Variation In Selection Pressures Research Articles

The evolutionary cost of homophily: Social stratification facilitates stable variant coexistence and increased rates of evolution in host-associated pathogens.

Coexistence of multiple strains of a pathogen in a host population can present significant challenges to vaccine development or treatment efficacy. Here we discuss a novel mechanism that can increase rates of long-lived strain polymorphism, rooted in the presence of social structure in a host population. We show that social preference of interaction, in conjunction with differences in immunity between host subgroups, can exert varying selection pressure on pathogen strains, creating a balancing mechanism that supports stable viral coexistence, independent of other known mechanisms. We use population genetic models to study rates of pathogen heterozygosity as a function of population size, host population composition, mutant strain fitness differences and host social preferences of interaction. We also show that even small periodic epochs of host population stratification can lead to elevated strain coexistence. These results are robust to varying social preferences of interaction, overall differences in strain fitnesses, and spatial heterogeneity in host population composition. Our results highlight the role of host population social stratification in increasing rates of pathogen strain diversity, with effects that should be considered when designing policies or treatments with a long-term view of curbing pathogen evolution.

Archaic introgression contributed to shape the adaptive modulation of angiogenesis and cardiovascular traits in human high-altitude populations from the Himalayas.

It is well established that several Homo sapiens populations experienced admixture with extinct human species during their evolutionary history. Sometimes, such a gene flow could have played a role in modulating their capability to cope with a variety of selective pressures, thus resulting in archaic adaptive introgression events. A paradigmatic example of this evolutionary mechanism is offered by the EPAS1 gene, whose most frequent haplotype in Himalayan highlanders was proved to reduce their susceptibility to chronic mountain sickness and to be introduced in the gene pool of their ancestors by admixture with Denisovans. In this study, we aimed at further expanding the investigation of the impact of archaic introgression on more complex adaptive responses to hypobaric hypoxia evolved by populations of Tibetan/Sherpa ancestry, which have been plausibly mediated by soft selective sweeps and/or polygenic adaptations rather than by hard selective sweeps. For this purpose, we used a combination of composite-likelihood and gene network-based methods to detect adaptive loci in introgressed chromosomal segments from Tibetan WGS data and to shortlist those presenting Denisovan-like derived alleles that participate to the same functional pathways and are absent in populations of African ancestry, which are supposed to do not have experienced Denisovan admixture. According to this approach, we identified multiple genes putatively involved in archaic introgression events and that, especially as regards TBC1D1, RASGRF2, PRKAG2, and KRAS, have plausibly contributed to shape the adaptive modulation of angiogenesis and of certain cardiovascular traits in high-altitude Himalayan peoples. These findings provided unprecedented evidence about the complexity of the adaptive phenotype evolved by these human groups to cope with challenges imposed by hypobaric hypoxia, offering new insights into the tangled interplay of genetic determinants that mediates the physiological adjustments crucial for human adaptation to the high-altitude environment.

Archaic introgression contributed to shape the adaptive modulation of angiogenesis and cardiovascular traits in human high-altitude populations from the Himalayas

It is well established that several Homo sapiens populations experienced admixture with extinct human species during their evolutionary history. Sometimes, such a gene flow could have played a role in modulating their capability to cope with a variety of selective pressures, thus resulting in archaic adaptive introgression events. A paradigmatic example of this evolutionary mechanism is offered by the EPAS1 gene, whose most frequent haplotype in Himalayan highlanders was proved to reduce their susceptibility to chronic mountain sickness and to be introduced in the gene pool of their ancestors by admixture with Denisovans. In this study, we aimed at further expanding the investigation of the impact of archaic introgression on more complex adaptive responses to hypobaric hypoxia evolved by populations of Tibetan/Sherpa ancestry, which have been plausibly mediated by soft selective sweeps and/or polygenic adaptations rather than by hard selective sweeps. For this purpose, we used a combination of composite-likelihood and gene network-based methods to detect adaptive loci in introgressed chromosomal segments from Tibetan WGS data and to shortlist those presenting Denisovan-like derived alleles that participate to the same functional pathways and are absent in populations of African ancestry, which are supposed to do not have experienced Denisovan admixture. According to this approach, we identified multiple genes putatively involved in archaic introgression events and that, especially as regards TBC1D1, RASGRF2, PRKAG2, and KRAS, have plausibly contributed to shape the adaptive modulation of angiogenesis and of certain cardiovascular traits in high-altitude Himalayan peoples. These findings provided unprecedented evidence about the complexity of the adaptive phenotype evolved by these human groups to cope with challenges imposed by hypobaric hypoxia, offering new insights into the tangled interplay of genetic determinants that mediates the physiological adjustments crucial for human adaptation to the high-altitude environment.

