Disruptive Selection Research Articles

Experimental evolution of adaptive divergence under varying degrees of gene flow.

Adaptive divergence is the key evolutionary process generating biodiversity by means of natural selection. Yet, the conditions under which it can arise in the presence of gene flow remain contentious. To address this question, we subjected 132 sexually reproducing fission yeast populations, sourced from two independent genetic backgrounds, to disruptive ecological selection and manipulated the level of migration between environments. Contrary to theoretical expectations, adaptive divergence was most pronounced when migration was either absent (allopatry) or maximal (sympatry), but was much reduced at intermediate rates (parapatry and local mating). This effect was apparent across central life-history components (survival, asexual growth and mating) but differed in magnitude between ancestral genetic backgrounds. The evolution of some fitness components was constrained by pervasive negative correlations (trade-off between asexual growth and mating), while others changed direction under the influence of migration (for example, survival and mating). In allopatry, adaptive divergence was mainly conferred by standing genetic variation and resulted in ecological specialization. In sympatry, divergence was mainly mediated by novel mutations enriched in a subset of genes and was characterized by the repeated emergence of two strategies: an ecological generalist and an asexual growth specialist. Multiple loci showed consistent evidence for antagonistic pleiotropy across migration treatments providing a conceptual link between adaptation and divergence. This evolve-and-resequence experiment shows that rapid ecological differentiation can arise even under high rates of gene flow. It further highlights that adaptive trajectories are governed by complex interactions of gene flow, ancestral variation and genetic correlations.

Genetic mapping of a novel powdery mildew resistance gene in wild emmer wheat from "Evolution Canyon" in Mt. Carmel Israel.

A single dominant powdery mildew resistance gene MlNFS10 was identified in wild emmer wheat and mapped within a 0.3cM genetic interval spanning a 2.1Mb physical interval on chromosome arm 4AL. Wheat powdery mildew caused by Blumeria graminis forma specialis tritici (Bgt) is a globally devastating disease. The use of powdery mildew resistance genes from wild relatives of wheat is an effective method of disease management. Our previous research has shown that disruptive ecological selection has driven the discrete adaptations of the wild emmer wheat population on the south facing slope (SFS) and north facing slope (NFS) at the microsite of "Evolution Canyon" at Mount Carmel, Israel and demonstrated that 16 accessions in the NFS population display high resistance to 11 powdery mildew isolates (collected from different wheat fields in China). Here, we constructed bi-parental population by crossing the accession NFS-10 (resistant to 22 Bgt races collected from China in seedling resistance screen) and the susceptible line SFS2-12. Genetic analysis indicated that NFS-10 carries a single dominant gene, temporarily designated MlNFS10. Ultimately, 13 markers were successfully located within the long arm of chromosome 4A, thereby delineating MlNFS10 to a 0.3cM interval covering 2.1Mb (729275816-731365462) in the Chinese Spring reference sequence. We identified disease resistance-associated genes based on the RNA-seq analysis of both parents. The tightly linked InDel marker XWsdau73447 and SSR marker XWsdau72928 were developed and used for marker-assisted selection when MlNFS10 was introgressed into a hexaploid wheat background. Therefore, MlNFS10 can be used for improvement of germplasm in breeding programs for powdery mildew resistant cultivars.

Hybrid Cuckoo Search for the Capacitated Vehicle Routing Problem

Having the best solution for Vehicle Routing Problem (VRP) is still in demand. Beside, Cuckoo Search (CS) is a popular metaheuristic based on the reproductive strategy of the Cuckoo species and has been successfully applied in various optimizations, including Capacitated Vehicle Routing Problem (CVRP). Although CS and hybrid CS have been proposed for CVRP, the performance of CS is far from the state-of-art. Therefore, this study proposes a hybrid CS with Simulated Annealing (SA) algorithm for the CVRP, consisting of three improvements—the investigation of 12 neighborhood structures, three selections strategy and hybrid it with SA. The experiment was conducted using 16 instances of the Augerat benchmark dataset. The results show that 6 out of 12 neighborhood structures were the best and the disruptive selection strategy is the best strategy. The experiments’ results showed that the proposed method could find optimal and near-optimal solutions compared with state-of-the-art algorithms.

