Emergent speciation by multiple Dobzhansky-Muller incompatibilities

Emergent speciation by multiple Dobzhansky-Muller incompatibilities

Tiago , Kevin E. Bassler, Ricardo B. R. Azevedo
doi: http://dx.doi.org/10.1101/008268

The Dobzhansky-Muller model posits that incompatibilities between alleles at different loci cause speciation. However, it is known that if the alleles involved in a Dobzhansky-Muller incompatibility (DMI) between two loci are neutral, the resulting reproductive isolation cannot be maintained in the presence of either mutation or gene flow. Here we propose that speciation can emerge through the collective effects of multiple neutral DMIs that cannot, individually, cause speciation-a mechanism we call emergent speciation. We investigate emergent speciation using a haploid neutral network model with recombination. We find that certain combinations of multiple neutral DMIs can lead to speciation. Complex DMIs and high recombination rate between the DMI loci facilitate emergent speciation. These conditions are likely to occur in nature. We conclude that the interaction between DMIs may be a root cause of the origin of species.

Exact solutions for the selection-mutation equilibrium in the Crow-Kimura evolutionary model

Exact solutions for the selection-mutation equilibrium in the Crow-Kimura evolutionary model

Yuri S. Semenov, Artem S. Novozhilov
(Submitted on 19 Aug 2014)

We reformulate the eigenvalue problem for the selection–mutation equilibrium distribution in the case of a haploid asexually reproduced population in the form of an equation for an unknown probability generating function of this distribution. The special form of this equation in the infinite sequence limit allows us to obtain analytically the steady state distributions for a number of particular cases of the fitness landscape. The general approach is illustrated by examples and theoretical findings are compared with numerical calculations.

Can the site-frequency spectrum distinguish exponential population growth from multiple-merger coalescents?

Can the site-frequency spectrum distinguish exponential population growth from multiple-merger coalescents?

Matthias Birkner, Jochen Blath, Bjarki Eldon, Fabian Freund
doi: http://dx.doi.org/10.1101/007690

The ability of the site-frequency spectrum (SFS) to reflect the particularities of gene genealogies exhibiting multiple mergers of ancestral lines as opposed to those obtained in the presence of exponential population growth is our focus. An excess of singletons is a well-known characteristic of both population growth and multiple mergers. Other aspects of the SFS, in particular the weight of the right tail, are, however, affected in specific ways by the two model classes. Using minimum-distance statistics, and an approximate likelihood method, our estimates of statistical power indicate that exponential growth can indeed be distinguished from multiple merger coalescents, even for moderate sample size, if the number of segregating sites is high enough. Additionally, we use a normalised version of the SFS as a summary statistic in an approximate bayesian computation (ABC) approach to distinguish multiple mergers from exponential population growth. The ABC approach gives further positive evidence as to the general eligibility of the SFS to distinguish between the different histories, but also reveals that suitable weighing of parts of the SFS can improve the distinction ability. The important issue of the difference in timescales between different coalescent processes (and their implications for the scaling of mutation parameters) is also discussed.

Reproductive isolation of hybrid populations driven by genetic incompatibilities

Reproductive isolation of hybrid populations driven by genetic incompatibilities

Molly Schumer, Rongfeng Cui, Gil G Rosenthal, Peter Andolfatto
doi: http://dx.doi.org/10.1101/007518

Despite its role in homogenizing populations, hybridization has also been proposed as a means to generate new species. The conceptual basis for this idea is that hybridization can result in novel phenotypes through recombination between the parental genomes, allowing a hybrid population to occupy ecological niches unavailable to parental species. A key feature of these models is that these novel phenotypes ecologically isolate hybrid populations from parental populations, precipitating speciation. Here we present an alternative model of the evolution of reproductive isolation in hybrid populations that occurs as a simple consequence of selection against incompatibilities. Unlike previous models, our model does not require small population sizes, the availability of new niches for hybrids or ecological or sexual selection on hybrid traits. We show that reproductive isolation between hybrids and parents evolves frequently and rapidly under this model, even in the presence of ongoing migration with parental species and strong selection against hybrids. Our model predicts that multiple distinct hybrid species can emerge from replicate hybrid populations formed from the same parental species, potentially generating patterns of species diversity and relatedness that mimic adaptive radiations.

Clonal interference and Muller’s ratchet in spatial habitats

Clonal interference and Muller’s ratchet in spatial habitats
Jakub Otwinowski, Joachim Krug
(Submitted on 18 Feb 2013 (v1), last revised 23 Jul 2014 (this version, v3))

Competition between independently arising beneficial mutations is enhanced in spatial populations due to the linear rather than exponential growth of clones. Recent theoretical studies have pointed out that the resulting fitness dynamics is analogous to a surface growth process, where new layers nucleate and spread stochastically, leading to the build up of scale-invariant roughness. This scenario differs qualitatively from the standard view of adaptation in that the speed of adaptation becomes independent of population size while the fitness variance does not. Here we exploit recent progress in the understanding of surface growth processes to obtain precise predictions for the universal, non-Gaussian shape of the fitness distribution for one-dimensional habitats, which are verified by simulations. When the mutations are deleterious rather than beneficial the problem becomes a spatial version of Muller’s ratchet. In contrast to the case of well-mixed populations, the rate of fitness decline remains finite even in the limit of an infinite habitat, provided the ratio Ud/s2 between the deleterious mutation rate and the square of the (negative) selection coefficient is sufficiently large. Using again an analogy to surface growth models we show that the transition between the stationary and the moving state of the ratchet is governed by directed percolation.

