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A colorful legacy of hybridization in wood-warblers includes frequent sharing of carotenoid genes among species and genera.

Introgression between species has the potential to shape evolutionary trajectories in important ways, but uncovering complex introgression dynamics has only recently been made possible by advances in genomics. Warblers of the avian family Parulidae exemplify rapid diversification and sexual trait divergence, and we endeavored to study historical introgression in the family. We sequenced multiple genomes of nearly every species, constructed a phylogeny for the family, and investigated gene flow across the genome and at genes known for controlling feather color. We found that DNA haplotypes including the gene BCO2, which encodes an enzyme that degrades yellow carotenoids, have spread among genera multiple times-from Vermivora to Geothlypis and from Leiothlypis to multiple Cardellina and Setophaga species. Patterns of inheritance in the latter case point to introgression that occurred 0.5 to 2 million years ago, and the shared haplotype among recipient species is less than 100 nucleotides long. Separately, we found evidence of introgression from red Cardellina species to both of the two red Myioborus species at BDH1L and from one red Myioborus species to the other at CYP2J19; both are key genes in the pathway that converts yellow carotenoids to red ketocarotenoids. Our results show that introgression is a common mechanism for the evolution of colorful plumage in parulid warblers and hint at complex histories of gene flow behind some of the Western Hemisphere's most colorful birds.

Animals↗

Modeling genetic networks and their evolution: a complex dynamical systems perspective.

After finishing the sequence of the human genome, a functional understanding of genome dynamics is the next major step on the agenda of the biosciences. New approaches, such as microarray techniques, and new methods of bioinformatics provide powerful tools aiming in this direction. In the last few years, important parts of genome organization and dynamics in a number of model organisms have been determined. However, an integrated view of gene regulation on a genomic scale is still lacking. Here, genome function is discussed from a complex dynamical systems perspective: which dynamical properties can a large genomic system exhibit in principle, given the local mechanisms governing the small subsystems that we know today? Models of artificial genetic networks are used to explore dynamical principles and possible emergent dynamical phenomena in networks of genetic switches. One observes evolution of robustness and dynamical self-organization in large networks of artificial regulators that are based on the dynamic mechanism of transcriptional regulators as observed in biological gene regulation. Possible biological observables and ways of experimental testing of global phenomena in genome function and dynamics are discussed. Models of artificial genetic networks provide a tool to address questions in genome dynamics and their evolution and allow simulation studies in evolutionary genomics.

Biological Evolution↗

What we have also learned: adaptive speciation is theoretically plausible.

A recent Perspectives article by Gavrilets (2003) on the theory of speciation ignored advances in understanding processes of adaptive speciation, in which the splitting of lineages is an adaptation caused by frequency-dependent selection. Adaptive, or sympatric, speciation has been modeled since the 1960s, but the large amount of attention from both empirical and theoretical biologists that adaptive speciation has received in recent years goes far beyond what was described in Gavrilets' paper. Due to conceptual advances based on the theory of adaptive dynamics, adaptive speciation has emerged as a theoretically plausible evolutionary process that can occur in many different ecological settings.

Adaptation, Biological↗

Advantage of storage in a fluctuating environment.

We will elaborate the evolutionary course of an ecosystem consisting of a population in a chemostat environment with periodically fluctuating nutrient supply. The organisms that make up the population consist of structural biomass and energy storage compartments. In a constant chemostat environment a species without energy storage always out-competes a species with energy reserves. This hinders evolution of species with storage from those without storage. Using the adaptive dynamics approach for non-equilibrium ecological systems we will show that in a fluctuating environment there are multiple stable evolutionary singular strategies (ss's): one for a species without, and one for a species with energy storage. The evolutionary end-point depends on the initial evolutionary state. We will formulate the invasion fitness in terms of Floquet multipliers for the oscillating non-autonomous system. Bifurcation theory is used to study points where due to evolutionary development by mutational steps, the long-term dynamics of the ecological system changes qualitatively. To that end, at the ecological time scale, the trait value at which invasion of a mutant into a resident population becomes possible can be calculated using numerical bifurcation analysis where the trait is used as the free parameter, because it is just a bifurcation point. In a constant environment there is a unique stable equilibrium for one species following the "competitive exclusion" principle. In contrast, due to the oscillatory dynamics on the ecological time scale two species may coexist. That is, non-equilibrium dynamics enhances biodiversity. However, we will show that this coexistence is not stable on the evolutionary time scale and always one single species survives.

Adaptation, Physiological↗

Evolution of plastid gene rps2 in a lineage of hemiparasitic and holoparasitic plants: many losses of photosynthesis and complex patterns of rate variation.

