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Genetic divergence of peripherally disjunct populations of the gastropod Batillariella estuarina in the Houtman Abrolhos Islands, Western Australia.

Geographically disjunct populations are unusual in marine species, but the Houtman Abrolhos Islands, Western Australia, provide opportunities to study highly disjunct peripheral isolates of several species. The intertidal snail Batillariella estuarina occurs in isolated tidal ponds in the Abrolhos Islands, where it is at its northern limit, disjunct from mainland populations by 600-900 km. The species is thus disjunct both geographically and among the peripherally isolated populations in the Abrolhos Islands. Comparisons of allozymes at 11 polymorphic loci were made among populations from 10 ponds in the Abrolhos Islands and six sites from relatively continuous tidal flats at Albany, 900 km away, the nearest major set of populations. Among all 16 populations, subdivision was high (FST = 0.455). Although there were subtle differences between the geographical regions, the large majority of divergence occurred among the isolated ponds in the Abrolhos (FST = 0.441), and divergence on the tidal flats at Albany was only moderate (FST = 0.085). Characteristic of peripheral isolates, the pond populations have less polymorphism and fewer alleles than the more connected populations at Albany. Combined with evidence of genetic divergence in the gastropods Bembicium vittatum and Austrocochlea constricta, which have very similar geographical distributions to that of B. estuarina, these results indicate the potential evolutionary significance of peripherally isolated marine populations in the unusual habitats of the Abrolhos Islands.

Alleles↗

Amount and structure of intra- and interspecific genetic variation in the moss genus Polytrichum.

Allozyme electrophoresis was used to determine amount and structure of genetic variation within and between five congeneric haploid moss species: Polytrichum formosum, P. commune, P. uliginosum, P. piliferum and P. juniperinum. For the different species, gene diversity within populations (HS) ranged from very low (0.025) to moderate values (0.127), being, on average, lower than those observed for vascular plants and many other moss species. Polytrichum piliferum and P. juniperinum showed significantly higher levels of HS than the other species examined, which possibly might be explained by sexual reproduction being more prevalent in these two species, that often live in more dynamic habitats, where turnover of individuals is more frequent. Genetic variability was observed to be structured in contrasting ways at different levels. At the intraspecific level genetic differentiation among populations of most Polytrichum species was low, FST </= 0.1, indicating a considerable degree of gene flow by wind-dispersed spores over substantial distances. At the interspecific level strong divergence, genetic identities being on average I=0.222, was observed between most species studied, despite low levels of divergence at the morphological level. These I-values are significantly lower than observed for congeneric vascular plant species and most congeneric bryophyte species. This suggests that the morphological conservatism is not due to lack of genetic variability and evolutionary potential.

Genetic Variation↗

Lack of response to artificial selection on the slope of reaction norms for seasonal polyphenism in the butterfly Bicyclus anynana.

The tropical butterfly Bicyclus anynana shows adaptive phenotypic plasticity in response to wet-dry seasonality. The wet season form (WSF) has a conspicuous wing pattern with large eyespots, whereas the dry season form (DSF) lacks eyespots and therefore has a more cryptic appearance. Temperature is the main factor controlling this difference: rearing larvae at a low (<19 degrees C) temperature in the laboratory results in the DSF, whereas rearing at a high (>23 degrees C) temperature induces the WSF. We applied truncation selection in opposite directions in successive generations reared at two alternating temperatures (18.5 degrees C and 23.5 degrees C) to increase (for two High Plasticity (HP) lines), and decrease (for two Low Plasticity (LP) lines) wing pattern plasticity. Plasticity was assessed by partitioning full-sib families over four rearing temperatures (18.5 degrees C, 20.5 degrees C, 21.5 degrees C and 23.5 degrees C). Several wing pattern elements were measured for which the first principal component (PC1) provides a useful summary. The slopes of reaction norms for PC1 were significantly steeper in the HP lines than in the LP lines; however, the selection lines did not always differ significantly from the unselected stock. The results of crosses between the replicates of the selection lines gave no indication for effects of inbreeding. We argue that high, positive genetic correlations across temperatures retard a response to selection in opposite directions in different environments. This is discussed with respect to potential evolutionary constraints in natural populations in these butterflies.

Adaptation, Biological↗

Does natural selection promote population divergence? A comparative analysis of population structure using amplified fragment length polymorphism markers and quantitative traits.

