Evolutionary Timescales Research Articles

Critical Role of Pressure for Chemo-Mechanical-Induced Stability of Sodium Metal Battery Anodes

In this work we demonstrate that cell pressure controls the morphology and stability of electroplated sodium metal deposits on carbon black nucleation layers in ether-based electrolytes. At pressures below 500 kPa we observe the presence of three-dimensional Na nuclei accompanied by low Coulombic efficiencies (CEs less than 98%). Conversely, at pressures between 500 and 1272 kPa we observe smooth, planar Na deposits, high CEs up to 99.9%, and stable electrochemical cycling. Through a series of tests conducted at elevated current densities and with or without rest stages, our findings elucidate the balance of important competing time scales for creep and morphology evolution under pressure and the rate of charge transfer that determines Na morphology and stability. This highlights how chemo-mechanical effects at pressure ranges relevant for battery packaging in coin and pouch cells are key factors in the design and operation of Na metal batteries.

How to be a fig nematode

Fig (Moraceae: Ficus) species host vast communities of organisms that are bound together by complicated ecological networks that have influenced community structure and dynamics over evolutionary timescales. Much attention has been paid to the mutualism between figs and their specialized pollinating fig wasps, as well as with often antagonistic non-pollinating fig wasps. Equally ubiquitous to fig systems, but much less understood are the multitude of nematode groups that have independently evolved obligate associations with pollinating fig wasps and proliferate inside fig syconia. In this review we describe what is currently known (and unknown) about these numerous and increasingly studied nematode taxa and how they interact with the fig systems they inhabit. We identify the groups that are currently understood to associate with fig pollinators and outline their known species distributions and evolutionary history, where possible. Special attention is paid to the life history of these nematode groups, especially which features of nematode biology are generalizable across groups and what idiosyncratic peculiarities exist within individual genera. We outline key biological features including host choice, dispersal, disembarkation, diet, mating, and proliferation within figs. We address biological conundrums that have been raised following observational work such as, why do nematodes sometimes infect the wrong host? What adaptations were required for nematodes to successfully adapt and coexist with figs and their pollinators for millions of years? How do nematodes overcome the constraints of low mating group size? Finally, we outline key considerations, gaps in the knowledge, and limitations to expand this field forward towards promising areas of future research. Through better understanding of fig nematodes, we stand to not only know more about Ficus communities, but also more about the evolution and maintenance of interspecific interactions, development, adaptation, and co-evolution in general.

Single-cell phenotypic plasticity modulates social behavior in Dictyostelium discoideum

In Dictyostelium chimeras, "cheaters" are strains that positively bias their contribution to the pool of spores, i.e., the reproductive cells resulting from development. On evolutionary time scales, the selective advantage; thus, gained by cheaters is predicted to undermine collective functions whenever social behaviors are genetically determined. Genotypes; however, are not the sole determinant of spore bias, but the relative role of genetic and plastic differences in evolutionary success is unclear. Here, we study chimeras composed of cells harvested in different phases of population growth. We show that such heterogeneity induces frequency-dependent, plastic variation in spore bias. In genetic chimeras, the magnitude of such variation is not negligible and can even reverse the classification of a strain's social behavior. Our results suggest that differential cell mechanicalproperties can underpin, through biases emerging during aggregation, a "lottery" in strains' reproductive success that may counter the evolution of cheating.

Evolution of a new form of haploid-specific gene regulation appearing in a limited clade of ascomycete yeast species.

Over evolutionary timescales, the logic and pattern of cell-type specific gene expression can remain constant, yet the molecular mechanisms underlying such regulation can drift between alternative forms. Here, we document a new example of this principle in the regulation of the haploid-specific genes in a small clade of fungal species. For most ascomycete fungal species, transcription of these genes is repressed in the a/α cell type by a heterodimer of two homeodomain proteins, Mata1 and Matα2. We show that in the species Lachancea kluyveri, most of the haploid-specific genes are regulated in this way, but repression of one haploid-specific gene (GPA1) requires, in addition to Mata1 and Matα2, a third regulatory protein, Mcm1. Model building, based on x-ray crystal structures of the three proteins, rationalizes the requirement for all three proteins: no single pair of the proteins is optimally arranged, and we show that no single pair can bring about repression. This case study exemplifies the idea that the energy of DNA binding can be "shared out" in different ways and can result in different DNA-binding solutions across different genes-while maintaining the same overall pattern of gene expression.

