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General models of multilocus evolution.

In 1991, Barton and Turelli developed recursions to describe the evolution of multilocus systems under arbitrary forms of selection. This article generalizes their approach to allow for arbitrary modes of inheritance, including diploidy, polyploidy, sex linkage, cytoplasmic inheritance, and genomic imprinting. The framework is also extended to allow for other deterministic evolutionary forces, including migration and mutation. Exact recursions that fully describe the state of the population are presented; these are implemented in a computer algebra package (available on the Web at http://helios.bto.ed.ac.uk/evolgen). Despite the generality of our framework, it can describe evolutionary dynamics exactly by just two equations. These recursions can be further simplified using a "quasi-linkage equilibrium" (QLE) approximation. We illustrate the methods by finding the effect of natural selection, sexual selection, mutation, and migration on the genetic composition of a population.

Emigration and Immigration↗

Play in evolution, culture, and individual adaptation: implications for therapy.

In this paper human play is examined from the perspective of its role in evolution. A model of social evolution is proposed to extend the model of biological evolution and to demonstrate the central role of play in the social-evolutionary process. Since in evolution play is a mechanism for its process, it demands species members who are playful. Thus, man is a player by virtue of species membership and the evolutionary pathway of the species. The implications of this deep-seated characteristic of playfulness are discussed in terms of human development. Finally, a set of principles is derived from the evolutionary-developmental characteristics of play. Application of these principles in clinical practice is demonstrated and discussed.

Adolescent↗

A molecular model for the evolution of endothermy in the theropod-bird lineage.

Ectothermy is a primitive state; therefore, a shared common ancestor of crocodiles, dinosaurs, and birds was at some point ectothermic. Birds, the extant descendants of the dinosaurs, are endothermic. Neither the metabolic transition within this lineage nor the place the dinosaurs held along the ectothermic-endothermic continuum is defined. This paper presents a conceptual model for the evolution of endothermy in the theropod-bird lineage. It is recognized that other animals (some fish, insects, etc.) are functionally endothermic. However, endothermy in other clades is beyond the scope of this paper, and we address the onset of endothermy in only the theropod/bird clade. The model begins with simple changes in a single gene of a common ancestor, and it includes a series of concomitant physiological and morphological changes, beginning perhaps as early as the first archosaurian common ancestor of dinosaurs and crocodiles. These changes continued to accumulate within the theropod-avian lineage, were maintained and refined through selective forces, and culminated in extant birds. Metabolic convergence or homoplasy is evident in the inherent differences between the endothermy of mammals and the endothermy of extant birds. The strength and usefulness of this model lie in the phylogenetic, genetic, evolutionary, and adaptive plausibility of each of the suggested developmental steps toward endothermy. The model, although conceptual in nature, relies on an extensive knowledge base developed by numerous workers in each of these areas. In addition, the model integrates known genetic, metabolic, and developmental aspects of extant taxa that phylogenetically bracket theropod dinosaurs for comparison with information derived from the fossil record of related extinct taxa.

Adaptation, Physiological↗

A theory of pragmatic information and its application to the quasi-species model of biological evolution.

'Standard' information theory says nothing about the semantic content of information. Nevertheless, applications such as evolutionary theory demand consideration of precisely this aspect of information, a need that has motivated a largely unsuccessful search for a suitable measure of an 'amount of meaning'. This paper represents an attempt to move beyond this impasse, based on the observation that the meaning of a message can only be understood relative to its receiver. Positing that the semantic value of information is its usefulness in making an informed decision, we define pragmatic information as the information gain in the probability distributions of the receiver's actions, both before and after receipt of a message in some pre-defined ensemble. We then prove rigorously that our definition is the only one that satisfies obvious desiderata, such as the additivity of information from logically independent messages. This definition, when applied to the information 'learned' by the time evolution of a process, defies the intuitions of the few previous researchers thinking along these lines by being monotonic in the uncertainty that remains after receipt of the message, but non-monotonic in the Shannon entropy of the input ensemble. It also follows that the pragmatic information of the genetic 'messages' in an evolving population is a global Lyapunov function for Eigen's quasi-species model of biological evolution. A concluding section argues that a theory such as ours must explicitly acknowledge purposeful action, or 'agency', in such diverse fields as evolutionary theory and finance.

Biological Evolution↗

A new syndrome with quadrupedal gait, primitive speech, and severe mental retardation as a live model for human evolution.