The strength and form of natural selection on transcript abundance in the wild.

Gene transcription variation is known to contribute to disease susceptibility and adaptation, but we currently know very little about how contemporary natural selection shapes transcript abundance. Here, we propose a novel analytical framework to quantify the strength and form of ongoing natural selection at the transcriptome level in a wild vertebrate. We estimated selection on transcript abundance in a cohort of a wild salmonid fish (Salmo trutta) affected by an extracellular myxozoan parasite (Tetracapsuloides bryosalmonae) through mark–recapture field sampling and the integration of RNA‐sequencing with classical regression‐based selection analysis. We show, based on fin transcriptomes of the host, that infection by the parasite and subsequent host survival is linked to upregulation of mitotic cell cycle process. We also detect a widespread signal of disruptive selection on transcripts linked to host immune defence, host–pathogen interactions, cellular repair and maintenance. Our results provide insights into how selection can be measured at the transcriptome level to dissect the molecular mechanisms of contemporary evolution driven by climate change and emerging anthropogenic threats. We anticipate that the approach described here will enable critical information on the molecular processes and targets of natural selection to be obtained in real time.

Sex differences in survival: viability selection in nymphal tree crickets from a conopid fly parasitoid

1. Variation in mortality due to parasitoids is expected to be affected by host sex and size.2. We use selection analyses to examine the effects of Stylogaster neglecta (Diptera: Conopidae), a recently discovered parasitoid of nymphal Oecanthus nigricornis (Orthoptera: Gryllidae), on sex and size viability in the host crickets.3. O. nigricornis nymphs were sampled across Ontario and New York State and dissected to determine whether they were parasitised. Logistic regression was used to examine parasitism by host body size and sex.4. There was strong evidence for an effect of sex and body size on parasitism, with males and smaller crickets being more likely to be parasitised and containing more parasitoids in them. There was also weaker evidence of an interaction effect between sex and body size, with body size being a significant predictor of parasitism only in males. Using a selection analysis, we found evidence of positive and disruptive selection on male body size.5. Causes of these viability differences may include sex differences in larval cricket immune function, riskier behaviour of male nymphs as they acquire resources necessary for adult trait development, as well as the effects of S. neglecta flies targeting hosts that maximise fitness.

Evolutionary trade‐offs may interact with physiological constraints to maintain color variation

AbstractAnimal coloration is a multifaceted trait with many ecological roles and related to a variety of developmental and physiological processes. Consequently, coloration is often subject to a variety of selective pressures, leading to the evolutionary maintenance of variation. In this study, we investigated hypotheses related to the maintenance of dorsal color variation in wood frogs (Rana sylvatica). First, we tested for multimodality, and whether color correlates with body size or condition or varies by sex or age class. We combined behavioral trials with visual modeling to test for sex recognition. We also considered visual models for predators and tested for an interaction between discriminability indexes (JND) of color channel (chromatic vs. achromatic) and predator type (birds vs. snakes), as well as for a within individual trade‐off between the JND of chromatic and achromatic coloration. Finally, we tested for disruptive viability selection on color using predation trials, and for antagonistic directional selection between viability selection and reproductive investment of females. We found that wood frogs present continuous color variation that does not correlate with body size or condition, but that changes with age. Wood frogs present subtle sexual dichromatism, but we found no evidence for a role of color in sex recognition. Instead, we discuss the possibility that sex differences might, at least in part, have a demographic explanation. Predator visual models indicated that wood frogs cannot solely rely on dorsal coloration for camouflage. Moreover, different predators might present selective pressures in different color channels, while individuals’ achromatic and chromatic coloration trade‐off in JND. Therefore, different selective pressures caused by different predators might interact with ontogenetic changes and developmental/physiological trade‐offs to maintain color variation. We found no relationship between color and survival or reproductive investment, suggesting further work is required to fully understand selection on color. Our results highlight the importance of understanding evolutionary trade‐offs and developmental/physiological constraints in combination with one another, and suggest the potential for an interaction between these proximate and ultimate mechanisms in the evolutionary maintenance of variation. These results likely extend beyond color expression in amphibians, and exemplify a more general process for such evolutionary outcomes.