Divergent natural selection is considered an important force in plant evolution leading to phenotypic differentiation between populations exploiting different environments. Extending an earlier greenhouse study of population differentiation in the selfing annual plant Senecio vulgaris, we estimated the degree of population divergence in several quantitative traits related to growth and life history and compared these estimates with those based on presumably neutral molecular markers (amplified fragment length polymorphisms; AFLPs). This approach allowed us to disentangle the effects of divergent selection from that of other evolutionary forces (e.g. genetic drift). Five populations were examined from each of two habitat types (ruderal and agricultural habitats). We found a high proportion of total genetic variance to be among populations, both for AFLP markers (phiST = 0.49) and for quantitative traits (range of QST: 0.26-0.77). There was a strong correlation between molecular and quantitative genetic differentiation between pairs of populations (Mantel's r = 0.59). However, estimates of population differentiation in several quantitative traits exceeded the neutral expectation (estimated from AFLP data), suggesting that divergent selection contributed to phenotypic differentiation, especially between populations from ruderal and agricultural habitats. Estimates of within-population variation in AFLP markers and quantitative genetic were poorly correlated, indicating that molecular marker data may be of limited value to predict the evolutionary potential of populations of S. vulgaris.

DNA, Plant↗

Nuclear DNA analyses in genetic studies of populations: practice, problems and prospects.

Population-genetic studies have been remarkably productive and successful in the last decade following the invention of PCR technology and the introduction of mitochondrial and microsatellite DNA markers. While mitochondrial DNA has proven powerful for genealogical and evolutionary studies of animal populations, and microsatellite sequences are the most revealing DNA markers available so far for inferring population structure and dynamics, they both have important and unavoidable limitations. To obtain a fuller picture of the history and evolutionary potential of populations, genealogical data from nuclear loci are essential, and the inclusion of other nuclear markers, i.e. single copy nuclear polymorphic (scnp) sequences, is clearly needed. Four major uncertainties for nuclear DNA analyses of populations have been facing us, i.e. the availability of scnp markers for carrying out such analysis, technical laboratory hurdles for resolving haplotypes, difficulty in data analysis because of recombination, low divergence levels and intraspecific multifurcation evolution, and the utility of scnp markers for addressing population-genetic questions. In this review, we discuss the availability of highly polymorphic single copy DNA in the nuclear genome, describe patterns and rate of evolution of nuclear sequences, summarize past empirical and theoretical efforts to recover and analyse data from scnp markers, and examine the difficulties, challenges and opportunities faced in such studies. We show that although challenges still exist, the above-mentioned obstacles are now being removed. Recent advances in technology and increases in statistical power provide the prospect of nuclear DNA analyses becoming routine practice, allowing allele-discriminating characterization of scnp loci and microsatellite loci. This certainly will increase our ability to address more complex questions, and thereby the sophistication of genetic analyses of populations.

Animals↗

Hybrid origin of a cichlid population in Lake Malawi: implications for genetic variation and species diversity.

The importance of species recognition to taxonomic diversity among Lake Malawi cichlids has been frequently discussed. Hybridization - the apparent breakdown of species recognition - has been observed sporadically among cichlids and has been viewed as both a constructive and a destructive force with respect to species diversity. Here we provide genetic evidence of a natural hybrid cichlid population with a unique colour phenotype and elevated levels of genetic variation. We discuss the potential evolutionary consequences of interspecific hybridization in Lake Malawi cichlids and propose that the role of hybridization in generating both genetic variability and species diversity of Lake Malawi cichlids warrants further consideration.

Animals↗

Heritability of resistance against ectoparasitism in the Drosophila-Macrocheles system.

Ectoparasites are abundant in natural communities, can have pronounced deleterious fitness consequences to their host and are important vectors of transmissible parasitic disease. Yet very few studies have estimated the magnitude of heritable genetic variation underlying resistance against ectoparasitism, which significantly limits our ability to predict the evolution of this ecologically important character. The present paper reports results of artificial selection for increased resistance in Drosophila nigrospiracula against ectoparasitic, haematophagous mites, Macrocheles subbadius. In this system, which occurs naturally in the Sonoran Desert of North America, ectoparasitism significantly damages the expression of host fitness traits, including longevity, fecundity and male mating success. In the present study, resistance, which was modelled as a threshold trait, responded significantly to selection applied on either sex. Realized heritability, calculated as a mean across four replicates, was estimated to be 0.152 +/- 0.014 (SE). The heritability estimate from selection on males did not differ from that on females, but both estimates differed significantly from zero. This documented presence of additive genetic variation for resistance, coupled with knowledge of the fitness consequences of ectoparasitism, indicates that the host population possesses significant evolutionary potential. Selection was applied on the pre-attachment phase, thereby targeting behavioural forms of defence. This study therefore establishes parallels between insects and other animals in their ability to protect themselves and evolve behavioural defences against ectoparasites.