Intraspecific plant chemodiversity at the individual and plot levels influences flower visitor groups with consequences for germination success

Abstract Chemodiversity, the diversity of specialised (secondary) metabolites, plays a crucial role in mediating interactions between plants and animals, including insect herbivores and mutualists. Chemodiversity can be observed at both the individual and the population level. However, the impacts of chemodiversity at these two levels on interactions between plants and floral visitors, mainly pollinators and floral herbivores (florivores), are only poorly understood. This study aimed to investigate the effects of chemodiversity at both individual (chemotype) and plot level on pollinators and florivores, examining their mutual interactions. To investigate these questions, we used individuals of the perennial Tanacetum vulgare differing in their terpenoid composition, representing different chemotypes. We planted individuals of five distinct chemotypes in a common garden design with homogeneous (five plants of the same chemotype) and heterogeneous (five different chemotypes) plots and observed flower visits in dependence of individual chemotype and plot type. Seeds were collected from a subset of plants and germination rates determined. Our findings revealed that chemodiversity at the plot level significantly influenced pollinator visitation, with more visits on plants of heterogeneous plots. We also observed marginally more pollinators on one of the chemotypes grown in homogeneous plots. In contrast, chemotype but not plot type had a significant effect on florivore visits. Pollinator and florivore species richness did not vary with plot type. Furthermore, a negative correlation was observed between pollinator and florivore visits in 1 year. Germination rates were positively correlated with pollinator visits and affected by chemotype but not by florivore visits or plot type. Synthesis. Our study emphasises the significance of the spatial level at which plant chemistry is perceived by flower visitors, potentially influencing the germination success of plants. Exploring the ecology of these visitors and the varying selection pressures they exert on floral chemistry can help to elucidate the evolutionary processes that maintain chemodiversity in natural environments. Read the free Plain Language Summary for this article on the Journal blog.

Assessment of Genetic Diversity and the Population Structure of Species from the Fusarium fujikuroi Species Complex Causing Fusarium Stalk Rot of Maize.

Fusarium stalk rot (FSR), caused by the Fusarium species complex, is an economic threat to maize cultivation all over the world. We investigated the population structure and genetic diversity of Fusarium species obtained from five major maize-growing regions in India. The Tef-1α locus was used for phylogenetic analysis of geographically distinct isolates of Fusarium verticillioides, F. andiyazi, F. proliferatum, F. nygamai, and F. acutatum causing FSR. A phylogenetic tree showed monophyletic, polyphyletic, and paraphyletic groupings reflecting the complex evolutionary history and genetic diversity within the genus. Monophyletic groupings depicting strong bootstrap support were shown to have a single common ancestor and genetic coherence with limited genetic divergence among sequences. Polyphyletic groupings also presented significant genetic differentiation within the F. verticillioides sequences from diverse ecological niches. Nucleotide diversity of moderate level 0.02471 reflected genetic variations within populations that were attributed to factors such as mutation, genetic drift, or varying selection pressures. The Fst value of 0.98205 is particularly indicative of high genetic differentiation, implying that most of the genetic variance is due to differences between populations rather than within them. F. verticillioides, with 57 sequences, showed low genetic diversity with three segregating sites and a low haplotype diversity of 0.19486, suggesting the founder effect, where a reduced population expands from a limited genetic pool. The total data estimates across all populations for haplotype analysis showed 72 sequences, 44 segregating sites, and 9 haplotypes with a haplotype diversity of 0.48513. The evolutionary dynamics showed genetic differentiations among Fusarium species causing FSR. AMOVA indicated high within-population variations, depicting a substantial genetic diversity within individual populations. The results offer a comprehensive framework for discussing the implications of genetic diversity in pathogen management and the evolutionary dynamics of the Fusarium species causing FSR in maize in the Indian subcontinent.

Varying Selection Pressure for a Na+ Sensing Site in Epithelial Na+ Channel Subunits Reflect Divergent Roles in Na+ Homeostasis.