Evidence of parallel evolution in the dental elements of Sweetognathus conodonts.

The repeated emergence of similar morphologies in the dental elements of Permian Sweetognathus conodonts has been a hypothesized example of parallel evolution. To test if morphological parallelisms occur between isolated Sweetognathus lineages, this study uses two-dimensional-based geometric morphometrics combined with a revised and expanded phylogeny of Permian Sweetognathus conodonts to quantify dental element trait distributions and compare the phenotypic trajectories between lineages. A hierarchical clustering method was used to identify recurrent species pairs based on principal component scores describing their morphological variation, with the further incorporation of widely used ecological metrics such as limiting similarity and morphological overlap. Our research implies that a major contributor to conodont diversity in Palaeozoic marine trophic networks is the emergence of recurrent parallel morphologies via disruptive and directional selection. This study illustrates the mechanisms through which conodonts achieved their status as hyper-diverse predators and scavengers, contributing substantially to the complexity of Palaeozoic marine communities.

Disruptive natural selection by male reproductive potential prevents underexpression of protein-coding genes on the human Y chromosome as a self-domestication syndrome

BackgroundIn population ecology, the concept of reproductive potential denotes the most vital indicator of chances to produce and sustain a healthy descendant until his/her reproductive maturity under the best conditions. This concept links quality of life and longevity of an individual with disease susceptibilities encoded by his/her genome. Female reproductive potential has been investigated deeply, widely, and comprehensively in the past, but the male one has not received an equal amount of attention. Therefore, here we focused on the human Y chromosome and found candidate single-nucleotide polymorphism (SNP) markers of male reproductive potential.ResultsExamining in silico (i.e., using our earlier created Web-service SNP_TATA_Z-tester) all 1206 unannotated SNPs within 70 bp proximal promoters of all 63 Y-linked genes, we found 261 possible male-reproductive-potential SNP markers that can significantly alter the binding affinity of TATA-binding protein (TBP) for these promoters. Among them, there are candidate SNP markers of spermatogenesis disorders (e.g., rs1402972626), pediatric cancer (e.g., rs1483581212) as well as male anxiety damaging family relationships and mother’s and children’s health (e.g., rs187456378). First of all, we selectively verified in vitro both absolute and relative values of the analyzed TBP–promoter affinity, whose Pearson’s coefficients of correlation between predicted and measured values were r = 0.84 (significance p < 0.025) and r = 0.98 (p < 0.025), respectively. Next, we found that there are twofold fewer candidate SNP markers decreasing TBP–promoter affinity relative to those increasing it, whereas in the genome-wide norm, SNP-induced damage to TBP–promoter complexes is fourfold more frequent than SNP-induced improvement (p < 0.05, binomial distribution). This means natural selection against underexpression of these genes. Meanwhile, the numbers of candidate SNP markers of an increase and decrease in male reproductive potential were indistinguishably equal to each other (p < 0.05) as if male self-domestication could have happened, with its experimentally known disruptive natural selection. Because there is still not enough scientific evidence that this could have happened, we discuss the human diseases associated with candidate SNP markers of male reproductive potential that may correspond to domestication-related disorders in pets.ConclusionsOverall, our findings seem to support a self-domestication syndrome with disruptive natural selection by male reproductive potential preventing Y-linked underexpression of a protein.

Resource competition shapes biological rhythms and promotes temporal niche differentiation in a community simulation.