A study of carbon-rich post-AGB stars in the Milky Way to understand the production of carbonaceous dust from evolved stars

Context. Knowledge of the Gaia, DR3 parallaxes of Galactic post-asymptotic giant branch (AGB) stars makes it possible to exploit these objects as tracers of AGB evolution, nucleosynthesis, and dust production as well as to use them to shed new light on still poorly known physical processes experienced by AGB stars. Aims. The goal of this study is to reconstruct the evolution and the dust formation processes during the final AGB phases of a sample of carbon-rich, post-AGB Galactic stars, with particular attention to the determination of the past mass-loss history. Methods. We study the IR excess of Galactic sources classified as post-AGB single stars by means of dust formation modelling where dust grains form and grow in a static wind and expand from the surface of the star. The method is applied to various evolutionary stages of the final AGB phase of stars with different masses and metallicities. The results from a spectral energy distribution (SED) fitting are used to infer information on mass loss, efficiency of dust formation, and wind dynamics. Results. The detailed analysis of the SED of the sources investigated, which included the derivation of the luminosities and the dust properties, allows us to confirm previous results, mostly based on the surface chemical composition, that most of the investigated sources descend from low-mass (M < 1.5 M⊙) progenitors that reached the C-star stage. Metal-poor carbon stars are characterised by higher IR excesses with respect to their more metal-rich counterparts of similar luminosity due to a higher surface carbon-to-oxygen excess. This work confirms previous conclusions based on a limited sample of carbon-rich post-AGB objects in the Magellanic Clouds, namely that more luminous stars descending from higher-mass progenitors are generally more opaque due to shorter evolutionary timescales that place the dust shell closer to the central object. Through the study of the dynamics of the outflow and results from stellar evolution modelling, we find that the mass-loss rate at the tip of the AGB phase of metal-rich low-mass carbon stars is approximately 1#x2212;1.5 × 10−5 M⊙ yr−1, whereas in the metal-poor domain Ṁ ∼ 4 − 5 × 10−5 M⊙ yr−1 is required. These results indicate the need for an upwards revision of the theoretical mass-loss rates of low-mass carbon stars in the available literature, which in turn require a revised determination of carbon dust yields by AGB stars.

Phenotypic selection in natural populations: what have we learned in 40 years?

In 1983, Russell Lande and Stevan Arnold published "The measurement of selection on correlated characters," which became a highly influential citation classic in evolutionary biology. This paper stimulated a cottage industry of field studies of natural and sexual selection in nature and resulted in several large-scale meta-analyses, statistical developments, and method papers. The statistical tools they suggested contributed to a breakdown of the traditional dichotomy between ecological and evolutionary time scales and stimulated later developments such as "eco-evolutionary dynamics". However, regression-based selection analyses also became criticized from philosophical, methodological, and statistical viewpoints and stimulated some still ongoing debates about causality in evolutionary biology. Here I return to this landmark paper by Lande and Arnold, analyze the controversies and debates it gave rise to and discuss the past, present, and future of selection analyses in natural populations. A remaining legacy of Lande & Arnold, 1983 is that studies of selection and inheritance can fruitfully be decoupled and be studied separately, since selection acts on phenotypes regardless of their genetic basis, and hence selection and evolutionary responses to selection are distinct processes.