The epithelial Na+ channel (ENaC) emerged early in vertebrates and has played a role in Na+ and fluid homeostasis throughout vertebrate evolution. We previously showed that proteolytic activation of the channel evolved at the water-to-land transition of vertebrates. Sensitivity to extracellular Na+, known as Na+ self-inhibition, reduces ENaC function when Na+ concentrations are high and is a distinctive feature of the channel. A fourth ENaC subunit, δ, emerged in jawed fishes from an α subunit gene duplication. Here, we analyzed 849 α and δ subunit sequences and found that a key Asp in a postulated Na+ binding site was nearly always present in the α subunit, but frequently lost in the δ subunit (e.g. human). Analysis of site evolution and codon substitution rates provide evidence that the ancestral α subunit had the site and that purifying selection for the site relaxed in the δ subunit after its divergence from the α subunit, coinciding with a loss of δ subunit expression in renal tissues. We also show that the proposed Na+ binding site in the α subunit is a bona fide site by conferring novel function to channels comprising human δ subunits. Together, our findings provide evidence that ENaC Na+ self-inhibition improves fitness through its role in Na+ homeostasis in vertebrates.

The evolutionary cost of homophily: social stratification facilitates stable variant coexistence and increased rates of evolution in host-associated pathogens.

Coexistence of multiple strains of a pathogen in a host population can present significant challenges to vaccine development or treatment efficacy. Here we discuss a novel mechanism that can increase rates of long-lived strain polymorphism, rooted in the presence of social structure in a host population. We show that social preference of interaction, in conjunction with differences in immunity between host subgroups, can exert varying selection pressure on pathogen strains, creating a balancing mechanism that supports stable viral coexistence, independent of other known mechanisms. We use population genetic models to study rates of pathogen heterozygosity as a function of population size, host population composition, mutant strain fitness differences and host social preferences of interaction. We also show that even small periodic epochs of host population stratification can lead to elevated strain coexistence. These results are robust to varying social preferences of interaction, overall differences in strain fitnesses, and spatial heterogeneity in host population composition. Our results highlight the role of host population social stratification in increasing rates of pathogen strain diversity, with effects that should be considered when designing policies or treatments with a long-term view of curbing pathogen evolution.

Seeing across variable ecological and social environments: comparative eye morphology of marine and terrestrial hermit crabs (Decapoda: Anomura: Coenobitidae, Paguridae)

Abstract Vision is a ubiquitous sensory modality adapted to vastly different environments, which place variable selection pressures on both macro- and microscopic dimensions of external eye morphology. The hermit crab clade (Crustacea: Decapoda: Anomura) spans environments from sea to land, yet, no systematic comparisons in eye morphology have been made between any species. We compared three species of hermit crabs inhabiting different ecological and social environments: two terrestrial species, one of which (Coenobita compressusH. Milne Edwards, 1837) is highly social and inhabits a flat, open diurnal habitat, and another (C. clypeatus Herbst, 1791) is its sister species that inhabits a rugged forested nocturnal habitat; and a third (Pagurus longicarpusSay, 1817) inhabits a marine intertidal habitat and is less social. We compared macroscopic eye morphology between species as well as microscopic differences in ommatidium-facet density and diameter within eyes. We found that eye dimensions (length, width, and thickness), but not overall volume, differed significantly between marine and terrestrial species. Furthermore, the highly social C. compressus had a significantly larger eye volume compared to its sister species, C. clypeatus, which is not as social. Larger eyes in C. compressus might be linked to its social lifestyle, where vision has been shown to be critical to finding conspecifics. All three species had a higher density of facets in the ventral compared to dorsal region of the eye. Our comparisons of external eye morphology revealed microscopic similarities but notable macroscopic variation between species, thus laying a foundation for future phylogenetically controlled tests within this species-rich clade, including comparisons of internal eye morphology.

Accumulation of resistance genes in Salmonella Typhimurium transmitted between poultry and dairy farms increases the risk to public health.

