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Development and evolution of lateral line placodes in amphibians. - II. Evolutionary diversification.

The amphibian lateral line system develops from a series of lateral line placodes. The different phases of development from early induction, to pattern formation, differentiation, morphogenesis, and metamorphic fate were summarized in the first part of this review (Schlosser, 2002a). Here, a survey of the diversity of lateral line systems in amphibians is presented indicating that most phases of lateral line development have been subject to evolutionary changes. Several trends suggest important roles for both adaptive changes and internal constraints in amphibian lateral line evolution. Many of these trends involved the coordinated modification of different derivatives of lateral line placodes suggesting that these placodes are not only autonomous developmental modules, but also units of evolutionary variation that tend to be modified in a coherent and largely context-independent fashion.

Journal Article↗

Phenotypic plasticity, sexual selection and the evolution of colour patterns.

When a population comes to occupy a new environment, phenotypically plastic responses alter the distribution of phenotypes, and hence affect both the direction and the intensity of selection. Rates of evolution can be accelerated or retarded compared to what would happen in the absence of plasticity. Plastic responses in one trait result in novel selection pressures on other traits, and this can lead to evolution in completely different directions than predicted in the absence of plasticity. In this paper I use the concept of the adaptive surface in order to identify conditions under which the various different outcomes are expected. I then discuss differences between sexually and naturally selected traits. Sexually selected traits are often expected to be plastic in their expression, with individuals in high condition developing greater elaboration. As examples of sexually selected traits I review the evolution of colour patterns in birds with a view to assessing the magnitude of plastic responses in their development, and to ask how such responses may have influenced genetic evolution. The common colour pigments in birds are carotenoids and melanins. Both are used in social signaling, and consequently are expected to evolve to be phenotypically plastic indicators of an individual's quality. Perhaps partly because they are condition indicators, the quantity of carotenoids in the plumage can be strongly influenced by diet. Examples are described where alterations of carotenoids in the diet are thought to have altered the phenotype, driving genetic evolution in novel directions. Melanin patterns seem to be less affected by diet, but the intensity of melanization after moult is affected by social interactions during the moult and by raising birds in humid conditions. Hormonal manipulations can have dramatic effects on both the kinds of melanin produced (eumelanin or phaeomelanin) as well as the patterns they form. Differences between species in melanin patterns resemble differences produced by environmental manipulations, as well as those produced by simple modulations of parameters in computer simulations of pattern formation. While phenotypic plasticity is one way that genetic change in plumage patterns (and other traits) could be driven, there are others, including the appearance of major mutations and selection on standing variation whose distribution is not altered in the new environment. I consider some evidence for the different alternatives, and ask when they might lead to qualitatively different evolutionary outcomes.

Adaptation, Physiological↗

Resolving the paradox of common, harmful, heritable mental disorders: which evolutionary genetic models work best?

Given that natural selection is so powerful at optimizing complex adaptations, why does it seem unable to eliminate genes (susceptibility alleles) that predispose to common, harmful, heritable mental disorders, such as schizophrenia or bipolar disorder? We assess three leading explanations for this apparent paradox from evolutionary genetic theory: (1) ancestral neutrality (susceptibility alleles were not harmful among ancestors), (2) balancing selection (susceptibility alleles sometimes increased fitness), and (3) polygenic mutation-selection balance (mental disorders reflect the inevitable mutational load on the thousands of genes underlying human behavior). The first two explanations are commonly assumed in psychiatric genetics and Darwinian psychiatry, while mutation-selection has often been discounted. All three models can explain persistent genetic variance in some traits under some conditions, but the first two have serious problems in explaining human mental disorders. Ancestral neutrality fails to explain low mental disorder frequencies and requires implausibly small selection coefficients against mental disorders given the data on the reproductive costs and impairment of mental disorders. Balancing selection (including spatio-temporal variation in selection, heterozygote advantage, antagonistic pleiotropy, and frequency-dependent selection) tends to favor environmentally contingent adaptations (which would show no heritability) or high-frequency alleles (which psychiatric genetics would have already found). Only polygenic mutation-selection balance seems consistent with the data on mental disorder prevalence rates, fitness costs, the likely rarity of susceptibility alleles, and the increased risks of mental disorders with brain trauma, inbreeding, and paternal age. This evolutionary genetic framework for mental disorders has wide-ranging implications for psychology, psychiatry, behavior genetics, molecular genetics, and evolutionary approaches to studying human behavior.