The plastid genomes of some nonphotosynthetic parasitic plants have experienced an extreme reduction in gene content and an increase in evolutionary rate of remaining genes. Nothing is known of the dynamics of these events or whether either is a direct outcome of the loss of photosynthesis. The parasitic Scrophulariaceae and Orobanchaceae, representing a continuum of heterotrophic ability ranging from photosynthetic hemiparasites to nonphotosynthetic holoparasites, are used to investigate these issues. We present a phylogenetic hypothesis for parasitic Scrophulariaceae and Orobanchaceae based on sequences of the plastid gene rps2, encoding the S2 subunit of the plastid ribosome. Parasitic Scrophulariaceae and Orobanchaceae form a monophyletic group in which parasitism can be inferred to have evolved once. Holoparasitism has evolved independently at least five times, with certain holoparasitic lineages representing single species, genera, and collections of nonphotosynthetic genera. Evolutionary loss of the photosynthetic gene rbcL is limited to a subset of holoparasitic lineages, with several holoparasites retaining a full length rbcL sequence. In contrast, the translational gene rps2 is retained in all plants investigated but has experienced rate accelerations in several hemi- as well as holoparasitic lineages, suggesting that there may be substantial molecular evolutionary changes to the plastid genome of parasites before the loss of photosynthesis. Independent patterns of synonymous and nonsynonymous rate acceleration in rps2 point to distinct mechanisms underlying rate variation in different lineages. Parasitic Scrophulariaceae (including the traditional Orobanchaceae) provide a rich platform for the investigation of molecular evolutionary process, gene function, and the evolution of parasitism.

Arabidopsis Proteins↗

Dynamics of disease resistance polymorphism at the Rpm1 locus of Arabidopsis.

The co-evolutionary 'arms race' is a widely accepted model for the evolution of host-pathogen interactions. This model predicts that variation for disease resistance will be transient, and that host populations generally will be monomorphic at disease-resistance (R-gene) loci. However, plant populations show considerable polymorphism at R-gene loci involved in pathogen recognition. Here we have tested the arms-race model in Arabidopsis thaliana by analysing sequences flanking Rpm1, a gene conferring the ability to recognize Pseudomonas pathogens carrying AvrRpm1 or AvrB. We reject the arms-race hypothesis: resistance and susceptibility alleles at this locus have co-existed for millions of years. To account for the age of alleles and the relative levels of polymorphism within allelic classes, we use coalescence theory to model the long-term accumulation of nucleotide polymorphism in the context of the short-term ecological dynamics of disease resistance. This analysis supports a 'trench warfare' hypothesis, in which advances and retreats of resistance-allele frequency maintain variation for disease resistance as a dynamic polymorphism.

Arabidopsis↗

Self-organized criticality in spatial evolutionary game theory.

Self-organized criticality is an important framework for understanding the emergence of scale-free natural phenomena. Cellular automata provide simple interesting models in which to study self-organized criticality. We consider the dynamics of a new class of cellular automata which are constructed as natural spatial extensions of evolutionary game theory. This construction yields a discrete one-parameter family of cellular automata. We show that there is a range of parameter values for which this system exhibits complex dynamics with long range correlations between states in both time and space. In this region the dynamics evolve to a self-organized critical state in which structures exist on all time and length scales, and the relevant statistical measures have power law behaviour.

Animals↗

Evolutionary self-organising modelling of a municipal wastewater treatment plant.

Building predictive models for highly time varying and complex multivariable aspects of the wastewater treatment plant is important both for understanding the dynamics of this complex system, and in the development of optimal control support and management schemes. This paper presents a new approach, which is called genetic programming as a self-organising modelling tool, to model dynamic performance of municipal activated-sludge wastewater treatment plants. Genetic programming evolves several process models automatically based on methods of natural selection ('survival of the fittest'), that could predict the dynamics of MLSS and suspended solids in the effluent. The predictive accuracy of the genetic programming approach was compared with a nonlinear state-space model with neural network and a well-known IAWQ ASM2. The genetic programming system evolved some models that were an improvement over the neural network and ASM2 and showed that the transparency of the model evolved may allow inferences about underlying processes to be made. This work demonstrates that dynamic nonlinear processes in the wastewater treatment plant may be successfully modelled through the use of evolutionary model induction algorithms in GP technique. Further, our results show that genetic programming can work as a cost-effective intelligent modelling tool, enabling us to create prototype process models quickly and inexpensively instead of an engineer developing the process model.

Automation↗

Ecological bistability and evolutionary reversals under asymmetrical competition.