Salmonella Typhimurium is a zoonotic pathogen that poses a major threat to public health. This generalist serotype can be found in many hosts and the environment where varying selection pressures may result in the accumulation of antimicrobial resistance determinants. However, the transmission of this serotype between food-producing hosts, specifically between poultry layer flocks and nearby dairy herds, was never demonstrated. We investigated an outbreak at a dairy in Israel to determine the role of nearby poultry houses to be sources of infection. The 2-month outbreak resulted in a 47% mortality rate among 15 calves born in that period. Routine treatment of fluid therapy, a nonsteroidal anti-inflammatory, and cefquinome was ineffective, and control was achieved by the introduction of vaccination of dry cows against Salmonella (Bovivac S, MSD Animal Health) and a strict colostrum regime. Whole genome sequencing and antimicrobial sensitivity tests were performed on S. Typhimurium strains isolated from the dairy (n = 4) and strains recovered from poultry layer farms (n = 10). We identified acquired antimicrobial-resistant genes, including the blaCTX-M-55 gene, conferring resistance to extended-spectrum cephalosporins, which was exclusive to dairy isolates. Genetic similarity with less than five single nucleotide polymorphism differences between dairy and poultry strains suggested a transmission link. This investigation highlights the severe impact of S. Typhimurium on dairy farms and the transmission risk from nearby poultry farms. The accumulation of potentially transferable genes conferring resistance to critically important antimicrobials underscores the increased public health risk associated with S. Typhimurium circulation between animal hosts.IMPORTANCESalmonella Typhimurium is one of the major causes of food-borne illness globally. Infections may result in severe invasive disease, in which antimicrobial treatment is warranted. Therefore, the emergence of multi-drug-resistant strains poses a significant challenge to successful treatment and is considered one of the major threats to global health. S. Typhimurium can be found in a variety of animal hosts and environments; however, its transmission between food-producing animals, specifically poultry layers flocks and dairy herds, was never studied. Here, we demonstrate the transmission of the pathogen from poultry to a nearby dairy farm. Alarmingly, the multi-drug-resistant strains collected during the outbreak in the dairy had acquired resistance to extended-spectrum cephalosporins, antibiotics critically important in treating Salmonellosis in humans. The findings of the study emphasize the increased risk to public health posed by zoonotic pathogens' circulation between animal hosts.

Expression of defensin genes across house fly (Musca domestica) life history gives insight into immune system subfunctionalization.

Animals encounter diverse microbial communities throughout their lifetime, which exert varying selection pressures. Antimicrobial peptides (AMPs), which lyse or inhibit microbial growth, are a first line of defense against some of these microbes. Here we examine how developmental variation in microbial exposure has affected the evolution of expression and amino acid sequences of Defensins (an ancient class of AMPs) in the house fly (Musca domestica). The house fly is a well-suited model for this work because it trophically associates with varying microbial communities throughout its life history and its genome contains expanded families of AMPs, including Defensins. We identified two subsets of house fly Defensins: one expressed in larvae or pupae, and the other expressed in adults. The amino acid sequences of these two Defensin subsets form distinct monophyletic clades, and they are located in separate gene clusters in the genome. The adult-expressed Defensins evolve faster than larval/pupal Defensins, consistent with different selection pressures across developmental stages. Our results therefore suggest that varied microbial communities encountered across life history can shape the evolutionary trajectories of immune genes.

Are dark-eyed dogs favoured by humans? Domestication as a potential driver of iris colour difference between dogs and wolves.

Comparative studies have shown that the eye morphology of primates has been shaped by a variety of selection pressures (e.g. communication, environmental factors). To comprehensively elucidate the complex links between ocular morphology and its evolutionary drive, attention should be paid to other phylogenetic groups. Here, we address a new question regarding the evolution of eye colour patterns in the oldest domesticated animal, namely, the domestic dog (Canis familiaris). In this study, we conducted an image analysis of dogs and their closest relatives, grey wolves (Canis lupus), to compare the colours of their irises, with the aim of assessing whether eye colours of dogs affect how humans perceived dogs. We found that the irises of dogs were significantly darker than those of wolves. We also found that facial images of dark-eyed dogs were perceived as more friendly and immature, potentially eliciting caregiving responses from humans. Our findings are consistent with our expectation that humans favour dark-eyed dogs over light-eyed ones and provide an updated hypothesis that dogs with dark eyes may have evolved by acquiring a facial trait that sends a non-threatening gaze signal to humans.

Microdiverse bacterial clades prevail across Antarctic wetlands.