Dynamics and stage-specificity of between-population gene expression divergence in the Drosophila melanogaster larval fat body

Gene expression variation is pervasive across all levels of organismal organization, including development. Few studies, however, have examined variation in developmental transcriptional dynamics among populations, or how it contributes to phenotypic divergence. Indeed, the evolution of gene expression dynamics when both the evolutionary and temporal timescale are comparatively short remains relatively uncharacterized. Here, we examined coding and non-coding gene expression in the fat body of an ancestral African and a derived European Drosophila melanogaster population across three developmental stages spanning ten hours of larval development. Between populations, expression divergence was largely stage-specific. We detected higher expression variation during the late wandering stage, which may be a general feature of this stage. During this stage, we also detected higher and more extensive lncRNA expression in Europe, suggesting that lncRNA expression may be more important in derived populations. Interestingly, the temporal breadth of protein-coding and lncRNA expression became more restricted in the derived population. Taken together with the signatures of potential local adaptation that we detected at the sequence level in 9-25% of candidate genes (those showing evidence of expression divergence between populations), this finding suggests that gene expression becomes more developmental stage-specific during adaptation to new environments. We further used RNAi to identify several candidate genes that likely contribute to known phenotypic divergence between these populations. Our results shed light on the evolution and dynamics of expression variation over short developmental and evolutionary timescales, and how this variation contributes to population and phenotypic divergence.

What’s in an age? Calculation and interpretation of ages and durations from U-Pb zircon geochronology of igneous rocks

Accurate assessment of the duration of zircon crystallization within igneous rocks is critical for constraining the time scales of magmatic evolution and storage, which have important implications for our understanding of magmatic fluxes and volcanic hazards. However, estimation of crystallization durations from finite geochronologic data sets is difficult and typically relies on numerous implicit assumptions. In this contribution, we evaluate these assumptions and provide recommendations for better interpretation of crystallization durations from individual samples by developing a simplified theoretical framework to relate zircon growth, nucleation, and armoring rates to zircon ages. We first investigate single zircon analyses and show that ages produced with methods that integrate the entire grain or grain fragments (e.g., chemical abrasion−isotope dilution−thermal ionization mass spectrometry [CA-ID-TIMS]) are inevitably biased toward the second half of the zircon growth interval, while subsampling of grains via microbeam approaches will only capture the majority of the zircon crystallization duration when the microbeam spot size is less than ∼25% of the zircon minor axis, and the analytical uncertainty of the measurement is less than ∼20% of the duration over which the individual zircon grew. We subsequently investigate the distribution of zircon mean ages produced through various combinations of zircon growth rate, nucleation rate, and the probability of zircon being armored by major phases. We show that zircon age distributions cannot be directly predicted from the rate of zircon mass crystallized, as many combinations of growth, nucleation, and armoring rates result in distinct age distributions, yet they produce nearly identical mass crystallization rates. Finally, we develop two equations that can be used to constrain the duration of crystallization observed within individual samples. In scenarios where the observed age dispersion is consistent with the reported analytical uncertainties, the first equation can be used to estimate the maximum duration. Otherwise, when the measured zircon population is overdispersed, a second equation constrains the minimum duration of zircon crystallization.

Global instability by runaway collisions in nuclear stellar clusters: numerical tests of a route for massive black hole formation

ABSTRACT The centres of galaxies host nuclear stellar clusters, supermassive black holes, or both. The origin of this dichotomy is still a mystery. Nuclear stellar clusters are the densest stellar system in the Universe, so they are ideal places for runaway collisions to occur. Previous studies have proposed the possible existence of a critical mass scale in such clusters, for which the occurrence of collisions becomes very frequent and leads to the formation of a very massive object. While it is difficult to directly probe this scenario with simulations, we here aim for a proof of concept using toy models where the occurrence of such a transition is shown based on simplified compact systems, where the typical evolution time-scales will be faster compared to the real Universe. Indeed our simulations confirm that such a transition takes place and that up to 50 per cent of the cluster mass can go into the formation of a central massive object for clusters that are above the critical mass scale. Our results thus support the proposed new scenario on the basis of idealized simulations. A preliminary analysis of observed nuclear star clusters shows similar trends related to the critical mass as in our simulations. We further discuss the caveats for the application of the proposed scenario in real nuclear star clusters.

Effect of Host-Switching on the Ecological and Evolutionary Patterns of Parasites.

