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Evolutionary consequences of coevolving targets

Most evolutionary optimization models incorporate a fitness evaluation that is based on a predefined static set of test cases or problems. In the natural evolutionary process, selection is of course not based on a static fitness evaluation. Organisms do not have to combat every existing disease during their lifespan; organisms of one species may live in different or changing environments; different species coevolve. This leads to the question of how information is integrated over many generations. This study focuses on the effects of different fitness evaluation schemes on the types of genotypes and phenotypes that evolve. The evolutionary target is a simple numerical function. The genetic representation is in the form of a program (i.e., a functional representation, as in genetic programming). Many different programs can code for the same numerical function. In other words, there is a many-to-one mapping between "genotypes" (the programs) and "phenotypes". We compare fitness evaluation based on a large static set of problems and fitness evaluation based on small coevolving sets of problems. In the latter model very little information is presented to the evolving programs regarding the evolutionary target per evolutionary time step. In other words, the fitness evaluation is very sparse. Nevertheless the model produces correct solutions to the complete evolutionary target in about half of the simulations. The complete evaluation model, on the other hand, does not find correct solutions to the target in any of the simulations. More important, we find that sparse evaluated programs are better generalizable compared to the complete evaluated programs when they are evaluated on a much denser set of problems. In addition, the two evaluation schemes lead to programs that differ with respect to mutational stability; sparse evaluated programs are less stable than complete evaluated programs.

Journal Article↗

Popular evolutionary psychology in the UK: an unusual case of science in the media?

This paper presents findings from quantitative analyses of UK press and print media coverage of evolutionary psychology during the 1990s. It argues that evolutionary psychology presents an interesting case for studies of science in the media in several different ways. First, press coverage of evolutionary psychology was found to be closely linked with the publications of popular books on the subject. Secondly, when compared to coverage of other subjects, a higher proportion of academics and authors wrote about evolutionary psychology in the press, contributing to the development of a scientific controversy in the public domain. Finally, it was found that evolutionary psychology coverage appeared in different areas of the daily press, and was rarely written about by specialist science journalists. The possible reason for these features are then explored, including the boom in popular science publishing during the 1990s, evolutionary psychology's status as a new subject of study and discussion, and the nature of the subject its as theoretically based and with a human, "everyday" subject matter.

Behavior↗

Apparent dependence of protein evolutionary rate on number of interactions is linked to biases in protein-protein interactions data sets.

BACKGROUND: Several studies have suggested that proteins that interact with more partners evolve more slowly. The strength and validity of this association has been called into question. Here we investigate how biases in high-throughput protein-protein interaction studies could lead to a spurious correlation. RESULTS: We examined the correlation between evolutionary rate and the number of protein-protein interactions for sets of interactions determined by seven different high-throughput methods in Saccharomyces cerevisiae. Some methods have been shown to be biased towards counting more interactions for abundant proteins, a fact that could be important since abundant proteins are known to evolve more slowly. We show that the apparent tendency for interactive proteins to evolve more slowly varies directly with the bias towards counting more interactions for abundant proteins. Interactions studies with no bias show no correlation between evolutionary rate and the number of interactions, and the one study biased towards counting fewer interactions for abundant proteins actually suggests that interactive proteins evolve more rapidly. In all cases, controlling for protein abundance significantly decreases the observed correlation between interactions and evolutionary rate. Finally, we disprove the hypothesis that small data set size accounts for the failure of some interactions studies to show a correlation between evolutionary rate and the number of interactions. CONCLUSIONS: The only correlation supported by a careful analysis of the data is between evolutionary rate and protein abundance. The reported correlation between evolutionary rate and protein-protein interactions cannot be separated from the biases of some protein-protein interactions studies to count more interactions for abundant proteins.

Bacterial Proteins↗

Toward a neutral evolutionary model of gene expression.

We introduce a stochastic model that describes neutral changes of gene expression over evolutionary time as a compound Poisson process where evolutionary events cause changes of expression level according to a given probability distribution. The model produces simple estimators for model parameters and allows discrimination between symmetric and asymmetric distributions of evolutionary expression changes along an evolutionary lineage. Furthermore, we introduce two measures, the skewness of expression difference distributions and relative difference of evolutionary branch lengths, which are used to quantify deviation from clock-like behavior of gene expression distances. Model-based analyses of gene expression profiles in primate liver and brain samples yield the following results: (1) The majority of gene expression changes are consistent with a neutral model of evolution; (2) along evolutionary lineages, upward changes in expression are less frequent but of greater average magnitude than downward changes; and (3) the skewness measure and the relative branch length difference confirm that an acceleration of gene expression evolution occurred on the human lineage in brain but not in liver. We discuss the latter result with respect to a neutral model of transcriptome evolution and show that a small number of genes expressed in brain can account for the observed data.