Antarctica's extreme environmental conditions impose selection pressures on microbial communities. Indeed, a previous study revealed that bacterial assemblages at the Cierva Point Wetland Complex (CPWC) are shaped by strong homogeneous selection. Yet which bacterial phylogenetic clades are shaped by selection processes and their ecological strategies to thrive in such extreme conditions remain unknown. Here, we applied the phyloscore and feature-level βNTI indexes coupled with phylofactorization to successfully detect bacterial monophyletic clades subjected to homogeneous (HoS) and heterogenous (HeS) selection. Remarkably, only the HoS clades showed high relative abundance across all samples and signs of putative microdiversity. The majority of the amplicon sequence variants (ASVs) within each HoS clade clustered into a unique 97% sequence similarity operational taxonomic unit (OTU) and inhabited a specific environment (lotic, lentic or terrestrial). Our findings suggest the existence of microdiversification leading to sub-taxa niche differentiation, with putative distinct ecotypes (consisting of groups of ASVs) adapted to a specific environment. We hypothesize that HoS clades thriving in the CPWC have phylogenetically conserved traits that accelerate their rate of evolution, enabling them to adapt to strong spatio-temporally variable selection pressures. Variable selection appears to operate within clades to cause very rapid microdiversification without losing key traits that lead to high abundance. Variable and homogeneous selection, therefore, operate simultaneously but on different aspects of organismal ecology. The result is an overall signal of homogeneous selection due to rapid within-clade microdiversification caused by variable selection. It is unknown whether other systems experience this dynamic, and we encourage future work evaluating the transferability of our results.

Ancistroceruswasps (Hymenoptera, Vespidae) from the Centre of Europe: Phylogeny, cryptic species, neutral and non‐neutral markers

AbstractThe aim of the study was to clarify the phylogenetic relationships among Northern EuropeanAncistrocerusand comparison of the applicability of evolutionarily neutral and non‐neutral markers for reconstruction of phylogeny. We used a 19,400 bp long dataset that included parts of mitochondrial DNA, nuclear rDNA operon, and 10 nuclear protein‐coding genes. Application of molecular barcoding and species delimitation algorithms unveiled a presence of cryptic species,A. balticussp. n., in the trap‐nesting wasp communities of the centre of Europe. We assessed the morphological, biological, and ecological differences of it from the siblingA. trifasciatusand updated the regional identification key. Phylogeny reconstruction using the neutral and the presumably non‐neutral markers resulted in different tree topologies. Evolutionary congruence of the rDNA operon with the other markers was relatively low. Evolutionary rate of the mitochondrial genes was 7–8 times as high as that of the exons of the nuclear genes, therefore, the mitochondrial markers overshadowed the nuclear ones in the phylogeny reconstructions. We assumed that at the speciation level, we might consider two different patterns of phylogeny: one based on evolutionary time and neutral changes, and the other based on adaptive evolutionary pathways under directional selection pressures. We assessed the effect of directional selection on the nuclear protein‐coding genes, applying the Spearman's rank correlation between pairwise phylogenetic distances among species, estimated using exons, and these distances, estimated using introns. One of these markers demonstrated a lack of positive correlation, implying a variable directional selection pressure on the coded protein. The publication has been registered on ZooBank: urn:lsid:zoobank.org:pub:13BD28D0‐736D‐4B2A‐B5CF‐4824BD4CDCFB.

Abstract 6621: Simulating the distortion of clonal fractions in ctDNA due to spatially heterogenous selection

Abstract Time-series circulating tumor DNA (ctDNA) sequencing has the potential to reveal emerging variants in the tissue in real time, and there is now substantial evidence that it can detect tumor growth and treatment-resistant mutations long before growth is visible. However, the reliability of ctDNA for profiling the primary tumor may be compromised by spatial heterogeneity in the tumor genetics, for example due to selective effects of differential drug penetration, immune infiltration, and oxygenation. We explore the influence of spatial factors on tumor genetics through a lattice-based branching process model of solid tumor evolution and ctDNA shedding with the goal of understanding where spatially variable selection pressure could lead to a significant difference between the clonal fractions in the blood and the tissue. The lattice model divides space into regions with different growth rates which result in different rates of driver mutation accumulation. The effects of regional differences in selection cause different apoptosis rates to correlate with the location of expanding driver clones. Using these rates to simulate ctDNA shedding, our model found that clones fit enough to invade a hostile region are temporarily amplified in the blood compared to their tissue frequency by as much as 300% for a limited window of time before the population expands further and the clonal fraction in the blood returns to close agreement with tissue. The results suggest evolutionary contexts in which liquid biopsy might provide highly biased estimates of tumor state, but also shows how this effect might be exploited to provide more sensitive detection of clonal expansion with implications for blood-based detection of recurrence and metastasis. Citation Format: Thomas W. Rachman, Oana Carja, Russell Schwartz. Simulating the distortion of clonal fractions in ctDNA due to spatially heterogenous selection [abstract]. In: Proceedings of the American Association for Cancer Research Annual Meeting 2023; Part 1 (Regular and Invited Abstracts); 2023 Apr 14-19; Orlando, FL. Philadelphia (PA): AACR; Cancer Res 2023;83(7_Suppl):Abstract nr 6621.