Competition for resources often contributes strongly to defining an organism's ecological niche. Endogenous biological rhythms are important adaptations to the temporal dimension of niches, but how other organisms influence such temporal niches has not been much studied, and the role of competition in particular has been even less examined. We investigated how interspecific competition and intraspecific competition for resources shape an organism's activity rhythms.To do this, we simulated communities of one or two species in an agent‐based model. Individuals in the simulation move according to a circadian activity rhythm and compete for limited resources. Probability of reproduction is proportional to an individual's success in obtaining resources. Offspring may have variance in rhythm parameters, which allow for the population to evolve over time.We demonstrate that when organisms are arrhythmic, one species will always be competitively excluded from the environment, but the existence of activity rhythms allows niche differentiation and indefinite coexistence of the two species. Two species which are initially active at the same phase will differentiate their phase angle of entrainment over time to avoid each other. When only one species is present in an environment, competition within the species strongly selects for niche expansion through arrhythmicity, but the addition of an interspecific competitor facilitates evolution of increased rhythmic amplitude when combined with additional adaptations for temporal specialization. Finally, if individuals preferentially mate with others who are active at similar times of day, then disruptive selection by intraspecific competition can split one population into two reproductively isolated groups separated in activity time.These simulations suggest that biological rhythms are an effective method to temporally differentiate ecological niches and that competition is an important ecological pressure promoting the evolution of rhythms and sleep. This is the first study to use ecological modeling to examine biological rhythms.

The components of directional and disruptive selection in heterogeneous group-structured populations

We derive how directional and disruptive selection operate on scalar traits in a heterogeneous group-structured population for a general class of models. In particular, we assume that each group in the population can be in one of a finite number of states, where states can affect group size and/or other environmental variables, at a given time. Using up to second-order perturbation expansions of the invasion fitness of a mutant allele, we derive expressions for the directional and disruptive selection coefficients, which are sufficient to classify the singular strategies of adaptive dynamics. These expressions include first- and second-order perturbations of individual fitness (expected number of settled offspring produced by an individual, possibly including self through survival); the first-order perturbation of the stationary distribution of mutants (derived here explicitly for the first time); the first-order perturbation of pairwise relatedness; and reproductive values, pairwise and three-way relatedness, and stationary distribution of mutants, each evaluated under neutrality. We introduce the concept of individual k-fitness (defined as the expected number of settled offspring of an individual for which k-1 randomly chosen neighbors are lineage members) and show its usefulness for calculating relatedness and its perturbation. We then demonstrate that the directional and disruptive selection coefficients can be expressed in terms individual k-fitnesses with k=1,2,3 only. This representation has two important benefits. First, it allows for a significant reduction in the dimensions of the system of equations describing the mutant dynamics that needs to be solved to evaluate explicitly the two selection coefficients. Second, it leads to a biologically meaningful interpretation of their components. As an application of our methodology, we analyze directional and disruptive selection in a lottery model with either hard or soft selection and show that many previous results about selection in group-structured populations can be reproduced as special cases of our model.

Sympatric ecological divergence with coevolution of niche preference.

Reinforcement, the increase of assortative mating driven by selection against unfit hybrids, is conditional on pre-existing divergence. Yet, for ecological divergence to precede the evolution of assortment, strict symmetries between fitnesses in niches must hold, and/or there must be low gene flow between the nascent species. It has thus been argued that conditions favouring sympatric speciation are rarely met in nature. Indeed, we show that under disruptive selection, violating symmetries in niche sizes and increasing strength of the trade-off in selection between the niches quickly leads to loss of genetic variation, instead of evolution of specialists. The region of the parameter space where polymorphism is maintained further narrows with increasing number of loci encoding the diverging trait and the rate of recombination between them. Yet, evolvable assortment and pre-existing assortment both substantially broaden the parameter space within which polymorphism is maintained. Notably, pre-existing niche preference speeds up further increase of assortment, thus facilitating reinforcement in the later phases of speciation. We conclude that in order for sympatric ecological divergence to occur, niche preference must coevolve throughout the divergence process. Even if populations come into secondary contact, having diverged in isolation, niche preference substantially broadens the conditions for coexistence in sympatry and completion of the speciation process.This article is part of the theme issue ‘Towards the completion of speciation: the evolution of reproductive isolation beyond the first barriers'.