Animals↗

[Evolutionary chromosomes and sex dimorphism].

In order to understand the phenomenon of sex, it is not enough to know about its reproductive role and role in recombination. In addition, knowledge of its evolutionary role is required. Sexual differentiation allows us to test evolutionary innovations in the male genome before they are transmitted to the female genome. This is possible in the framework of dichronous evolution, when the evolution of the male sex goes faster than the evolution of the female sex. This implies that there clearly should be male genes, which are already present in the male genome, but are not yet in the female genome; and vice versa, there should exist female genes, which have already been lost by the male sex, but still remain in females. Dichronicity results from different reaction norm of the sexes. This interpretation implies (1) the nomadic behavior of genes in the genome (2) the existence of evolutionary sexual dimorphism, which begins in the form of modificational changes in the female sex followed by selection-derived changes in the male sex; (3) the discovery of evolutionary roles of chromosomes and the proposition of a new concept, according to which the Y-chromosome represents a "gateway" for environment-related information coming to the genome, i.e., place of "birth" and testing of new genes, which comprise the basis of evolutionary sexual dimorphism--its initiator, accelerator, and regulator. The X-chromosome is the "transport" chromosome that transfers new genes from the Y-chromosomes into autosomes and therefore serves as a stabilizer, relaxer, and liquidator of evolutionary sexual dimorphism; it also serves as repository of the outgoing genes that await elimination. This concept provides for a new interpretation of the appearance, localization, and movement of genes along chromosomes and between them, and it sheds new light on the phenomena of chromosomal inactivation, mobile genes, association of Y-chromosome with stress, viruses, etc.

Animals↗

[Evolutionary physiology and related sciences].

A brief historical review of development of the evolutionary physiology as an independent discipline. Description of the problems and methodology of the evolutionary physiology in the historical perspective. Interrelation of the evolutionary physiology and the related physiological and biological disciplines, specific features of the evolutionary physiology as an independent research discipline. The proceedings of 10 workshops on the evolutionary physiology, and the national and international bibliographic publications on evolution of the organic world are used to demonstrate the structure and the changes in time of the thematic spectrum explored by the evolutionary physiology.

Biological Evolution↗

Myosin superfamily evolutionary history.

The superfamily of myosin proteins found in eukaryotic cells is known to contain at least 18 different classes. Members are classified based on the phylogenetic analysis of the head domains located at the amino terminus of the polypeptide. While phylogenetic relationships provide insights into the functional relatedness of myosins within and between families, the evolutionary history of the myosin superfamily is not revealed by such studies. In order to establish the evolutionary history of the superfamily, we analyzed the representation of myosin gene families in a range of organisms covering the taxonomic spectrum. The amino acid sequences of 232 myosin heavy chains, as well as 65 organisms representing the protist, plant, and animal kingdoms, were included in this study. A phylogenetic tree of organisms was constructed based on several complementary taxonomic classification schemes. The results of the analysis support an evolutionary hypothesis in which myosins II and I evolved the earliest of all the myosin groups. Myosins V and XI evolved from a common myosin II-like ancestor, but the two families diverged to either the plant (XI) or animal (V) lineage. Class VII myosin appeared fourth among the families, and classes VI and IX appeared later during the early period of metazoan radiation. Myosins III, XV, and XVIII appeared after this group, and X appeared during the formative phases of vertebrate evolution. The remaining members of the myosin superfamily (IV, VI, XII, XIII, XIV, XVI, and XVII) are limited in distribution to one or more groups of organisms. The evolutionary data permits one to predict the likelihood that myosin genes absent from a given species are either missing (not found yet because of insufficient data) or lost due to a mutation that removed the gene from an organism's lineage. In conclusion, an analysis of the evolutionary history of the myosin superfamily suggests that early-appearing myosin families function as generalists, carrying out a number of functions in a variety of cell types, while more recently evolved myosin families function as specialists and are limited to a few organisms or a few cell types within organisms.

Biological Evolution↗

From the "Modern Synthesis" to cybernetics: Ivan Ivanovich Schmalhausen (1884-1963) and his research program for a synthesis of evolutionary and developmental biology.