Evolutionary footprints of cold adaptation in arctic-alpine Cochlearia (Brassicaceae) – Evidence from freezing experiments and electrolyte leakage

As global warming progresses, plants may be forced to adapt to drastically changing environmental conditions. Arctic-alpine plants have been among the first to experience the effects of climate change. As a result, cold acclimation and freezing tolerance may become increasingly crucial for the survival as winter warming events and earlier snowmelt will cause increased exposure to occasional frost. The tribe Cochlearieae in the mustard family (Brassicaceae) offers an instructive system for studying cold adaptation in evolutionary terms, as the two sister genera Ionopsidium and Cochlearia are distributed among different ecological habitats throughout the European continent and the far north into circumarctic regions. By applying an electrolyte leakage assay to leaves obtained from plants cultivated under controlled temperature regimes in growth chambers, the freezing tolerance of different Ionopsidium and Cochlearia species was assessed measuring lethal freezing temperature values (LT50 and LT100), thereby allowing for a comparison across different species and accessions in their responses to cold. We hypothesized that, owing to varying selection pressures, geographically distant species would differ in freezing tolerance. Despite Ionopsidium occurring under warm and dry Mediterranean conditions and Cochlearia species distributed often at cold habitats, all accessions exhibited similar cold responses. The results may indicate that physiological adaptations of primary metabolic pathways to different stressors, such as salinity and drought, may confer an additional tolerance to cold; this is because all these stressors induce osmotic challenges.

Genomic Comparisons Reveal Selection Pressure and Functional Variation Between Nutritional Endosymbionts of Cave-Adapted and Epigean Hawaiian Planthoppers.

Planthoppers in the family Cixiidae (Hemiptera: Auchenorrhyncha: Fulgoromorpha) harbor a diverse set of obligate bacterial endosymbionts that provision essential amino acids and vitamins that are missing from their plant-sap diet. "Candidatus Sulcia muelleri" and "Ca. Vidania fulgoroidea" have been associated with cixiid planthoppers since their origin within the Auchenorrhyncha, whereas "Ca. Purcelliella pentastirinorum" is a more recent endosymbiotic acquisition. Hawaiian cixiid planthoppers occupy diverse habitats including lava tube caves and shrubby surface landscapes, which offer different nutritional resources and environmental constraints. Genomic studies have focused on understanding the nutritional provisioning roles of cixiid endosymbionts more broadly, yet it is still unclear how selection pressures on endosymbiont genes might differ between cixiid host species inhabiting such diverse landscapes, or how variation in selection might impact symbiont evolution. In this study, we sequenced the genomes of Sulcia, Vidania, and Purcelliella isolated from both surface and cave-adapted planthopper hosts from the genus Oliarus. We found that nutritional biosynthesis genes were conserved in Sulcia and Vidania genomes in inter- and intra-host species comparisons. In contrast, Purcelliella genomes retain different essential nutritional biosynthesis genes between surface- and cave-adapted planthopper species. Finally, we see the variation in selection pressures on symbiont genes both within and between host species, suggesting that strong coevolution between host and endosymbiont is associated with different patterns of molecular evolution on a fine scale that may be associated with the host diet.

Genetic Covariation Between the Vertically Transmitted Endophyte Epichloë canadensis and Its Host Canada Wildrye.

Symbiotic mutualisms are thought to be stabilized by correlations between the interacting genotypes which may be strengthened via vertical transmission and/or reduced genetic variability within each species. Vertical transmission, however, may weaken interactions over time as the endosymbionts would acquire mutations that could not be purged. Additionally, temporal variation in a conditional mutualism could create genetic variation and increased variation in the interaction outcome. In this study, we assessed genetic variation in both members of a symbiosis, the endosymbiotic fungal endophyte Epichloë canadensis and its grass host Canada wildrye (Elymus canadensis). Both species exhibited comparable levels of diversity, mostly within populations rather than between. There were significant differences between populations, although not in the same pattern for the two species, and the differences were not correlated with geographic distance for either species. Interindividual genetic distance matrices for the two species were significantly correlated, although all combinations of discriminant analysis of principle components (DAPC) defined multilocus genotype groups were found suggesting that strict genotype matching is not necessary. Variation in interaction outcome is common in grass/endophyte interactions, and our results suggest that the accumulation of mutations overtime combined with temporal variation in selection pressures increasing genetic variation in the symbiosis may be the cause.

