time plasticity in the experiment overall. Thus, when explored with contemporary methods, the prescient work of Westerman & Lawrence (1970) provides new insights about evolutionary constraints on the evolution of plasticity.
 
Article
Ecological selection against hybrids between populations occupying different habitats might be an important component of reproductive isolation during the initial stages of speciation. The strength and directionality of this barrier to gene flow depends on the genetic architecture underlying divergence in ecologically relevant phenotypes. We here present line cross analyses of inheritance for two key foraging-related morphological traits involved in adaptive divergence between stickleback ecotypes residing parapatrically in lake and stream habitats within the Misty Lake watershed (Vancouver Island, Canada). One main finding is the striking genetic dominance of the lake phenotype for body depth. Selection associated with this phenotype against first- and later-generation hybrids should therefore be asymmetric, hindering introgression from the lake to the stream population but not vice versa. Another main finding is that divergence in gill raker number is inherited additively and should therefore contribute symmetrically to reproductive isolation. Our study suggests that traits involved in adaptation might contribute to reproductive isolation qualitatively differently, depending on their mode of inheritance.
 
Article
The increase in diversity towards the equator arises from latitudinal variation in rates of cladogenesis, extinction, immigration and/or emigration of taxa. We tested the relative contribution of all four processes to the latitudinal gradient in 26 marine invertebrate orders with extensive fossil records, examined previously by David Jablonski. Coupling Jablonski's estimates of latitudinal variation in cladogenesis with new data on patterns of extinction and current distributions, we show that the present-day gradient in diversity is caused by higher rates of cladogenesis and subsequent range expansion (immigration) at lower latitudes. In contrast, extinction and emigration were not important in the creation of the latitudinal gradient in ordinal richness. This work represents one of the first simultaneous tests of the role of all four processes in the creation of the latitudinal gradient in taxonomic richness, and suggests that low tropical extinction rates are not essential to the creation of latitudinal diversity gradients.
 
Article
Some mitochondrial introgression is common between closely related species, but distinct species rarely show substantial introgression in their entire distribution range. In this study, however, we report a complete lack of mitochondrial divergence between two sympatric species of flat periwinkles (Littorina fabalis and Littorina obtusata) which, based on previous allozyme studies, diverged approximately 1 Ma. We re-examined their species status using both morphology (morphometric analysis) and neutral genetic markers (microsatellites) and our results confirmed that these species are well separated. Despite this, the two species shared all common cytochrome-b haplotypes throughout their NE Atlantic distribution and no deep split between typical L. fabalis and L. obtusata haplotypes could be found. We suggest that incomplete lineage sorting explains most of the lack of mitochondrial divergence between these species. However, coalescent-based analyses and the sympatric sharing of unique haplotypes suggest that introgressive hybridization also has occurred.
 
Article
Several statements by Pouydebat et al. (2008) do not adequately represent views of authors cited, in part because they reflect confusion in the literature about terminology regarding precision gripping. We address these problems, by tracing definitions of precision grips through the literature on manipulative behaviour and identifying the grip that is central to the Pouydebat et al. (2008) study. This allows us to offer a clarification of the statements by Pouydebat et al. (2008) regarding the sequence of appearance of human grip capabilities and possible morphological correlates to these capabilities in extant species.
 
Article
A recent article presents a study of pollinator visitation behaviour that is used to evaluate the selective pressure that pollinator visitation rate might have on the timing of the production of nonrewarding flowers. Here we take issue with the conclusions of the paper that there should be selection pressure for nonrewarding flowers to be available earlier in the season in order to avoid dissimilar sympatric rewarding species. Consideration of selection pressure must take into account temporal variation in total pollinator availability, pollinator longevity and unlearned response, and the stability of plant communities over time, as well as the learned responses of individual pollinators that the original study focused on. Learning alone would not necessarily select for early flowering by nonrewarders if temporal variation in pollinator numbers is strong or naïve pollinators consistently appear throughout the flowering season. Further, we argue that early flowering could simply be a natural corollary of longevity of flowers needed to combat negative frequency-dependent selection and low overall visitation rates by pollinators, rather than a trait that has been specifically selected to reduce temporal overlap with competing rewarding species.
 