How geographic productivity patterns affect food-web evolution

It is well recognized that spatial heterogeneity and overall productivity have important consequences for the diversity and community structure of food webs. Yet, few, if any, studies have considered the effects of heterogeneous spatial distributions of primary production. Here, we theoretically investigate how the variance and autocorrelation length of primary production affect properties of evolved food webs consisting of one autotroph and several heterotrophs. We report the following findings. (1) Diversity increases with landscape variance and is unimodal in autocorrelation length. (2) Trophic level increases with landscape variance and is unimodal in autocorrelation length. (3) The extent to which the spatial distribution of heterotrophs differ from that of the autotroph increases with landscape variance and decreases with autocorrelation length. (4) Components of initial disruptive selection experienced by the ancestral heterotroph predict properties of the final evolved communities. Prior to our study reported here, several authors had hypothesized that diversity increases with the landscape variance of productivity. Our results support their hypothesis and contribute new facets by providing quantitative predictions that also account for autocorrelation length and additional properties of the evolved communities.

Functional Tradeoffs Carry Phenotypes Across the Valley of the Shadow of Death.

Functional tradeoffs are often viewed as constraints on phenotypic evolution, but they can also facilitate evolution across the suboptimal valleys separating performance peaks. I explore this process by reviewing a previously published model of how disruptive selection from competing functional demands defines an intermediate performance optimum for morphological systems that cannot simultaneously be optimized for all of the functional roles they must play. Because of the inherent tradeoffs in such a system, its optimal morphology in any particular environmental context will usually be intermediate between the performance peaks of the competing functions. The proportional contribution of each functional demand can be estimated by maximum likelihood from empirically observed morphologies, including complex ones measured with multivariate geometric morphometrics, using this model. The resulting tradeoff weight can be mapped onto a phylogenetic tree to study how the performance optimum has shifted across a functional landscape circumscribed by the function-specific performance peaks. This model of tradeoff evolution is sharply different from one in which a multipeak Ornstein-Uhlenbeck (OU) model is applied to a set of morphologies and a phylogenetic tree to estimate how many separate performance optima exist. The multi-peak OU approach assumes that each branch is pushed toward one of two or more performance peaks that exist simultaneously and are separated by valleys of poor performance, whereas the model discussed here assumes that each branch tracks a single optimal performance peak that wanders through morphospace as the balance of functional demands shifts. That the movements of this net performance peak emerge from changing frequencies of selection events from opposing functional demands are illustrated using a series of computational simulations. These simulations show how functional tradeoffs can carry evolution across putative performance valleys: even though intermediate morphologies may not perform optimally for any one function, they may represent the optimal solution in any environment in which an organism experiences competing functional demands.

Ecological opportunity and upward prey-predator radiation cascades

A general goal in community ecology and evolutionary biology is to understand how diversity has arisen. In our attempts to reach such goals we become increasingly aware of interacting ecological and evolutionary processes shaping biodiversity. Ecological opportunity and adaptive radiations can, for example, drive diversification in competitive communities but little is known about how such processes propagate through trophic levels in adaptive radiation cascades. I use an eco-evolutionary model of trait-based ecological interactions and micro-evolutionary processes to investigate the macro-evolutionary aspects of predator diversification in such cascades. Prey diversification facilitates predator radiation through predator feeding opportunity and disruptive selection. Predator radiation, however, often disconnects from the prey radiation as the diversification progresses. Only when predators have an intermediate niche width, high predatory efficiency, and high evolutionary potential can radiation cascades be maintained over macro-evolutionary time scales. These results provide expectations for predator response to prey divergence and insight into eco-evolutionary feedbacks between trophic levels. Such expectations are crucial for future studies that aim for a better understanding of how diversity is generated and maintained in complex communities.