The author has discovered a new syndrome with quadrupedal gait, flexed head and body, primitive speech, severe mental retardation, and mild cerebellar signs with a disturbed conscious experience. This syndrome was exhibited by 5 of 19 children from a consanguineous family. The pedigree demonstrated a typical autosomal-recessive inheritance. The genetic nature of this syndrome suggests a backward stage in human evolution, which is most probably caused by a genetic mutation, rendering, in turn, the transition from quadrupedality to bipedality. This would then be consistent with theories of punctuated evolution. On the other hand, the extensor motor system causing a resistance of the body against the gravity may actually be subjected to evolutionary forces. This new syndrome may be used as a live model for human evolution. An accompanying video clip for this article is available as a downloadable file accompanying the official online version of International Journal of Neuroscience. To access it, click on the issue link for 116(3), then select this article. A download option appears at the bottom of this abstract.

Adolescent↗

A kin selection model for the evolution of virulence.

The costs and benefits of parasite virulence are analysed in an evolutionarily stable strategy (ESS) model. Increased host mortality caused by disease (virulence) reduces a parasite's fitness by damaging its food supply. The fitness costs of high virulence may be offset by the benefits of increased transmission or ability to withstand the host's defences. It has been suggested that multiple infections lead to higher virulence because of competition among parasite strains within a host. A quantitative prediction is given for the ESS virulence rate as a function of the coefficient of relatedness among co-infecting strains. The prediction depends on the quantitative relation between the costs of virulence and the benefits of transmission or avoidance of host defences. The particular mechanisms by which parasites can increase their transmission or avoid host defences also have a key role in the evolution of virulence when there are multiple infections.

Animals↗

A model for the evolution of networks of genes.

An organism persists through the activity of structural genes, which is co-ordinated by clusters of coupled regulatory genes. During evolution, changes of coupling within a cluster can increase the reliability with which its structural genes perform a task. To study the evolution of coupling, we have simulated and analyzed a stochastic model for a simple problem. The assumptions of the model are these: A network of regulatory genes co-ordinates the synthesis of four structural proteins, which associate in distinct heterodimers that form a heterotetramer. Mutation in cis-regulatory regions produces transitions among 64 types of network. In a population, each network reproduces in proportion to its fitness, which depends on its probability (reliability) of synthesizing the tetramer. Fitness-dependent attrition keeps the size of the population constant. Regulatory genes occur in a sequence of levels; each level is associated with a different family of transcription factors. The following results emerge: Because different messengers within a family can give networks with the same connectivity, the 64 types of networks cluster into eight equivalence classes. During evolution with a low mutation rate, high-fitness classes can be approached through various paths on a fitness landscape. With a higher mutation rate, networks remain more uniformly distributed among the 64 types, and lower-fitness networks remain preponderant. An initially homogeneous population becomes more heterogeneous through mutation, but selection according to fitness later reduces its diversity. During this process the dispersion of the population over the possible networks increases, then decreases as the population approaches a unique steady state.

Animals↗

A general model for the evolution of mutualisms.

The evolution of mutualisms presents a puzzle. Why does selection favour cooperation among species rather than cheaters that accept benefits but provide nothing in return? Here we present a general model that predicts three key factors will be important in mutualism evolution: (i) high benefit to cost ratio, (ii) high within-species relatedness and (iii) high between-species fidelity. These factors operate by moderating three types of feedback benefit from mutualism: cooperator association, partner-fidelity feedback and partner choice. In defining the relationship between these processes, our model also allows an assessment of their relative importance. Importantly, the model suggests that phenotypic feedbacks (partner-fidelity feedback, partner choice) are a more important explanation for between-species cooperation than the development of genetic correlations among species (cooperator association). We explain the relationship of our model to existing theories and discuss the empirical evidence for our predictions.

Biological Evolution↗

Time evolution of the Partridge-Barton model.

The time evolution of the Partridge-Barton model in the presence of the pleiotropic constraint and deleterious somatic mutations is exactly solved for arbitrary fecundity in the context of a matricial formalism. Analytical expressions for the time dependence of the mean survival probabilities are derived. Using the fact that the asymptotic behavior for large time t is controlled by the largest matrix eigenvalue, we obtain the steady state values for the mean survival probabilities and the Malthusian growth exponent. The mean age of the population exhibits a t-1 power law decayment. Some Monte Carlo simulations were also performed and they corroborated our theoretical results.

Biological Evolution↗

A model for the evolution of reproductive skew without reproductive suppression

Reproductive skew is a measure of the way breeding is distributed among the members of an animal society or group. Up to now, explanations of patterns of skew have been limited to one particular model, which assumes that a single dominant has full control over the distribution of subordinate reproduction. If this control is incomplete or absent, however, unsanctioned breeding by subordinate females will increase the total number of young produced. Here I present a new model for the evolution of skew that considers the effect of brood size on the inclusive fitness of dominants and subordinates. By augmenting brood size, a subordinate female reduces the per capita fitness of a dominant's offspring, so the net benefits of producing young are lower for related subordinates. I consider the stable level of skew when both dominant and subordinate attempt to maximize their inclusive fitness under two conditions: (1) when the dominant is unable to anticipate that a subordinate will add to her brood; and (2) the dominant does anticipate subordinate reproduction and can respond by adjusting her own brood size. In the first case, the model predicts that reproductive skew will increase with relatedness between breeders, because related subordinates are selected to add fewer young to the dominant's brood. In the second case, the dominant's optimal response to the presence of a second breeder exaggerates the relationship between relatedness and skew: dominants should produce more young when breeding with related compared with unrelated subordinates. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Cloning of the HSP70 gene from Halobacterium marismortui: relatedness of archaebacterial HSP70 to its eubacterial homologs and a model for the evolution of the HSP70 gene.