Animals↗

Phylogenetic analysis of the ecological correlates of dioecy in angiosperms.

We report on a phylogenetic analysis of correlations between the occurrence of dioecy and several ecological and life-history attributes: tropical distribution, woody growth form, abiotic pollination, small inconspicuous flowers and inflorescences, many-flowered inflorescences and fleshy fruits. Various hypotheses have been proposed to explain why associations occur between dioecy and several of these attributes, yet most assume that dioecy originates more often in clades with these traits than in clades with alternative character states. To investigate correlations between dioecy and these attributes, and to provide insights into the potential evolutionary pathways that have led to these associations, we assigned states of these traits to genera on a large-scale molecular phylogeny of the angiosperms; we then used maximum-likelihood analysis to analyse the presence of correlations and the sequence of acquisition of traits. Phylogenetic analysis revealed correlations between dioecy and six of the seven attributes; only many-flowered inflorescences exhibiting no association with the dioecious condition. The particular correlations that were revealed and the strength of the association differed among the three main monophyletic groups of angiosperms (Rosids, Asterids, and Eumagnoliids). Our analysis provided no general support for the hypothesis that dioecy is more likely to evolve in lineages already possessing the seven attributes we considered. Further analysis of the intercorrelations of the seven attributes provided evidence for non-independence between some of the traits, implying that functional associations among these traits have influenced the ecology and evolution of dioecious species.

Ecology↗

Double-segment defining role of even-skipped homologs along the evolution of insect pattern formation.

Recent studies on insect patterning suggest that the genetic hierarchy may be roughly conserved in phylogenetically divergent species, but pair-rule genes may not function identically in all insects. In order to understand potential evolutionary changes in the role of the pair-rule genes, a Bombyx even-skipped homolog was cloned and its expression pattern during early embryogenesis studied. Eight stripes of Bombyx even-skipped were progressively expressed in an antero-posterior order. Later, these stripes disappeared anteriorly. Under this detection system, Bombyx even-skipped stripes clearly do not resolve into the corresponding secondary stripes, an obvious difference from Drosophila and Tribolium. These results suggest that Bombyx even-skipped may serve a double-segment defining role and may determine the odd-numbered engrailed stripes.

Amino Acid Sequence↗

Developmental flexibility and the effect of social environment on fertility and fecundity in parthenogenetic reproduction.

One specialized environment that can influence development arises in the context of social interactions, including the environment contributed by a sexual partner during sexual reproduction. It is often difficult, however, to separate out the effect of mating (fertilization) from the effect of social environment. In the study reported here we examine the effect of social environment mediated by a pheromonal signal on the fertility and fecundity of the facultatively parthenogenetic cockroach Nauphoeta cinerea. By examining parthenogenetically reproducing females, we isolate the effects of social environment in the absence of mating or fertilization. Females exposed to male odors are more likely to produce parthenogenetic offspring. Further, increased exposure to the male pheromone increases the number of offspring produced. Variation in timing of reproduction is also dependent on the male. Thus, social environments are a mechanism by which males contribute to the development of their offspring, resulting in variation in development. This study illustrates the potential evolutionary importance of social environments in development, because a requirement for male-contributed environments may be a constraint to evolving asexual reproduction from a sexually reproducing species.

Analysis of Variance↗

Rapid evolution of a polyphenic threshold.