Adaptation, Physiological↗

A family competition evolutionary algorithm for automated docking of flexible ligands to proteins.

In this paper, we study an evolutionary algorithm for flexible ligand docking. Based on family competition and adaptive rules, the proposed approach consists of global and local strategies by integrating decreasing mutations and self-adaptive mutations. To demonstrate the robustness of the proposed approach, we apply it to the problems of the first international contests on evolutionary optimization. Following the description of function optimization, our approach is applied to a dihydrofolate reductase enzyme with the anti-cancer drug methotrexate and with two analogs of the antibacterial drug trimethoprim. Our numerical results indicate that the proposed approach is robust. The docked lowest energy structures have rms derivations ranging from 0.72 A to 1.98 A with respect to the corresponding crystal structure.

Algorithms↗

Evolutionary studies on an alpha-amylase gene segment in bats and other mammals.

Comparative studies of salivary glands showed that they maybe related to the adaptive radiation of bats, especially in the family Phylostomidae. In this study we have been searching for a likely relationship between different feeding habits found in bats and possible adaptive changes in a coding segment of the alpha-amylase enzyme. We have also tested some hypothesis about the phylogenetic relationship of bats and other mammals. A 663 bp segment of the alpha-amylase gene, corresponding to the exon 4 and part of the intron c, was sequenced in nine bat species. The exon 4 was also sequenced in further ten mammalian species. The phylogenetic trees generated with different methods produced the same results. When the intron c and the exon 4 were independently analyzed, they showed distinct topologies involving the bat species Sturnira lilium, different from the traditional bat phylogeny. Phylogenetic analysis of bats, primates and rodents supports the Euarchontoglires-Laurasiatheria hypothesis about the relationship among these groups. Selection tests showed that the alpha-amylase exon 4 is under strong purifying selection, probably caused by functional constraints. The conflicting bat phylogenies could not be explained by evolutionary convergence due to adaptive forces, and the different topologies may be likely due to the retention of plesiomorphic characters or the independent acquisition by evolutionary parallelism.

Amino Acid Sequence↗

Population and landscape genomics provide insights into the adaptive genetic variation and future climate-induced vulnerability of the endangered tree species Phoebe bournei.

Elucidating the genomic underpinnings of adaptive variation is highly important for the conservation, landscape application, and management of ornamental trees against the backdrop of global climate change. However, research on the genetic mechanisms underlying climate adaptation in Phoebe bournei-a near-threatened subtropical tree species endemic to China, which is endowed with exceptionally high ornamental and ecological value-remains scarce. Whole-genome resequencing was conducted on 362 individuals from 27 natural populations across the geographical range of the species. Genome-environment association analyses were employed to identify 1556 climate-associated variants and 167 candidate genes associated with temperature and precipitation variables. Through functional annotation and expression profiling, pivotal genes, including TRX-M4 and FBD1, were identified as integral to drought and heat stress responses, with adaptive alleles displaying distinct geographic frequency distributions and significant phenotypic differentiation. Divergent evolutionary trajectories were deduced among populations, with southeastern populations distinguished by elevated genetic diversity and strong signatures of local adaptation. Nevertheless, projections derived from the Risk of Non-Adaptedness and gradient forest models suggest that these southeastern populations will face substantial genomic offset under future climate scenarios, signaling heightened vulnerability and the need for prioritized conservation and management. This study provides the first genome-wide perspective into the adaptive evolution of P. bournei and offers a robust foundation for its conservation and climate-resilient management.

Journal Article↗

Host genetics and viral infections: immunology taught by viruses, virology taught by the immune system.

Viruses are prototypic obligate intracellular pathogens, and are therefore, by necessity, highly pre-adapted to surviving the host immune response. Nevertheless, host genetic factors remain an important determinant of disease outcome, particularly in the case of viruses that have encountered humans in the more recent past (e.g. human immunodeficiency virus). Recent studies have identified an increasingly rich network of functionally relevant polymorphic immune factors, including major MHC alleles, killer immunoglobulin-like receptors and functional chemokine receptor polymorphisms. Moreover, genetic variation is increasingly appreciated beyond the single genotype level, incorporating extended haplotypes as well as regions of segmental genetic duplication. These issues can be considered within an evolutionary perspective that acknowledges the crucial role of adaptive host-viral relationships in shaping both host and pathogen genetic diversity.