Fixation properties of subdivided populations with balancing selection

Fixation properties of subdivided populations with balancing selection

Pierangelo Lombardo, Andrea Gambassi, Luca Dall’Asta
Comments: 17 pages, 10 figures
Subjects: Populations and Evolution (q-bio.PE); Statistical Mechanics (cond-mat.stat-mech); Biological Physics (physics.bio-ph)

In subdivided populations, migration acts together with selection and genetic drift and determines their evolution. Building up on a recently proposed method, which hinges on the emergence of a time scale separation between local and global dynamics, we study the fixation properties of subdivided populations in the presence of balancing selection. The approximation implied by the method is accurate when the effective selection strength is small and the number of subpopulations is large. In particular, it predicts a phase transition between species coexistence and biodiversity loss in the infinite-size limit and, in finite populations, a nonmonotonic dependence of the mean fixation time on the migration rate. In order to investigate the fixation properties of the subdivided population for stronger selection, we introduce an effective coarser description of the dynamics in terms of a voter model with intermediate states, which highlights the basic mechanisms driving the evolutionary process.

Stress-Induced Mutagenesis and Complex Adaptation

Stress-Induced Mutagenesis and Complex Adaptation
Yoav Ram, Lilach Hadany
(Submitted on 14 Jul 2014)

Because mutations are mostly deleterious, mutation rates should be reduced by natural selection. However, mutations also provide the raw material for adaptation. Therefore, evolutionary theory suggests that the mutation rate must balance between adaptability – the ability to adapt – and adaptedness – the ability to remain adapted. We model an asexual population crossing a fitness valley and analyze the rate of complex adaptation with and without stress-induced mutagenesis – the increase of mutation rates in response to stress or maladaptation. We show that stress-induced mutagenesis increases the rate of complex adaptation without reducing the population mean fitness, thus breaking the evolutionary trade-off between adaptability and adaptedness. Our theoretical results support the hypothesis that stress-induced mutagenesis promotes adaptation and provide quantitative predictions of the rate of complex adaptation with different mutational strategies.

On the number of ranked species trees producing anomalous ranked gene trees

On the number of ranked species trees producing anomalous ranked gene trees
Filippo Disanto, Noah A. Rosenberg
Subjects: Populations and Evolution (q-bio.PE)

Analysis of probability distributions conditional on species trees has demonstrated the existence of anomalous ranked gene trees (ARGTs), ranked gene trees that are more probable than the ranked gene tree that accords with the ranked species tree. Here, to improve the characterization of ARGTs, we study enumerative and probabilistic properties of two classes of ranked labeled species trees, focusing on the presence or avoidance of certain subtree patterns associated with the production of ARGTs. We provide exact enumerations and asymptotic estimates for cardinalities of these sets of trees, showing that as the number of species increases without bound, the fraction of all ranked labeled species trees that are ARGT-producing approaches 1. This result extends beyond earlier existence results to provide a probabilistic claim about the frequency of ARGTs.

The site frequency spectrum of dispensable genes

The site frequency spectrum of dispensable genes
Franz Baumdicker
Comments: 24 pages, 8 figures
Subjects: Populations and Evolution (q-bio.PE); Probability (math.PR)

The differences between DNA-sequences within a population are the basis to infer the ancestral relationship of the individuals. Within the classical infinitely many sites model, it is possible to estimate the mutation rate based on the site frequency spectrum, which is comprised by the numbers $C_1,…,C_{n-1}$, where n is the sample size and $C_s$ is the number of site mutations (Single Nucleotide Polymorphisms, SNPs) which are seen in $s$ genomes. Classical results can be used to compare the observed site frequency spectrum with its neutral expectation, $E[C_s]= \theta_2/s$, where $\theta_2$ is the scaled site mutation rate. In this paper, we will relax the assumption of the infinitely many sites model that all individuals only carry homologous genetic material. Especially, it is today well-known that bacterial genomes have the ability to gain and lose genes, such that every single genome is a mosaic of genes, and genes are present and absent in a random fashion, giving rise to the dispensable genome. While this presence and absence has been modeled under neutral evolution within the infinitely many genes model in previous papers, we link presence and absence of genes with the numbers of site mutations seen within each gene. In this work we derive a formula for the expectation of the joint gene and site frequency spectrum, denotes $G_{k,s}$ the number of mutated sites occurring in exactly $s$ gene sequences, while the corresponding gene is present in exactly $k$ individuals. We show that standard estimators of $\theta_2$ for dispensable genes are biased and that the site frequency spectrum for dispensable genes differs from the classical result.

Convergent Evolution During Local Adaptation to Patchy Landscapes

Convergent Evolution During Local Adaptation to Patchy Landscapes
Peter L. Ralph, Graham Coop

Species often encounter, and adapt to, many patches of locally similar environmental conditions across their range. Such adaptation can occur through convergent evolution as different alleles arise and spread in different patches, or through the spread of alleles by migration acting to synchronize adaptation across the species. The tension between the two reflects the degree of constraint imposed on evolution by the underlying genetic architecture versus how how effectively selection acts to inhibit the geographic spread of locally adapted alleles. This paper studies a model of the balance between these two routes to adaptation in continuous environments with patchy selection pressures. We address the following questions: How long does it take for a new, locally adapted allele to appear in a patch of habitat where it is favored through new mutation? Or, through migration from another, already adapted patch? Which is more likely to occur, as a function of distance between the patches? How can we tell which has occurred, i.e.\ what population genetic signal is left by the spread of migrant alleles and how long does this signal persist for? To answer these questions we decompose the migration–selection equilibrium surrounding an already adapted patch into families of migrant alleles, in particular treating those rare families that reach new patches as spatial branching processes. This provides a way to understand the role of geographic separation between patches in promoting convergent adaptation and the genomic signals it leaves behind. We illustrate these ideas using the convergent evolution of cryptic coloration in the rock pocket mouse, Chaetodipus intermedius, as an empirical example.