Polyphenisms are thought to play an important role in the evolution of phenotypic diversity and the origin of morphological and behavioral novelties. However, the extent to which polyphenic developmental mechanisms evolve in natural populations is unknown. Here we contrast patterns of male phenotype expression in native and exotic and ancestral and descendant populations of the horn polyphenic beetle, Onthophagus taurus. Males in this species express two alternative morphologies in response to larval feeding conditions. Favorable conditions cause males to grow larger than a threshold body size and to develop a pair of horns on their heads. Males that encounter relatively poor conditions do not reach this threshold size and remain hornless. We show that exotic and native populations of O. taurus differ significantly in the body size threshold that separates alternative male phenotypes. Comparison with archival museum collections and additional samples obtained from the native range of O. taurus suggests that allometric differences between exotic and native populations do not reflect preexisting variation in the native range of this species. Instead, our data suggest that threshold divergences between exotic and native populations have evolved in less than 40 years since the introduction to a new habitat and have proceeded in opposite directions in two exotic ranges of this species. Finally, we show that the kind and magnitude of threshold divergence between native and exotic populations are similar to differences normally observed between species. Our results support the view that certain components of the developmental control mechanism that underlie polyphenic development can evolve rapidly in natural populations and may provide important avenues for phenotypic differentiation and diversification in nature. We discuss the role of developmental control mechanisms in the origin of allometric diversification and explore potential evolutionary mechanisms that could drive scaling relationship evolution in nature.

Adaptation, Biological↗

Most species are not driven to extinction before genetic factors impact them.

There is controversy concerning the role of genetic factors in species extinctions. Many authors have asserted that species are usually driven to extinction before genetic factors have time to impact them, but few studies have seriously addressed this issue. If this assertion is true, there will be little difference in genetic diversity between threatened and taxonomically related nonthreatened species. We compared average heterozygosities in 170 threatened taxa with those in taxonomically related nonthreatened taxa in a comprehensive metaanalysis. Heterozygosity was lower in threatened taxa in 77% of comparisons, a highly significant departure from the predictions of the no genetic impact hypothesis. Heterozygosity was on average 35% lower (median 40%) in threatened taxa than in related nonthreatened ones. These differences in heterozygosity indicate lowered evolutionary potential, compromised reproductive fitness, and elevated extinction risk in the wild. Independent evidence from stochastic computer projections has demonstrated that inbreeding depression elevates extinction risk for threatened species in natural habitats when all other threatening processes are included in the models. Thus, most taxa are not driven to extinction before genetic factors affect them adversely.

Animals↗

Herbivores cause a rapid increase in hereditary symbiosis and alter plant community composition.

Microbial symbioses are ubiquitous in nature. Hereditary symbionts warrant particular attention because of their direct effects on the evolutionary potential of their hosts. In plants, hereditary fungal endophytes can increase the competitive ability, drought tolerance, and herbivore resistance of their host, although it is unclear whether or how these ecological benefits may alter the dynamics of the endophyte symbiosis over time. Here, we demonstrate that herbivores alter the dynamics of a hereditary symbiont under field conditions. Also, we show that changes in symbiont frequency were accompanied by shifts in the overall structure of the plant community. Replicated 25-m2 plots were enriched with seed of the introduced grass, Lolium arundinaceum at an initial frequency of 50% infection by the systemic, seed-transmitted endophyte Neotyphodium coenophialum. Over 54 months, there was a significantly greater increase in endophyte-infection frequency in the presence of herbivores (30% increase) than where mammalian and insect herbivory were experimentally reduced by fencing and insecticide application (12% increase). Under ambient mammalian herbivory, the above-ground biomass of nonhost plant species was reduced compared with the mammal-exclusion treatment, and plant composition shifted toward greater relative biomass of infected, tall fescue grass. These results demonstrate that herbivores can drive plant-microbe dynamics and, in doing so, modify plant community structure directly and indirectly.

Animals↗

Mimicking natural evolution in metallo-beta-lactamases through second-shell ligand mutations.

Metallo-beta-lactamases (MBLs) represent the latest generation of beta-lactamases. The structural diversity and broad substrate profile of MBLs allow them to confer resistance to most beta-lactam antibiotics. To explore the evolutionary potential of these enzymes, we have subjected the Bacillus cereus MBL (BcII) to a directed evolution scheme, which resulted in an increased hydrolytic efficiency toward cephalexin. A systematic study of the hydrolytic profile, substrate binding, and active-site features of the evolved lactamase reveal that directed evolution has shaped the active site by means of remote mutations to better hydrolyze cephalosporins with small, uncharged C-3 substituents. One of these mutations is found in related enzymes from pathogenic bacteria and is responsible for the increase in that enzyme's hydrolytic profile. The mutations lowered the activation energy of the rate-limiting step rather than improved the affinity of the enzyme toward these substrates. The following conclusions can be made: (i) MBLs are able to expand their substrate spectrum without sacrificing their inherent hydrolytic capabilities; (ii) directed evolution is able to mimic mutations that occur in nature; (iii) the metal-ligand strength is tuned by second-shell mutations, thereby influencing the catalytic efficiency; and (iv) changes in the position of the second Zn(II) ion in MBLs affect the substrate positioning in the active site. Overall, these results show that the evolution of enzymatic catalysis can take place by remote mutations controlling reactivity.