Speciation via host-switching is a macroevolutionary process that emerges from a microevolutionary dynamic where individual parasites switch hosts, establish a new association, and reduce reproductive contact with the original parasite lineage. Phylogenetic distance and geographic distribution of the hosts have been shown to be determinants of the capacity and opportunity of the parasite to change hosts. Although speciation via host-switching has been reported in many host-parasite systems, its dynamic on the individual, population and community levels is poorly understood. Here we propose a theoretical model to simulate parasite evolution considering host-switching events on the microevolutionary scale, taking into account the macroevolutionary history of the hosts, to evaluate how host-switching can affect ecological and evolutionary patterns of parasites in empirical communities at regional and local scales. In the model, parasite individuals can switch hosts under variable intensity and have their evolution driven by mutation and genetic drift. Mating is sexual and only individuals that are sufficiently similar can produce offspring. We assumed that parasite evolution occurs at the same evolutionary time scale as their hosts, and that the intensity of host-switching decreases as the host species differentiate. Ecological and evolutionary patterns were characterized by the turnover of parasite species among host species, and parasite evolutionary tree imbalance respectively. We found a range of host-switching intensity that reproduces ecological and evolutionary patterns observed in empirical communities. Our results showed that turnover decreased as host-switching intensity increased, with low variation among the model replications. On the other hand, tree imbalance showed wide variation and non-monotonic tendency. We concluded that tree imbalance was sensitive to stochastic events, whereas turnover may be a good indicator of host-switching. We found that local communities corresponded to higher host-switching intensity when compared to regional communities, highlighting that spatial scale is a limitation for host-switching. [Dispersal of parasites, opportunity and capacity of interaction, phylogenetic conservatism, and community structure.].

Exceptional endemicity of Aotearoa New Zealand biota shows how taxa dispersal traits, but not phylogeny, correlate with global species richness

ABSTRACT Species’ with more limited dispersal and consequently less gene flow are more likely to form new spatially segregated species and thus contribute disproportionally to endemic biota and global species richness. Aotearoa New Zealand has exceptional endemicity, with 52% of its 54,000 named species endemic, including 32%, 39% and 68% for freshwater, marine and terrestrial environments respectively. The lower endemicity of freshwater biota (excluding insects) is attributed to their need to disperse between habitats that are temporary on evolutionary timescales. The percent endemicity of higher taxa (Order to Kingdom), a measure of phylogenetic relationships, was not correlated with regional and global species richness. However, there was a positive correlation between endemicity and species richness across dispersal trait groups based on their environment, typical body size, mobility (including flight), and if marine, whether pelagic or benthic. Typically flighted taxa had high endemicity contrary to the dispersal-endemicity hypothesis, but reflecting exceptional isolation by distance and time, and reduced flight ability as occurs on islands. It is proposed that the high richness and endemicity of mobile macrofauna is caused by a combination of niche specialisation opportunities and predation limiting dispersal respectively. Thus, dispersal traits better predicted endemicity and global species richness than phylogeny.

Back from Exile? First Records of Chewing Lice (Lutridia exilis; Ischnocera; Mallophaga) in Growing Eurasian Otter (Lutra lutra) Populations from Northern Germany.

Arthropod ectoparasites of aquatic wildlife often have complex relationships with their host species that have developed over long evolutionary time scales. Specialist parasite occurrence might depend on these hosts' distributions. Eurasian otter (Lutra lutra) populations are recovering in Northern German federal states, such as Schleswig-Holstein and Lower Saxony. Chewing lice (Lutridia exilis; Ischnocera; Mallophaga) are considered otter-specific yet rare parasites in their known range. In 2022, they were recorded for the first time on nine otters found dead in Northern Germany. All otters originated from the years 2021-2022 and were dissected during population health monitoring programs in 2022. Females (n = 6) were 0-5.5 years old and showed signs of disease in five cases. Males (n = 3), in contrast, were 0-1.6 years old and showed disease in a single case. Individual lice intensity of infection ranged from 1 to 75 specimens per otter. No direct adverse health effects of chewing lice on the otters were noted. Lutridia exilis morphological characteristics were documented and measurements were taken to study specialized adaptations that allow lice to attach to semi-aquatic otters. In addition, morphology was compared between lice from different geographical regions and specimens from previous reports. A region of the COI mDNA was amplified to molecularly characterize L. exilis for the first time and detect genetic differences between otter lice populations in Germany. It is believed that specialist parasites reduce in numbers even before their host populations decline. Recovering otter populations in Northern Germany could be an example of a reverse effect, where the comeback of a host species results in the return of a specialist parasite, which reflects an ultimate boost in overall species biodiversity.