Ivan I. Schmalhausen was one of the central figures in the Russian development of the "Modern Synthesis" in evolutionary biology. He is widely cited internationally even today. Schmalhausen developed the main principles of his theory facing the danger of death in the totalitarian Soviet Union. His great services to evolutionary and theoretical biology are indisputable. However, the received view of Schmalhausen's contributions to evolutionary biology makes an unbiased reading of his texts difficult. Here we show that taking all of his works into consideration (including those only available in Russian) paints a much more dynamic and exciting picture of what he tried to achieve. Schmalhausen pioneered the integration of a developmental perspective into evolutionary thinking. A main tool for achieving this was his approach to living objects as complex multi-level self-regulating systems. Schmalhausen put enormous effort into bringing this idea into fruition during the final stages of his career by combining evolutionary theory with cybernetics. His results and ideas remain thought-provoking, and his texts are of more than just historical interest.

Biological Evolution↗

The ConSurf-HSSP database: the mapping of evolutionary conservation among homologs onto PDB structures.

The HSSP (Homology-Derived Secondary Structure of Proteins) database provides multiple sequence alignments (MSAs) for proteins of known three-dimensional (3D) structure in the Protein Data Bank (PDB). The database also contains an estimate of the degree of evolutionary conservation at each amino acid position. This estimate, which is based on the relative entropy, correlates with the functional importance of the position; evolutionarily conserved positions (i.e., positions with limited variability and low entropy) are occasionally important to maintain the 3D structure and biological function(s) of the protein. We recently developed the Rate4Site algorithm for scoring amino acid conservation based on their calculated evolutionary rate. This algorithm takes into account the phylogenetic relationships between the homologs and the stochastic nature of the evolutionary process. Here we present the ConSurf-HSSP database of Rate4Site estimates of the evolutionary rates of the amino acid positions, calculated using HSSP's MSAs. The database provides precalculated evolutionary rates for nearly all of the PDB. These rates are projected, using a color code, onto the protein structure, and can be viewed online using the ConSurf server interface. To exemplify the database, we analyzed in detail the conservation pattern obtained for pyruvate kinase and compared the results with those observed using the relative entropy scores of the HSSP database. It is reassuring to know that the main functional region of the enzyme is detectable using both conservation scores. Interestingly, the ConSurf-HSSP calculations mapped additional functionally important regions, which are moderately conserved and were overlooked by the original HSSP estimate. The ConSurf-HSSP database is available online (http://consurf-hssp.tau.ac.il).

Algorithms↗

Human evolutionary psychology and animal behaviour.

Homo sapiens is increasingly being studied within the evolutionary (adaptationist, selectionist) framework favoured by animal behaviour researchers. There are various labels for such work, including evolutionary psychology, human behavioural ecology and human sociobiology. Collectively, we call these areas 'human evolutionary psychology' (HEP) because their shared objective is an evolutionary understanding of human information processing and decision making. Sexual selection and sex differences have been especially prominent in recent HEP research, but many other topics have been addressed, including parent-offspring relations, reciprocity and exploitation, foraging strategies and spatial cognition. Many HEP researchers began their scientific careers in animal behaviour, and in many ways, HEP research is scarcely distinguishable from other animal behaviour research. Currently controversial issues in HEP, such as the explanation(s) for observed levels of heritable diversity, the kinds of data needed to test adaptationist hypotheses, and the characterization of a species-typical 'environment of evolutionary adaptedness', are issues in animal behaviour as well. What gives HEP a distinct methodological flavour is that the research animal can talk, an ability that has both advantages and pitfalls for researchers. The proper use of self-reports and other verbal data in HEP might usefully become a subject of future research in its own right. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Escape from evolutionary stasis by transposon-mediated deleterious mutations.

Evolution within a rugged fitness landscape is limited by the tendency for organisms to become trapped on local optima resulting in evolutionary stasis. It is presently unclear how founder populations escape from an adaptive peak to found a new species. Insertion sequences, transposons and other mobile DNA elements are found in all species of eukaryotes, bacteria and archaebacteria, where they have been sought and are usually considered to be genomic parasites or selfish genes. However, many transposons and other mobile repetitive DNA are remarkably species or phyla-specific, indicating that infection with transposable elements coincides with speciation events and is involved in promoting evolutionary change. We propose here a model in which transposable elements are involved in speciation events by their ability to produce irreversible deleterious mutations that promote escape from evolutionary stasis. We have constructed a genetic algorithm designed to model both spontaneous and transposon-mediated mutations in populations of asexual digital organisms. We use this model to investigate the effect of transposon-mediated mutations on the rate of evolution of digital organisms as they compete for resources within an artificial adaptive landscape. In the absence of transposon mutations the seed organisms quickly evolve to occupy the nearest adaptive peak but thereafter evolutionary stasis ensues and adjacent empty peaks are left unoccupied. In the presence of transposon mutations, evolution is again dominated by stasis but is punctuated by bursts of rapid evolution in with consecutive unoccupied adaptive peaks are filled with organisms derived from single transposition events. Rapid evolutionary events leading to founding of new biological species, may be similarly initiated by irreversible deleterious mutations induced by transposition.