Anthropogenic and natural size-related selection act in concert during brown trout (Salmo trutta) smolt river descent

By hindering migration and inducing direct turbine mortality during downstream migration, hydropower is regarded as one of the most serious threats to anadromous salmonids. Yet, little attention has been paid to long-term turbine-induced selection mechanisms effecting fish populations. This work evaluates turbine and post-turbine survival of PIT-tagged wild brown trout smolts. By estimating individual river and sea survival rates, we were able to compare survival rates of smolts that had migrated through the turbine with smolts that had bypassed the turbine, as well as investigate both natural and anthropogenic size-selective mechanisms operative on the population. Total river-descent survival probability was 0.20 for turbine migrants and 0.44 for bypass migrants. The surviving turbine migrants were significantly smaller than their bypass counterparts and more exposed to predation from Northern pike. The estimated mean-adjusted selection gradient was − 0.76 for turbine migrants and + 1.85 for the bypass migrants. The resulting disruptive selection may ultimately lead to increased phenotypic smolt size variation provided sufficient additive genetic variance associated with smolt size. Mitigation measures at hydropower plants are thus essential for preserving sustainable populations of anadromous fish and maintaining population genetic variation.

Biogeography of shell morphology in over‐exploited shellfish reveals adaptive trade‐offs on human‐inhabited islands and incipient selectively driven lineage bifurcation

AbstractAimHumans are unintentionally affecting the evolution of fishery species directly through exploitation and indirectly by altering climate. We aim to test for a relationship between biogeographic patterns in the shell phenotypes of an over‐exploited shellfish and the presence of humans to identify human‐mediated adaptive trade‐offs. The implications of these trade‐offs are discussed with respect to the sustainability of the fishery.TaxonThe endemic Hawaiian intertidal limpet, ‘opihi makaiauli (Patellagastropoda, Nacellidae, Cellana exarata)MethodsWe surveyed phenotypic characters associated with temperature and predation avoidance across the entire species range and tested for differences in the relationship between these characters and latitude, on islands with and without humans.ResultsAmong all limpets surveyed, there was a bimodal distribution in shell colour (light, dark) and a parapatric pattern of shell coloration across the archipelago with lighter shells being prevalent on the uninhabited islands and darker, more camouflaged shells being prevalent on the inhabited islands. On the cooler, uninhabited islands, all morphometric characters associated with thermal avoidance (surface area, height and doming) increased with decreasing latitude. On the hotter, inhabited islands, however, shells were flatter, less variable and less adapted for avoiding thermal stress than predation.Main ConclusionsThe biogeographic patterns in shell phenotype and previous genetic studies suggest that the population is beginning to bifurcate in response to disruptive and directional selection as well as geographic isolation between the islands with and without humans. Decreased phenotypic and genetic diversity on the inhabited islands despite much larger populations of ‘opihi suggests a prominent historical bottleneck. The prevalence of maladaptive dark, flat phenotypes for thermal avoidance on the inhabited islands suggests that predation is a stronger selective force, driving adaptive trade‐offs in shape and colour. We propose that this is likely a case of fisheries‐induced evolution and a millennium of harvesting is the most likely selective pressure driving the observed biogeographic patterns in shell morphology. The flatter, darker shells will allow body temperatures to rise higher in direct sunlight, therefore we hypothesize that the thermal niche of ‘opihi is narrower on inhabited islands and will continue to narrow as Earth warms.

Evolution of dispersal in a spatially heterogeneous population with finite patch sizes

Dispersal is one of the fundamental life-history strategies of organisms, so understanding the selective forces shaping the dispersal traits is important. In the Wright's island model, dispersal evolves due to kin competition even when dispersal is costly, and it has traditionally been assumed that the living conditions are the same everywhere. To study the effect of spatial heterogeneity, we extend the model so that patches may receive different amounts of immigrants, foster different numbers of individuals, and give different reproduction efficiency to individuals therein. We obtain an analytical expression for the fitness gradient, which shows that directional selection consists of three components: As in the homogeneous case, the direct cost of dispersal selects against dispersal and kin competition promotes dispersal. The additional component, spatial heterogeneity, more precisely the variance of so-called relative reproductive potential, tends to select against dispersal. We also obtain an expression for the second derivative of fitness, which can be used to determine whether there is disruptive selection: Unlike the homogeneous case, we found that divergence of traits through evolutionary branching is possible in the heterogeneous case. Our numerical explorations suggest that evolutionary branching is promoted more by differences in patch size than by reproduction efficiency. Our results show the importance of the existing spatial heterogeneity in the real world as a key determinant in dispersal evolution.