Amino Acid Substitution↗

Variable host-pathogen compatibility in Mycobacterium tuberculosis.

Mycobacterium tuberculosis remains a major cause of morbidity and mortality worldwide. Studies have reported human pathogens to have geographically structured population genetics, some of which have been linked to ancient human migrations. However, no study has addressed the potential evolutionary consequences of such longstanding human-pathogen associations. Here, we demonstrate that the global population structure of M. tuberculosis is defined by six phylogeographical lineages, each associated with specific, sympatric human populations. In an urban cosmopolitan environment, mycobacterial lineages were much more likely to spread in sympatric than in allopatric patient populations. Tuberculosis cases that did occur in allopatric hosts disproportionately involved high-risk individuals with impaired host resistance. These observations suggest that mycobacterial lineages are adapted to particular human populations. If confirmed, our findings have important implications for tuberculosis control and vaccine development.

Adaptation, Physiological↗

Ameiotic recombination in asexual lineages of Daphnia.

Despite the enormous theoretical attention given to the evolutionary consequences of sexual reproduction, the validity of the key assumptions on which the theory depends rarely has been evaluated. It is often argued that a reduced ability to purge deleterious mutations condemns asexual lineages to an early extinction. However, most well characterized asexual lineages fail to exhibit the high levels of neutral allelic divergence expected in the absence of recombination. With purely descriptive data, it is difficult to evaluate whether this pattern is a consequence of the rapid demise of asexual lineages, an unusual degree of mutational stability, or recombination. Here, we show in mutation-accumulation lines of asexual Daphnia that the rate of loss of nucleotide heterozygosity by ameiotic recombination is substantially greater than the rate of introduction of new variation by mutation. This suggests that the evolutionary potential of asexual diploid species is not only a matter of mutation accumulation and reduced efficiency of selection, and [corrected] it underscores the limited utility of using neutral allelic divergence as an indicator of ancient asexuality.

Animals↗

Genomic signatures of dairy adaptation in Saccharomyces cerevisiae from traditional Yaghnob goat-cheese fermentation.

The growing interest in studying Saccharomyces cerevisiae strains from previously unexplored niches is greatly expanding our understanding of this yeast's ecology and evolution. While strains involved in alcoholic fermentation are the most studied, S. cerevisiae has also been isolated from milk fermentations and their products, suggesting a potential evolutionary specialization for dairy environments. These fermentations are characterized by the predominant presence of lactose, a carbon source that S. cerevisiae cannot metabolize directly but can exploit through the enzymatic activity of co-occurring microorganisms that convert lactose into fermentable substrates, such as glucose and galactose. In this study, we analyzed S. cerevisiae strains isolated from an unexplored and remote niche: traditional goat fermented milk produced by the Yaghnob people, an ethnically and geographically partly isolated population living in the Upper Zarafshan area of the Republic of Tajikistan. Comparative analyses with published S. cerevisiae genomes positioned the Yaghnob strains at the base of the phylogenetic dairy clade. These strains revealed distinctive coding sequences and strain-specific single-nucleotide variants present in all Yaghnob strains but absent from the other 1,053 strains analyzed. Further investigation of variants in key genes involved in galactose metabolism provided insights into the genomic and protein-level evolution of Yaghnob strains, uncovering unique genomic signatures of adaptation to the dairy environment.

Saccharomyces cerevisiae↗

The genetic system in parthenogenetic strains of Drosophila mercatorum.

Five parthenogenetic laboratory strains of Drosophila mercatorum have been examined for their mode of diploidization in the eggs of females. Diploidization occurs either by pronuclear duplication leading to complete homozygosity or by nuclear fusion. If the latter occurs between pronuclei from different secondary oocytes, heterozygosity can be maintained. Each of the five strains shows some nuclear fusion and three show significantly different amounts, ranging from about 21% to less than 1%. Even in strains with a strong tendency towards homozygosity, it is argued that this dual system of nuclear reconstitution provides for genetic recombination; thus, the strains retain considerable evolutionary potential.

Animals↗