Target-capture probes for phylogenomics of the Caenogastropoda.

Target-capture approaches have facilitated a rapid growth in the field of phylogenomics but few probe sets exist for mollusks, an exceptionally rich phylum with unparalleled ecological and morphological diversity. We designed and tested the first universal probe set using Phyluce to capture ultraconserved elements (UCEs) and exon loci from the Subclass Caenogastropoda - one of six major lineages of gastropods. The probe set consists of 29,441 probes designed to target 1,142 UCE loci and 1,933 exon loci (3,075 total). In silico analyses of our probe set yielded an average of 2,110 loci from genomes and 1,389 loci from transcriptomes of diverse caenogastropods, from which an average of 1,669 and 849 loci were retained respectively after screening to remove those that matched multiple contigs. Phylogenetic analyses of the loci extracted from transcriptomes produced well-supported trees very similar to those published based on transcriptomic analyses. Phylogenetic relationships estimated from loci extracted from genomes recover similar phylogenetic relationships, and indicate that the loci targeted with this probe set are informative for resolving deep phylogenetic relationships. An in vitro analysis of the probe set with the Epitoniidae, a diverse caenogastropod family of uncertain affinity and with poorly resolved evolutionary relationships, recovered a total of 2,850 loci. Although preliminary, the analysis of loci captured by our probe set for a small number of epitoniid taxa produced a well-resolved tree indicating that this probe set is also able to resolve relationships at shallower hierarchical scales. Together, the in silico and in vitro analyses indicate that target-capture enrichment with this probe set is a useful tool for reconstructing phylogenetic relationships across taxonomic levels and evolutionary time scales.

Craniodental divergence associated with bite force between hybridizing pine squirrels (Tamiasciurus).

Bite force can be a limiting factor in foraging and can significantly affect the competitive ability and lifetime fitness of mammals. Tamiasciurus squirrels feed primarily on conifer seeds and have a strong bite force to mechanically extract seeds from conifer cones with their mouths. In the North Cascades region, Douglas squirrels (Tamiasciurus douglasii) and red squirrels (T. hudsonicus) occupy ecologically different forests with different hardnesses in conifer cones. The ranges of these species overlap in a narrow hybrid zone where these forests meet near the crest of the North Cascades. We examined interspecific divergence in dietary ecomorphology in allopatry, in sympatry within the hybrid zone, and between hybrids and each parental species. We focused on three craniodental traits, including the incisor-strength index as a proxy measure for maximal bite force, cranial-suture complexity, and mandible shape. We find that these sister squirrel species differ in bite force and suture complexity in allopatry and sympatry and that mandible shape changes with the expected hardness of accessed food items, but is not significantly different between species. Furthermore, we find that hybrids display morphologies that overlap with hybrid zone red squirrels but not with hybrid zone Douglas squirrels. This work shows how important ecological processes at shallow evolutionary timescales can impact the divergence of morphological traits in taxa with extreme conservation of craniomandibular shape.