Adaptation, Physiological↗

Evolutionary dynamics of seed size and seedling competitive ability.

We present a model for the evolutionary dynamics of seed size when there is a trade-off between seed size and seed number, and seedlings from large seeds are better competitors and have a higher precompetitive survival than seedlings from small seeds. We find that strong competitive asymmetry, high resource levels, and intermediate harshness of the precompetitive environment favor coexistence of plants with different seed sizes. If the evolution of seed size is mutation-limited and single mutations have only a small phenotypic effect, then an initially monomorphic population reaches the final evolutionarily stable polymorphic state through one or more discrete evolutionary branching events. At each such branching event, a given lineage already present in the population divides into two phenotypically diverging daughter lines, each with its own seed size. If the precompetitive survival of seeds and seedlings is high for small and large seeds alike, however, evolutionary branching may be followed by the extinction of one or more lineages. Various results presented here are model-independent and point the way to a more general evolutionary bifurcation theory describing how the number and stability properties of evolutionary equilibria may change as a consequence of changes in model parameters.

Biological Evolution↗

Evolutionary engineering of industrially important microbial phenotypes.

The tremendous complexity of dynamic interactions in cellular systems often impedes practical applications of metabolic engineering that are largely based on available molecular or functional knowledge. In contrast, evolutionary engineering follows nature's 'engineering' principle by variation and selection. Thus, it is a complementary strategy that offers compelling scientific and applied advantages for strain development and process optimization, provided a desired phenotype is amenable to direct or indirect selection. In addition to simple empirical strain development by random mutation and direct selection on plates, evolutionary engineering also encompasses recombination and continuous evolution of large populations over many generations. Two distinct evolutionary engineering applications are likely to gain more relevance in the future: first, as an integral component in metabolic engineering of strains with improved phenotypes, and second, to elucidate the molecular basis of desired phenotypes for subsequent transfer to other hosts. The latter will profit from the broader availability of recently developed methodologies for global response analysis at the genetic and metabolic level. These methodologies facilitate identification of the molecular basis of evolved phenotypes. It is anticipated that, together with novel analytical techniques, bioinformatics, and computer modeling of cellular functions and activities, evolutionary engineering is likely to find its place in the metabolic engineer's toolbox for research and strain development. This review presents evolutionary engineering of whole cells as an emerging methodology that draws on the latest advances from a wide range of scientific and technical disciplines.

Biological Evolution↗

A protein alignment scoring system sensitive at all evolutionary distances.

Protein sequence alignments generally are constructed with the aid of a "substitution matrix" that specifies a score for aligning each pair of amino acids. Assuming a simple random protein model, it can be shown that any such matrix, when used for evaluating variable-length local alignments, is implicitly a "log-odds" matrix, with a specific probability distribution for amino acid pairs to which it is uniquely tailored. Given a model of protein evolution from which such distributions may be derived, a substitution matrix adapted to detecting relationships at any chosen evolutionary distance can be constructed. Because in a database search it generally is not known a priori what evolutionary distances will characterize the similarities found, it is necessary to employ an appropriate range of matrices in order not to overlook potential homologies. This paper formalizes this concept by defining a scoring system that is sensitive at all detectable evolutionary distances. The statistical behavior of this scoring system is analyzed, and it is shown that for a typical protein database search, estimating the originally unknown evolutionary distance appropriate to each alignment costs slightly over two bits of information, or somewhat less than a factor of five in statistical significance. A much greater cost may be incurred, however, if only a single substitution matrix, corresponding to the wrong evolutionary distance, is employed.

Algorithms↗

Strain evolution in Caenorhabditis elegans: transposable elements as markers of interstrain evolutionary history.