How does the process of life-history evolution interplay with population dynamics? Almost all models that have addressed this question assume that any combination of phenotypic traits uniquely determine the ecological population state. Here we show that if multiple ecological equilibria can exist, the evolution of a trait that relates to competitive performance can undergo adaptive reversals that drive cyclic alternation between population equilibria. The occurrence of evolutionary reversals requires neither environmentally driven changes in selective forces nor the coevolution of interactions with other species. The mechanism inducing evolutionary reversals is twofold. First, there exist phenotypes near which mutants can invade and yet fail to become fixed; although these mutants are eventually eliminated, their transitory growth causes the resident population to switch to an alternative ecological equilibrium. Second, asymmetrical competition causes the direction of selection to revert between high and low density. When ecological conditions for evolutionary reversals are not satisfied, the population evolves toward a steady state of either low or high abundance, depending on the degree of competitive asymmetry and environmental parameters. A sharp evolutionary transition between evolutionary stasis and evolutionary reversals and cycling can occur in response to a smooth change in ecological parameters, and this may have implications for our understanding of size-abundance patterns.

Animals↗

The origin of hematopoietic cell type diversity.

The hematopoietic system remains robust with regards to extrinsic perturbations, in sharp contrast with the stochastic behavior of hematopoeitic stem cells (HSCs) at the single cell level, suggesting that stability may be achieved within a stem cell system that undergoes constant self-renewal, commitment to differentiation and generates cell type diversification. Converging evidence at the interface of cellular, molecular and numerical studies suggests that diversity is generated by the chaotic dynamics of transcription factor networks within a cell and of the combination of growth factors and cytokines in the environment, both involving cooperation and competition. Current evidence indicates that HSCs are primed for multilineage gene expression. A subtle shift in transcription factor dosage is sufficient to perturb this equilibrium and to drive lineage commitment that involves a resolution of complexity at the molecular level and a transition towards less chaotic behavior. This dynamical instability establishes a state of responsiveness to extrinsic signals. Evolutionary conserved environmental cues that drive pattern formation or migratory behavior during embryonic development operate in the adult to influence the decision between self-renewal and differentiation in HSCs, as exemplified by the role of Notch1, Wnt proteins, BMPs and VEGF. In contrast, a network of cytokines uniquely present in mammalians influences later developmental stages, from progenitors with more restricted potentials (tri-, bi- or unipotent) to mature functional cells. These cytokines have co-opted the ancient Jak-STAT pathway but also appear to trigger lineage-affiliated transcription factors, thus linking environmental signaling to cell fate decisions.

Acute Disease↗

Diversity, biodiversity, conservation, and sustainability.

The concepts of diversity and biodiversity are analysed regarding their historical emergence, and their intrinsic meaning and differences are discussed. Through a brief synopsis, difficulties usually experienced by statisticians in capturing the dynamics of diversity are analysed and main problems identified. The shift from diversity to the more holistic biodiversity as a working concept is appraised in terms of the novelty involved. Through a number of examples, the way the two concepts capture natural cyclic changes is analysed, and their reciprocal and complementary relations are approached theoretically. The way diversity could develop from the stores of biodiversity as its active expression through selective and evolutionary processes is described. Through the use of a very simple dynamic model, the concepts of diversity and biodiversity are analysed in extremely opposite hypothetical scenarios. Comparisons with natural situations are made and the theoretical implications from the conservation point of view are discussed. These support the opinion that conservation undertaken in restricted and protected areas is not self-sustainable, needing permanent external intervention to regulate internal processes, and in the long run will most probably lead in the direction of obsolescence and extinction. Finally, the relations between diversity, biodiversity, and sustainability are approached. The vagueness of the sustainability concept is discussed. Preservation of biodiversity is then defended as one of the best available indicators to assist us in fixing boundaries which may help to provide a more precise definition of sustainability.

Animals↗

Dynamic and non-additive gene regulation shapes maize responses to simultaneous salt and cold stress.

Salt and cold stresses often occur together in nature and severely impact crop productivity, yet their transcriptional regulation remains poorly understood. Here, we conducted a time-series transcriptomic analysis of maize under salt, cold, and their combination at 0, 6, 12, and 24 h. Differential expression analysis revealed dynamic, condition-specific gene responses grouped into eight distinct temporal patterns. Promoter motif analysis of genes within each pattern identified 5-39 significantly enriched motifs, with over 40% lacking known counterparts, suggesting the involvement of previously uncharacterized cis-regulatory elements in stress-responsive transcriptional regulation. By comparing combined stress responses to the sum of single-stress effects, we found that about 74% of DEGs showed non-additive patterns, suggesting that combined stress triggers a distinct transcriptional program. Evolutionary analysis showed that additive DEGs tend to be more recently evolved, subject to weaker purifying selection, and enriched in transposed duplications, contrasting with the stronger constraint observed in non-additive DEGs. WGCNA identified 24 co-expression modules, among which 65 hub DEGs were detected in modules significantly correlated with specific stress conditions. Furthermore, we reconstructed 228, 20, and 200 sequential transcription factor cascades spanning 6 h, 12 h, and 24 h under cold, salt, and combined stress, respectively, with no cascade shared across all three conditions. Together, these results reveal that maize responses to combined salt and cold stress are largely non-additive and temporally dynamic, with distinct evolutionary patterns underlying different response types, offering insights and candidate regulators for enhancing crop stress resilience.