Orbital precession in short-period hot Jupiter exoplanet systems

ABSTRACT In several exoplanet systems the stellar rotation axis is not aligned with the normal to the orbital plane. For the class of ‘hot Jupiters’, a significant fraction of total angular momentum resides in the orbit. Orbit precessional motion has been observed in several such systems. We expect the tides raised by the orbiting exoplanet to induce normal mode oscillations in the host star, with the possibility of normal mode-orbit resonance. Gravity modes possess frequencies in the range of typical orbital Keplerian frequencies and their overtones. These resonances, confined to very narrow ranges of frequency space, would be highly improbable unless ‘resonance locking’ occurs, driven, for example, by structural and spin rate changes of the host star, operating on nuclear evolutionary time-scales. Resonance locking amplifies the amplitude of tidally driven oscillations, possibly by orders of magnitude, compared to the equilibrium tide displacements. We address the problem of precession and nutation in a system with a single exoplanet, with non-aligned spin and orbit axes, coupling the gravitational perturbations of normal mode distortions with orbital motion. Resonant modes are expected to have large amplitudes, contributing significantly to the gravitational perturbation already present due to rotational distortion of the star that gives rise to uniform orbital precession. The relative magnitude of rotational distortion and normal-mode perturbations is estimated. For Kepler-13Ab, estimates of their influence on transit time variations are given, and suggest they may be discernible with modern space telescope missions.

The Evolution of Mitochondrial Genomes between Two Cymbidium Sister Species: Dozens of Circular Chromosomes and the Maintenance and Deterioration of Genome Synteny.

Plant mitochondrial genomes (mitogenomes) exhibit fluid genome architectures, which could lead to the rapid erosion of genome synteny over a short evolutionary time scale. Among the species-rich orchid family, the leafy Cymbidium lancifolium and leafless Cymbidium macrorhizon are sister species with remarkable differences in morphology and nutritional physiology. Although our understanding of the evolution of mitochondria is incomplete, these sister taxa are ideal for examining this subject. In this study, the complete mitogenomes of C. lancifolium and C. macrorhizon, totaling 704,244 bp and 650,751 bp, respectively, were assembled. In the 2 mitogenomes, 38 protein-coding genes, 18 cis- and 6 trans-spliced introns, and approximately 611 Kb of homologous sequences are identical; overall, they have 99.4% genome-wide similarity. Slight variations in the mitogenomes of C. lancifolium and C. macrorhizon in repeat content (21.0 Kb and 21.6 Kb, respectively) and mitochondrial DNA of plastid origin (MIPT; 38.2 Kb and 37.5 Kb, respectively) were observed. The mitogenome architectures of C. lancifolium and C. macrorhizon are complex and comprise 23 and 22 mini-circular chromosomes, respectively. Pairwise comparisons indicate that the two mitogenomes are largely syntenic, and the disparity in chromosome numbers is likely due to repeat-mediated rearrangements among different chromosomes. Notably, approximately 93.2 Kb C. lancifolium mitochondrial sequences lack any homology in the C. macrorhizon mitogenome, indicating frequent DNA gains and losses, which accounts mainly for the size variation. Our findings provide unique insights into mitogenome evolution in leafy and leafless plants of sister species and shed light on mitogenome dynamics during the transition from mixotrophy to mycoheterotrophy.

Reverse Algols and hydrogen-rich Wolf-Rayet stars from very massive binaries

Massive star feedback affects the evolution of galaxies, where the most massive stars may have the largest impact. The majority of massive stars are born as members of close binary systems. In this work, we investigated detailed evolutionary models of very massive binaries (30−90 M⊙) with Large Magellanic Cloud (LMC) metallicity. We identify four effects defying the conventional knowledge of binary evolution, which are all related to the proximity of the models to the Eddington limit. We find that the majority of systems undergo mass transfer during core hydrogen burning. During the ensuing nuclear timescale evolution, many mass donors remain more massive than their companions (‘reverse Algols’), and nuclear timescale mass transfer may be interrupted or absent altogether. Furthermore, due to the elevated luminosity-to-mass ratio, many of the core-hydrogen-burning donors may develop Wolf-Rayet-type winds at luminosities where single stars would not. We identify observational counterparts of very massive reverse Algol binaries in the LMC and discuss their contribution to the observed hydrogen-rich Wolf-Rayet stars. We argue that understanding very massive Algol systems is key to predicting the advanced evolution of very massive binaries, including their ability to evolve into observable gravitational wave sources.