Sympatric speciation of wild emmer wheat driven by ecology and chromosomal rearrangements

In plants, the mechanism for ecological sympatric speciation (SS) is little known. Here, after ruling out the possibility of secondary contact, we show that wild emmer wheat, at the microclimatically divergent microsite of "Evolution Canyon" (EC), Mt. Carmel, Israel, underwent triple SS. Initially, it split following a bottleneck of an ancestral population, and further diversified to three isolated populations driven by disruptive ecological selection. Remarkably, two postzygotically isolated populations (SFS1 and SFS2) sympatrically branched within an area less than 30 m at the tropical hot and dry savannoid south-facing slope (SFS). A series of homozygous chromosomal rearrangements in the SFS1 population caused hybrid sterility with the SFS2 population. We demonstrate that these two populations developed divergent adaptive mechanisms against severe abiotic stresses on the tropical SFS. The SFS2 population evolved very early flowering, while the SFS1 population alternatively evolved a direct tolerance to irradiance by improved ROS scavenging activity that potentially accounts for its evolutionary fate with unstable chromosome status. Moreover, a third prezygotically isolated sympatric population adapted on the abutting temperate, humid, cool, and forested north-facing slope (NFS), separated by 250 m from the SFS wild emmer wheat populations. The NFS population evolved multiple resistant loci to fungal diseases, including powdery mildew and stripe rust. Our study illustrates how plants sympatrically adapt and speciate under disruptive ecological selection of abiotic and biotic stresses.

Evolutionary Geography of Root Nodule Bacteria: Speciation Directed by the Host Plants

Compared to free-living relatives, bacterial symbionts of plants exhibit higher rates of population diversification (microevolution), as well as of speciation and formation of super-species taxa (macroevolution). Analysis of rhizobia, the N2-fixing symbionts of leguminous plants, suggests that both evolutionary modes are determined by the hosts, which have mutagenic and recombinogenic effects on their microsymbionts, apart from activating the selective processes that determine the fate of newly emerged genotypes in their populations. Migration induced by leguminous plants plays a key role in the evolution of rhizobia, including: (1) joint movement of the symbionts and their hosts into new ecological zones and (2) local circulation of the symbionts in plant‒soil systems. In the first case, allopatric speciation occurs in rhizobia due to transfer of the symbiotically specialized (sym) genes from the introduced strains into local bacteria, which are transformed into novel symbionts. In the second case, a sympatric divergence of Rhizobium populations occurs, resulting in formation of intraspecies taxa (biovars and symbiovars) contrasting in their host ranges. In the evolutionarily advanced rhizobia, Rhizobium leguminosarum and Neorhizobium galegae interacting with galegoid legumes (tribes Fabeae, Galegae, and Trifolieae), these processes are associated with diversification of the sym genes representing the accessory genome parts due to the interaction between bacteria and the hosts differing in their symbiotic affinities. In parallel, formation of new cryptic (genomic) species was revealed in local rhizobia populations, which is associated with divergence of the core genome parts. It may be caused either by genetic drift or by selection for resistance against local edaphic stresses, which in combination with disruptive selection can sufficiently enhance the rhizobia diversity and change the structures of their pangenomes.

Peculiarities of microevolution processes in Trollius asiaticus in South Siberia Mountains

In the paper, we analyse the distribution of species and forms closely-related to Trollius asiaticus L. growing in the mountains of South Siberia between the Altai Mountain Country and Baikalia. The interrelationship between the endemic species and forms of high-mountains and parental lowland forms are adjacent-sympatric. We suppose the ways of disruptive selection to be of the same type, thus resulting in T. sajanensis and T. altaicus speciation, which are twin-species to each other. Between the parental and daughter species, there are interlayers of morphological forms being introgressive to both species. Key words: characters, populations, disruptive selection, introgression, speciation.