Balancing selection on genomic deletion polymorphisms in humans.

A key question in biology is why genomic variation persists in a population for extended periods. Recent studies have identified examples of genomic deletions that have remained polymorphic in the human lineage for hundreds of millennia, ostensibly owing to balancing selection. Nevertheless, genome-wide investigation of ancient and possibly adaptive deletions remains an imperative exercise. Here, we demonstrate an excess of polymorphisms in present-day humans that predate the modern human-Neanderthal split (ancient polymorphisms), which cannot be explained solely by selectively neutral scenarios. We analyze the adaptive mechanisms that underlie this excess in deletion polymorphisms. Using a previously published measure of balancing selection, we show that this excess of ancient deletions is largely owing to balancing selection. Based on the absence of signatures of overdominance, we conclude that it is a rare mode of balancing selection among ancient deletions. Instead, more complex scenarios involving spatially and temporally variable selective pressures are likely more common mechanisms. Our results suggest that balancing selection resulted in ancient deletions harboring disproportionately more exonic variants with GWAS (genome-wide association studies) associations. We further found that ancient deletions are significantly enriched for traits related to metabolism and immunity. As a by-product of our analysis, we show that deletions are, on average, more deleterious than single nucleotide variants. We can now argue that not only is a vast majority of common variants shared among human populations, but a considerable portion of biologically relevant variants has been segregating among our ancestors for hundreds of thousands, if not millions, of years.

Intraspecific variation in morphology of spiny pollen grains along an altitudinal gradient in an insect-pollinated shrub.

Intraspecific variations in pollen morphological traits are poorly studied. Interspecific variations are often associated with pollination systems and pollinator types. Altitudinal environmental changes, which can influence local pollinator assemblages, provide opportunities to explore differentiation in pollen traits of a single species over short distances. The aim of this study is to examine intraspecific variations in pollen traits ofan insect-pollinated shrub, Weigela hortensis (Caprifoliaceae), along an altitudinal gradient. Pollen spine phenotypes (length, number and density), pollen diameter, lipid mass (pollenkitt) around pollen grains, pollen production per flower and pollinator assemblages were compared at four sites at different altitudes. Spine length and the spine length/diameter ratio of pollen grains were greater at higher altitudes but not correlated with flower or plant size. Spine number and density increased as flower size increased, and pollen lipid mass decreased as plant size increased. Bees were the predominant pollinators at low-altitude sites whereas flies, specifically Oligoneura spp. (Acroceridae), increased in relative abundance with increasing altitude. The results of this study suggest that the increase in spine length with altitude was the result of selection favouring longer spines at higher-altitude sites and/or shorter spines at lower-altitude sites. The altitudinal variation in selection pressure on spine length could reflect changes in local pollinator assemblages with altitude.

Correlated expression of phenotypic and extended phenotypic traits across stingless bee species: worker eye morphology, foraging behaviour, and nest entrance architecture

Stingless bees are the most species-rich group of eusocial bees and show great diversity in behaviour, ecology, nest architecture, colony size, and worker morphology. How this variation relates to varying selection pressures and constraints is not well understood. Variation can be caused by selection acting on behavioural or morphological traits, both alone and in correlation across traits. Here we tested whether behavioural and morphological traits important for foraging and defence are linked to nest-entrance architecture, an extended phenotype relevant to both foraging and nest defence. Using 23 species we investigated whether eye size, nest entrance size, landing behaviour and foraging method show cross-species correlations. A phylogenetically-controlled comparative analysis revealed that species with relatively smaller eyes build relatively larger entrances, which in turn are associated with faster landing approaches and fewer landing errors by foragers, both of which could reduce predation risk. Concerning foraging, mass-recruiting species have c. 10-times larger entrance holes than species with a solitary foraging strategy. Larger entrances could help species with mass recruitment to rapidly increase forager traffic or mount a strong defensive response when under attack. Our results show that studying correlations among different traits helps understand phenotypic diversity in species rich groups.