Between-population differentiation measured by the ratio of between- to within-population genotypic variance  as a function of mean within-population genotypic variance  in different landscapes. Lines are analytical predictions from eqn 13, for different values of the net migration rate. Open (filled) symbols correspond to simulation results with pure pollen (seed) dispersal, for two values of the genomic mutation rate (U = 10−2, 10−1) and five values of the net migration rate; crosses and continuous lines: ; squares and dotted lines: ); circles and short-dashed lines: ); triangles and dot-dashed lines: ); diamonds and long-dashed lines: ). Left panels (a,c,e) show moderately fragmented landscapes of n = 4 populations of N = 200 individuals. Right panels (b,d,f) show highly fragmented landscapes of n = 32 populations of N = 25 individuals. Habitat heterogeneity increases from top to bottom: (a,b) Δθ = 0, (c,d) Δθ = 1, (e,f) Δθ = 3. Free recombination between loci. Confidence intervals are smaller than size of symbols. Confidence intervals on the x-axis range from 10−4 to 9 × 10−3. Some simulation points are missing (20 over 120 parameters sets) due to lack of convergence to a stable equilibrium (see Appendix S2). Missing points correspond to the following parameter sets: (N = 25, n=32, , for all values of the seed dispersal rate, mutation rate and habitat heterogeneity) (N = 200, , U = 10−1, for all values of the seed dispersal rate and habitat heterogeneity) (N = 200, , U = 10−2, Δθ = 3 only for pure pollen dispersal) (N = 200, , U = 10−1, Δθ = 3 only for pure seed dispersal).
Mean genetic load as a function of mean within-population genotypic variance  in different landscapes. Lines are analytical predictions from eqn 15, for different values of the net migration rate. Open (filled) dots correspond to simulation results with pure pollen (seed) dispersal for two values of the mutation rate (U = 10−2, 10−1) and five values of the migration rate (same values of the migration rate and associated symbols as in ). Left panels (a,c,e) show moderately fragmented landscapes of n = 4 populations of N = 200 individuals. Right panels (b,d,f) show highly fragmented landscapes of n = 32 populations of N = 25 individuals. Habitat heterogeneity increases from top to bottom: (a,b) Δθ = 0, (c,d) Δθ = 1, (e,f) Δθ = 3 Free recombination between loci. Confidence intervals are smaller than size of symbols. Confidence intervals on the x-axis range from 10−4 to 9 × 10−3. Missing data points are as in .
Within-population genotypic variance and its components as a function of landscape heterogeneity and dispersal mode in a highly fragmented landscape of 32 populations of 25 individuals. Open (filled) symbols correspond to pure pollen (pure seed) dispersal. (a) Total within-population genotypic variance (), (b) mean genic variance at HWE (), (c) mean part of the genotypic variance contributed by gametic linkage disequilibrium at HWE (), (d) mean part of the variance due to deviations of genotypic frequencies with respect to HWE (). See eqns 7–10 for definitions. Genomic mutation rate U = 0.01. Free recombination between loci. Net migration rate . Confidence intervals are smaller than plotting symbols.
Within-population genotypic variance and its components as a function of seed migration rate ms in a highly fragmented landscape of 32 populations of 25 individuals. (a) Top panel shows mean within-population total genotypic variance  for different levels of landscape heterogeneity; continuous line and open circles: homogeneous landscape (Δθ = 0); dashed line and open triangles: slightly heterogeneous landscape (Δθ = 1); dotted line and crosses: highly heterogeneous landscape (Δθ = 3); the point on the left shows the level of within-population genotypic variance without migration; (b,c,d) bottom panels show within-population genotypic variance components as a function of the net seed migration rate; white: genic variance at HWE (); grey: part of the genotypic variance contributed by gametic linkage disequilibrium at HWE (); hatched: part of the variance due to deviations from HWE (). See eqns 7–10 for definitions. Landscape heterogeneity increases from left to right. Pure seed dispersal: mp = 0. Genomic mutation rate U = 0.01. Free recombination between loci.
Partition of the within-population genotypic variance as a function of landscape heterogeneity, mutation and recombination rates in a highly fragmented landscape of 4 populations of 200 individuals. F = free recombination between loci. L = all loci are linked on the same chromosome with a recombination probability of 0.01 between adjacent loci. In each panel, the two first bars correspond to a genomic mutation rate U = 10−2, and U = 10−1 for the latter two bars. White, grey and hatched bars, respectively, correspond to the mean genic variance at HWE , the mean part of the genotypic variance contributed by gametic linkage disequilibrium at HWE () and the mean part of the variance due to deviations of genotypic frequencies with respect to HWE (). Dots indicate the mean within-population genotypic variance . Habitat heterogeneity measured by the difference in optimal phenotypic values in the two habitats increases from left to right: Δθ = 0, Δθ = 1 and Δθ = 3. Migration occurs through seeds only, with . Confidence intervals are plotted for  only and are smaller than symbols.
Article
Evolution of local adaptation depends critically on the level of gene flow, which, in plants, can be due to either pollen or seed dispersal. Using analytical predictions and individual-centred simulations, we investigate the specific influence of seed and pollen dispersal on local adaptation in plant populations growing in patchy heterogeneous landscapes. We study the evolution of a polygenic trait subject to stabilizing selection within populations, but divergent selection between populations. Deviations from linkage equilibrium and Hardy-Weinberg equilibrium make different contributions to genotypic variance depending on the dispersal mode. Local genotypic variance, differentiation between populations and genetic load vary with the rate of gene flow but are similar for seed and pollen dispersal, unless the landscape is very heterogeneous. In this case, genetic load is higher in the case of pollen dispersal, which appears to be due to differences in the distribution of genotypic values before selection.
 
Top-cited authors
Roger K Butlin
  • The University of Sheffield
Anders Pape Moller
  • French National Centre for Scientific Research
Michael D Jennions
  • Australian National University
Nicolas Bierne
  • Université de Montpellier
James Mallet
  • UCL, and Harvard University