HIV Infections↗

Patterns and mechanisms of genetic and phenotypic differentiation in marine microbes.

Microbes in the ocean dominate biogeochemical processes and are far more diverse than anticipated. Thus, in order to understand the ocean system, we need to delineate microbial populations with predictable ecological functions. Recent observations suggest that ocean communities comprise diverse groups of bacteria organized into genotypic (and phenotypic) clusters of closely related organisms. Although such patterns are similar to metazoan communities, the underlying mechanisms for microbial communities may differ substantially. Indeed, the potential among ocean microbes for vast population sizes, extensive migration and both homologous and illegitimate genetic recombinations, which are uncoupled from reproduction, challenges classical population models primarily developed for sexually reproducing animals. We examine possible mechanisms leading to the formation of genotypic clusters and consider alternative population genetic models for differentiation at individual loci as well as gene content at the level of whole genomes. We further suggest that ocean bacteria follow at least two different adaptive strategies, which constrain rates and bounds of evolutionary processes: the 'opportunitroph', exploiting spatially and temporally variable resources; and the passive oligotroph, efficiently using low nutrient concentrations. These ecological lifestyle differences may represent a fundamental divide with major consequences for growth and predation rates, genome evolution and population diversity, as emergent properties driving the division of labour within microbial communities.

Adaptation, Biological↗

Negative effect of genetic bottlenecks on the adaptability of vesicular stomatitis virus.

Muller's ratchet is a principle of evolutionary genetics describing mutant accumulation in populations that are repeatedly subjected to genetic bottleneck. The immediate effect of Muller's ratchet, overall loss of fitness, has been confirmed in several viral systems belonging to different groups. This report shows that in addition to fitness loss, genetic bottlenecks also have longer-term effects, namely changes in the capacity of viral populations to adapt. Thus, vesicular stomatitis virus strains with a history of genetic bottleneck have lower adaptability than strains maintained at relatively large population sizes. This lower adaptability is illustrated by their reduced ability to regain fitness and by their inability to outcompete wild-type populations in situations where the initial fitness of the bottlenecked mutant is the same or even higher than the initial fitness of the wild-type.

Animals↗

Adaptedness, evolution and a hierarchical concept of fitness.

Fitness is a consequence of the adaptedness of an entity to its environment. The fitness of an evolutionary entity, when the entity is defined by a set of attributes determining how it interacts with its environment, is manifested as persistence of those attributes. Two measures of fitness are presented to explicate this concept: (1) an extensive measure of the persistence of the original set of attributes in the individual entity and its descendants, but corrected for evolution; this includes as a special case fitness as used in population genetics; and, (2) an intensive measure that is independent of the abundance of descendant exemplars. Fitness as conceived here is a function of survival time, degree of evolution, and when applicable, reproductive contribution. The rate of fitness increase of an entity will vary inversely with the degree of evolutionary change experienced by the entity, its descendants, or both through time. Adaptive evolution can increase the length of time that fitness accumulates by increasing the survival of descendants possessing all or some of the ancestral attributes. Reproduction, where possible, can increase the number of descendants. This concept of fitness is simple, unambiguous, coherent, and applicable to entities at any hierarchical level of interest to ecologists and evolutionary biologists.

Adaptation, Physiological↗

Dietary restriction and aging: comparative tests of evolutionary hypotheses.

Dietary restriction (DR) increases life span in many types of animals. The response to chronic DR may be an adaptation to environments with variable food levels. This study uses the comparative method to test evolutionary predictions about the origin of the response to DR, using data from 10 species of rotifers. Most species, but not all, responded to DR by increasing mean life span, maximum life span, reproductive life span, mortality rate doubling time, and initial mortality rate. Interspecific comparisons did not show the predicted correlations between the strength of the response to DR and either reproductive life span, age of first reproduction, or total reproduction. There was support for the idea that the response to chronic DR is associated with changes in reproductive allocation during short-term periods of starvation: species that reduced reproduction when starved increased their life spans under DR, whereas species that continued to reproduce when starved decreased their life spans under DR.

Aging↗

Widespread geographical distribution of mitochondrial haplotypes in rock-dwelling cichlid fishes from Lake Tanganyika.