Zea mays↗

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

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

Animals↗

Evolutionary cheating in Escherichia coli stationary phase cultures.

Starved cultures of Escherichia coli are highly dynamic, undergoing frequent population shifts. The shifts result from the spread of mutants able to grow under conditions that impose growth arrest on the ancestral population. To analyze competitive interactions underlying this dynamic we measured the survival of a typical mutant and the wild type during such population shifts. Here we show that the survival advantage of the mutant at any given time during a takeover is inversely dependent on its frequency in the population, its growth adversely affects the survival of the wild type, and its ability to survive in stationary phase at fixation is lower than that of its ancestor. These mutants do not enter, or exit early, the nondividing stationary-phase state, cooperatively maintained by the wild type. Thus they end up overrepresented as compared to their initial frequency at the onset of the stationary phase, and subsequently they increase disproportionately their contribution in terms of progeny to the succeeding generation in the next growth cycle, which is a case of evolutionary cheating. If analyzed through the game theory framework, these results might be explained by the prisoner's dilemma type of conflict, which predicts that selfish defection is favored over cooperation.

Cell Division↗

From hawks and doves to self-consistent games of territorial behavior.

Explaining the "prior-residence effect" (automatic owner status of individuals who arrived first in an area) was one of the very first applications of game theory in animal behavior. These models, however, predict paradoxical solutions where intruders always win, with no satisfactory explanation for the absence of such cases in nature. We propose a solution based on new developments in evolutionary game theory. A self-consistent model with feedbacks between individual behavior and population dynamics produces qualitatively different frequency-dependent selection on intruders (floaters) than on territory owners. Starting with an ancestral population with no respect for ownership, the most likely evolutionary end point is complete or partial respect. Conventional rules of conflict resolution thus can rely on "uncorrelated asymmetries" without differences in resource-holding power or territory value, although they will be strengthened by such differences. We also review the empirical literature on animal contests, testing whether asymmetries in resource-holding power are required to explain the observations. Despite much empirical effort, results remain inconclusive, because experiments are often unable to distinguish between the motivation of individuals to fight and the behavioral outcome of a contest. To help arrive at conclusive answers, we suggest a standardized empirical approach to quantify prior-residence effects.

Animals↗

How should we define goodness?--reputation dynamics in indirect reciprocity.

Theory of indirect reciprocity is important in explaining cooperation between humans. Since a partner of a social interaction often changes, an individual should assess his partner by using social information such as reputation and make decisions whether to help him or not. To those who have 'good' social reputation does a player give aid as reciprocation, whereas he has to refuse to help those who have 'bad' reputation. Otherwise benefits of altruism is easily exploited by them. Little has been known, however, about the definition of 'goodness' in reputation. What kind of actions are and should be regarded as good and what kind of actions bad? And what sort of goodness enables sustaining exchange of altruism? We herein challenge this question with an evolutionary perspective. We generalize social reputation as 'Honor-score' (H-score) and examine the conditions under which individuals in a group stably maintain cooperative relationships based on indirect reciprocity. We examine the condition for evolutionarily stable strategies (ESSs) over 4096 possible cases exhaustively. Mathematical analysis reveals that only eight cases called 'leading eight' are crucial to the evolution of indirect reciprocity. Each in the leading eight shares two common characteristics: (i) cooperation with good persons is regarded as good while defection against them is regarded as bad, and (ii) defection against bad persons should be regarded as a good behavior because it works as sanction. Our results give one solution to the definition of goodness from an evolutionary viewpoint. In addition, we believe that the formalism of reputation dynamics gives general insights into the way social information is generated, handled, and transmitted in animal societies.

Altruism↗

Evolutionary reconstruction of networks.

Can a graph specifying the pattern of connections of a dynamical network be reconstructed from statistical properties of a signal generated by such a system? In this model study, we present a Metropolis algorithm for reconstruction of graphs from their Laplacian spectra. Through a stochastic process of mutations and selection, evolving test networks converge to a reference graph. Applying the method to several examples of random graphs, clustered graphs, and small-world networks, we show that the proposed stochastic evolution allows exact reconstruction of relatively small networks and yields good approximations in the case of large sizes.

Journal Article↗

Evolutionary stability of ecological hierarchy.

A self-similar hierarchical solution that is both dynamically and evolutionarily stable is found for the multidimensional Lotka-Volterra equation with a single chain of prey-predator relations. This gives a simple and natural explanation to the key features of hierarchical ecosystems, such as its ubiquity, pyramidal population distribution, and higher aggressiveness among higher trophic levels.

Animals↗