The dynamic adaptive landscape of cetacean body size

Adaptive landscapes are central to evolutionary theory, forming a conceptual bridge between micro- and macroevolution.1,2,3,4 Evolution by natural selection across an adaptive landscape should drive lineages toward fitness peaks, shaping the distribution of phenotypic variation within and among clades over evolutionary timescales.5 The location and breadth of these peaks in phenotypic space can also evolve,4 but whether phylogenetic comparative methods can detect such patterns has largely remained unexplored.6 Here, we characterize the global and local adaptive landscape for total body length in cetaceans (whales, dolphins, and relatives), a trait that spans an order of magnitude, across their ∼53-million-year evolutionary history. Using phylogenetic comparative methods, we analyze shifts in long-term mean body length7 and directional changes in average trait values8 for 345 living and fossil cetacean taxa. Remarkably, we find that the global macroevolutionary adaptive landscape of cetacean body length is relatively flat, with very few peak shifts occurring after cetaceans entered the oceans. Local peaks are more numerous and manifest as trends along branches linked to specific adaptations. These results contrast with previous studies using only extant taxa,9 highlighting the vital role of fossil data for understanding macroevolution.10,11,12 Our results indicate that adaptive peaks are dynamic and are associated with subzones of local adaptations, creating moving targets for species adaptation. In addition, we identify limits in our ability to detect some evolutionary patterns and processes and suggest that multiple approaches are required to characterize complex hierarchical patterns of adaptation in deep time.

Dynamic changes in Shiga toxin (Stx) 1 transducing phage throughout the evolution of O26:H11 Stx-producing Escherichia coli

Shiga toxin (Stx) is the key virulence factor of Stx-producing Escherichia coli (STEC). All known Stxs (Stx1 and Stx2) are encoded by bacteriophages (Stx phages). Although the genetic diversity of Stx phages has frequently been described, systematic analyses of Stx phages in a single STEC lineage are limited. In this study, focusing on the O26:H11 STEC sequence type 21 (ST21) lineage, where the stx1a gene is highly conserved, we analysed the Stx1a phages in 39 strains representative of the entire ST21 lineage and found a high level of variation in Stx1a phage genomes caused by various mechanisms, including replacement by a different Stx1a phage at the same or different locus. The evolutionary timescale of events changing Stx1a phages in ST21 was also determined. Furthermore, by using an Stx1 quantification system developed in this study, we found notable variations in the efficiency of Stx1 production upon prophage induction, which sharply contrasted with the conserved iron regulated Stx1 production. These variations were associated with the Stx1a phage alteration in some cases but not in other cases; thus, Stx1 production in this STEC lineage was determined by differences not only in Stx1 phages but also in host-encoded factors.

Sexually antagonistic selection maintains genetic variance when sexual dimorphism evolves.

Genetic variance (VG) in fitness related traits is often unexpectedly high, evoking the question how VG can be maintained in the face of selection. Sexually antagonistic (SA) selection favouring alternative alleles in the sexes is common and predicted to maintain VG, while directional selection should erode it. Both SA and sex-limited directional selection can lead to sex-specific adaptations but how each affect VG when sexual dimorphism evolves remain experimentally untested. Using replicated artificial selection on the seed beetle Callosobruchus maculatus body size we recently demonstrated an increase in size dimorphism under SA and male-limited (ML) selection by 50% and 32%, respectively. Here we test their consequences on genetic variation. We show that SA selection maintained significantly more ancestral, autosomal additive genetic variance than ML selection, while both eroded sex-linked additive variation equally. Ancestral female-specific dominance variance was completely lost under ML, while SA selection consistently sustained it. Further, both forms of selection preserved a high genetic correlation between the sexes (rm,f). These results demonstrate the potential for sexual antagonism to maintain more genetic variance while fuelling sex-specific adaptation in a short evolutionary time scale, and are in line with predicted importance of sex-specific dominance reducing sexual conflict over alternative alleles.