The spectacularly diverse cichlid fish species flocks of the East African Rift Lakes have elicited much debate on the potential evolutionary mechanisms responsible for the origin of these adaptive radiations. An historical perspective on population structure may offer insights into the processes driving population differentiation and possibly speciation. Here, we examine mitochondrial DNA (mtDNA) sequence variation in two endemic species of rock-dwelling cichlids, Simochromis babaulti and S. diagramma, from Lake Tanganyika. Phylogeographic analyses were used to infer what factors might have been important in the genetic structuring of Simochromis populations. Patterns of mtDNA differentiation in Simochromis were compared to those of other rock-dwelling cichlids to distinguish between competing hypotheses concerning the processes underlying their evolution. In striking contrast to previous findings, populations of Simochromis, even those separated by up to 300 km, were found to share mitochondrial DNA haplotypes. There is no correspondence between mtDNA genealogies and the geographical distribution of populations. Only S. babaulti, but not S. diagramma was found to have a significant association between genetic and geographic distance. These phylogeographic patterns suggest that the evolutionary effects of abiotic and biotic factors shaping population genetic structure may differ substantially even among closely related species of rock-dwelling cichlids. Physical events and barriers to gene flow that are believed to have had a major impact on the geographical distribution and intralacustrine speciation of Tropheus do not seem to have equally strongly affected its close relative Simochromis. These findings emphasize that no single mechanism can be responsible for the formation of population structure, speciation, and the adaptive radiation of all cichlid fishes.

Africa, Eastern↗

Online evolution for a self-adapting robotic navigation system using evolvable hardware.

Great interest has been shown in the application of the principles of artificial life to physically embedded systems such as mobile robots, computer networks, home devices able continuously and autonomously to adapt their behavior to changes of the environments. At the same time researchers have been working on the development of evolvable hardware, and new integrated circuits that are able to adapt their hardware autonomously and in real time in a changing environment. This article describes the navigation task for a real mobile robot and its implementation on evolvable hardware. The robot must track a colored ball, while avoiding obstacles in an environment that is unknown and dynamic. Although a model-free evolution method is not feasible for real-world applications due to the sheer number of possible interactions with the environment, we show that a model-based evolution can reduce these interactions by two orders of magnitude, even when some of the robot's sensors are blinded, thus allowing us to apply evolutionary processes online to obtain a self-adaptive tracking system in the real world, when the implementation is accelerated by the utilization of evolvable hardware.

Biological Evolution↗

Origin, world-wide dispersion, bio-geographical diversification, radiation and recombination: an evolutionary history of Yam mild mosaic virus (YMMV).

We developed an evolutionary epidemiological approach to understand the regional and world-wide dispersion of Yam mild mosaic virus (YMMV) by retracing its evolutionary history. Analyses of the distribution and the prevalence of YMMV in the Caribbean islands of Guadeloupe and Martinique, and in French Guyana revealed that YMMV has a wide repartition and different prevalence on Dioscorea alata L. (Asian and Oceanic origin), on D. cayenensis Lam.-D. rotundata Poir. (African origin) and on D. trifida L. (Amazon and the Caribbean origin) in this region. Considering the data on the current dispersion of the virus and the evolution and the history of the yams, the phylogenetic analysis of the 3' terminal part of the YMMV genome gave a consistent support of the Asian-Pacific origin of YMMV from D. alata species. The YMMV phylogenetic tree is star-like, suggesting an early split of the genetic lineages. An important part of the clades is constituted by a single lineage arisen by recombination. The largest emerging monophyletic group illustrates well YMMV geographical dispersion. This evolutionary pattern contrasts with the one revealed by the African distinct lineages and by the second significant monophyletic group, for which a host adaptation to D. trifida is suggested. The analysis of the pattern of nucleotide substitutions in the CP gene revealed that purifying selection dominates the evolution of the CP of potyviruses and strongly operates on the YMMV. Switching events, radiation, host and geographical adaptation and recombination events are proposed as major traits of YMMV evolutionary history.

3' Flanking Region↗

Perspective: From mutants to mechanisms? Assessing the candidate gene paradigm in evolutionary biology.