Evolutionary relationships across taxa can be deduced from sequence divergence of proteins, RNA, or DNA; sequences which diverge rapidly, such as those of mitochondrial genes, have been especially useful for comparisons of closely related species, and--within limits--of strains within a species. We have utilized the transposable element Tc1 as a polymorphic marker to evaluate the evolutionary relationships among nine Caenorhabditis elegans strains. For five low-Tc1-copy strains, we compared patterns of restriction fragments hybridizing to a cloned Tc1 probe. Twenty of the 40 Tc1 insertion sites thus characterized were common to all five strains, and so presumably preceded strain divergence; the 20 differential bands were used to construct a maximum-parsimony tree relating these strains. In four high-copy-number stocks (three wild-type strains and a subline), we determined occupancy of 35 individual Tc1 insertion sites by a polymerase chain reaction assay. Surprisingly, the high-copy strains share a common subset of these Tc1 insertions, and the chromosomal distribution of conserved Tc1 sites is "clustered" with respect to the other elements tested. These data imply a close evolutionary relationship among the high-copy strains, such that two of these strains appear to have been derived from the highest-copy-number lineage (represented by two stocks) through crossing with a low-Tc1 strain. Abundances of Tc1 elements were also estimated for the four high-copy-number stocks, at approximately 200-500 copies per haploid genome, by quantitative dot-blot hybridization relative to two low-copy strains. Annealing with 32P-labeled probes corresponding to full-length Tc1, an oligonucleotide within the Tc1 terminal inverted repeats, and an internal Tc1 oligonucleotide, gave essentially identical results--indicating that Tc1 termini exist in the genome primarily as components of full-length Tc1 elements. A composite evolutionary tree is proposed, based on the locations and numbers of Tc1 elements in these strains, which is consistent with a four-branch intraspecific tree deduced previously by maximum-parsimony analyses of mitochondrial sequence changes; it also serves to elucidate the evolutionary history of transposon mobility.

Animals↗

A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences.

Some simple formulae were obtained which enable us to estimate evolutionary distances in terms of the number of nucleotide substitutions (and, also, the evolutionary rates when the divergence times are known). In comparing a pair of nucleotide sequences, we distinguish two types of differences; if homologous sites are occupied by different nucleotide bases but both are purines or both pyrimidines, the difference is called type I (or "transition" type), while, if one of the two is a purine and the other is a pyrimidine, the difference is called type II (or "transversion" type). Letting P and Q be respectively the fractions of nucleotide sites showing type I and type II differences between two sequences compared, then the evolutionary distance per site is K = -(1/2) ln [(1-2P-Q) square root of 1-2Q]. The evolutionary rate per year is then given by k = K/(2T), where T is the time since the divergence of the two sequences. If only the third codon positions are compared, the synonymous component of the evolutionary base substitutions per site is estimated by K'S = -(1/2) ln (1-2P-Q). Also, formulae for standard errors were obtained. Some examples were worked out using reported globin sequences to show that synonymous substitutions occur at much higher rates than amino acid-altering substitutions in evolution.

Animals↗

Site-specific evolutionary rate inference: taking phylogenetic uncertainty into account.

The evolutionary rate at an amino acid site is indicative of how conserved this site is and, in turn, allows evaluating the importance of this site in maintaining the structure/function of the protein. When evolutionary rates are estimated, one must reconstruct the phylogenetic tree describing the evolutionary relationship among the sequences under study. However, if the inferred phylogenetic tree is incorrect, it can lead to erroneous site-specific rate estimates. Here we describe a novel Bayesian method that uses Markov chain Monte Carlo methodology to integrate over the space of all possible trees and model parameters. By doing so, the method considers alternative evolutionary scenarios weighted by their posterior probabilities. We show that this comprehensive evolutionary approach is superior over methods that are based on only a single tree. We illustrate the potential of our algorithm by analyzing the conservation pattern of the potassium channel protein family.

Algorithms↗

Chromosome Y microsatellites: population genetic and evolutionary aspects.

By means of a multicenter study, a large number of males have been characterized for Y-chromosome specific short tandem repeats (STRs) or microsatellites. A complete summary of the allele frequency distributions for these Y-STRs is presented in the Appendix. This manuscript describes in more detail some of the population genetic and evolutionary aspects for a restricted set of seven chromosome Y STRs in a selected number of population samples. For all the chromosome Y STRs markedly different region-specific allele frequency distributions were observed, also when closely related populations were compared. Haplotype analyses using AMOVA showed that when four different European male groups (Germans, Dutch, Swiss, Italians) were compared, less than 10% of the total genetic variability was due to differences between these populations. Nevertheless, these pairwise comparisons revealed significant differences between most population pairs. Assuming a step-wise mutation model and a mutation frequency of 0.21%, it was estimated that chromosome Y STR-based evolutionary lines of descent can be reliably inferred over a time-span of only 1950 generations (or about 49,000 years). This reduces the reliability of the inference of population affinities to a historical, rather than evolutionary time scale. This is best illustrated by the construction of a human evolutionary tree based on chromosome Y STRs in which most of the branches connect in a markedly different way compared with trees based on classical protein polymorphisms and/or mtDNA sequence variation. Thus, the chromosome Y STRs seem to be very useful in comparing closely related populations which cannot probably be separated by e.g. autosomal STRs. However, in order to be used in an evolutionary context they need to be combined with more stable Y-polymorphisms e.g. base-substitutions.

Biological Evolution↗