The generation of mutants in model organisms by geneticists and developmental biologists over the last century has occasionally produced phenotypes that are startlingly reminiscent of those seen in other species. Such extreme mutations have generally been dismissed by evolutionary geneticists since the "modern synthesis" as irrelevant to adaptation and speciation. But only in recent years has information on the molecular bases of mutant phenotypes become widely available, and thus work on testing the relevance of such extreme mutations to the generation of phylogenetic diversity has just begun. Here we evaluate whether evolutionary mimics are, in fact, useful for pinpointing the genetic differences that distinguish morphological variants generated during evolution. Examples come from both plants and animals, and range from intraspecific to interordinal taxonomic ranges. The use of mutationally defined candidate genes to predict evolutionary mechanisms has so far been most fruitful in explaining intraspecific variants, where it has been effective in both plants and animals. In several cases these efforts were facilitated or supported by parallel results from quantitative trait loci studies, in which natural alleles controlling continuous variation in developmental model organisms were mapped to mutationally defined genes. However, despite these successes the approach's utility seems to rapidly decay as a function of phylogenetic distance. This suggests that the divergence of developmental genetic systems is great even in closely related organisms and may become intractable at larger distances. We discuss this result in the context of what it teaches us about development, the future prospects of the candidate gene approach, and the historical debate over process in micro- and macroevolution.

Animals↗

Effects of ecological differentiation on Lotka-Volterra systems for species with behavioral adaptation and variable growth rates.

We study the properties of a n2-dimensional Lotka-Volterra system describing competing species that include behaviorally adaptive abilities. We indicate as behavioral adaptation a mechanism, based on a kind of learning, which is not viewed in the evolutionary sense but is intended to occur over shorter time scales. We consider a competitive adaptive n species Lotka-Volterra system, n > or = 3, in which one species is made ecologically differentiated with respect to the others by carrying capacity and intrinsic growth rate. The symmetry properties of the system and the existence of a certain class of invariant subspaces allow the introduction of a 7-dimensional reduced model, where n appears as a parameter, which gives full account of existence and stability of equilibria in the complete system. The reduced model is effective also in describing the time-dependent regimes for a large range of parameter values. The case in which one species has a strong ecological advantage (i.e. with a carrying capacity higher than the others), but with a varying growth rate, has been analyzed in detail, and time-dependent behaviors have been investigated in the case of adaptive competition among four species. Relevant questions, as species survival/exclusion, are addressed focusing on the role of adaptation. Interesting forms of species coexistence are found (i.e. competitive stable equilibria, periodic oscillations, strange attractors).

Adaptation, Biological↗

Disaptation and recovery in the evolution of Antarctic fishes.

The radiation of notothenioid fishes provides an excellent system to explore issues of evolution and adaptation. Most studies emphasize adaptation to the extreme Antarctic environment; however, recent work provides cogent examples of disaptation or evolutionary loss of function. The nature and extent of regressive change is revealed by subsequent adaptive recovery. Ancestral notothenioids were benthic but some became secondarily pelagic through the retention of larval characters. Paedomorphosis has produced detrimental changes in lateral-line sensory systems that have been made good by compensatory adaptation. In the icefish family, compensatory adaptation has followed the loss of the oxygen-binding pigments haemoglobin and myoglobin.

Journal Article↗

On the origin and evolution of major morphological characters.

Although the mathematical principles underpinning population-level evolution are now well studied, the origin and evolution of morphological novelties has received far less attention. Here, a broad but general theory for how this sort of change takes place is outlined, relying on functional continuity, least-constrained components of morphology, redundancy and preadaptation. At least four distinct sorts of redundancy are identified: (i) redundancy arising through duplication (amplification); (ii) redundancy arising through regionalisation (parcellation); (iii) redundancy arising through functional convergence; and (iv) redundancy arising from shared function (functional degeneracy). Although organisms are here recognised to be functionally constrained ("burdened", in Riedl's terminology), these constraints can be overcome through the combination of the four principles given above. Contrary to its common treatment, functional constraint is neither an ever-increasing restriction on the scope of evolution, nor does it require drastic events to overcome or "break" it. Rather, it is an evolutionary quantity, subject to selection at some level. The rules that govern morphological evolution are the primary controls on what is allowed to happen in the evolution of the overall genotype-phenotype system, suggesting strong controls on the sorts of developmental changes that might be associated with macroevolution. This model, then, sees organism functionality as the primary control on evolvability, with exact genetic make-up being of secondary importance. It should prove possible to recast traditional notions of body-plan evolution into the formulations of complex system analysis, which in the future may prove a unifying discipline for fields as disparate as palaeontology and gene regulatory networks. In particular, understanding how morphology can evolve may provide the critical link between the ecological and morphological networks that are currently the intense focus of evolutionary investigations.

Adaptation, Physiological↗