Heat shock induces the synthesis of a set of proteins in Halobacterium marismortui whose molecular sizes correspond to the known major heat shock proteins. By using the polymerase chain reaction and degenerate oligonucleotide primers for conserved regions of the 70-kDa heat shock protein (HSP70) family, we have successfully cloned and sequenced a gene fragment containing the entire coding sequence for HSP70 from H. marismortui. HSP70 from H. marismortui shows between 44 and 47% amino acid identity with various eukaryotic HSP70s and between 51 and 58% identity with its eubacterial and archaebacterial homologs. On the basis of a comparison of all available HSP70 sequences, we have identified a number of unique sequence signatures in this protein family that provide a clear distinction between eukaryotic organisms and prokaryotic organisms (archaebacteria and eubacteria). The archaebacterial (viz., H. marismortui and Methanosarcina mazei) HSP70s have been found to contain all of the signature sequences characteristic of eubacteria (particularly the gram-positive bacteria), which suggests a close evolutionary relationship between these groups. In addition, detailed analyses of HSP70 sequences that we have carried out have revealed a number of additional novel features of the HSP70 protein family. These include (i) the presence of an insertion of about 25 to 27 amino acids in the N-terminal quadrants of all known eukaryotic and prokaryotic HSP70s except those from archaebacteria and the gram-positive group of bacteria, (ii) significant sequence similarity in HSP70 regions comprising its first and second quadrants from organisms lacking the above insertion, (iii) highly significant similarity between a protein, MreB, of Escherichia coli and the N-terminal half of HSP70s, (iv) significant sequence similarity between the N-terminal quadrant of HSP70 (from gram-positive bacteria and archaebacteria) and the m-type thioredoxin of plant chloroplasts. To account for these and other observations, a model for the evolution of HSP70 proteins involving gene duplication is proposed. The model proposes that HSP70 from archaebacteria (H. marismortui and M. mazei) and the gram-positive group of bacteria constitutes the ancestral form of the protein and that all other HSP70s (viz., other eubacteria as well as eukaryotes) containing the insert have evolved from this ancient protein.

Amino Acid Sequence↗

Xenopus as an experimental model for studying evolution of hsp--immune system interactions.

The frog Xenopus provides a unique model system for studying the evolutionary conservation of the immunological properties of heat shock proteins (hsps). General methods for maintaining and immunizing isogenetic clones of defined MHC genotypes are presented together with more recently developed protocols for exploring hsp-mediated immune responses in vitro (proliferative and cytotoxic assays) and in vivo (adoptive cell transfer and antibody treatment) in adults and in naturally MHC class I-deficient larvae. Finally, techniques to study modalities of expression of the endoplasmic reticulum resident gp96 at the cell surface of tumor and normal lymphocytes are considered.

Animals↗

Genetic variation in the Heterodoxus octoseriatus group (Phthiraptera): a test of Price's model of parasite evolution.

Most of the genetic variation in the H. octoseriatus group is present as fixed gene differences between species which have been described on morphological criteria. Based on allozymes, the taxonomic status of some species was challenged. There was insufficient evidence, however, to demonstrate that these were not 'good' biological species. Overall, the limited intraspecific variation was present as fixed gene differences among lice from different hosts and from different colonies of hosts; heterozygotes were rare. Two predictions derived from Price's model of parasite evolution were met: populations of lice were genetically homogeneous and, where genetic markers were present, we found substantial genetic variation among populations. These data contrast with those for endoparasitic helminths, where, in general, the amount of genetic variation is similar to that of free-living invertebrates.

Animals↗

[Modelling of spatial evolution and dynamics of a population of healthy then rabies infected foxes].

The authors describe the main feature of a computer model which helps to simulate the evolution of a rabies epi-enzootic in foxes. They show first the goals and interests of the study, then the originality of used methodology. Their results deal successively with dynamic evolution of a healthy population of foxes, then with this same population infected with rabies and, at last, spatial and temporal evolution of the enzootics. Simulated results are discussed by comparison with those observed in the field.

Animals↗

Genetic diversity and models of viral evolution for the hepatitis C virus.

In this review we discuss the application of theoretical frameworks to the interpretation of viral gene sequence data, with particular reference to the hepatitis C virus (HCV). The increasing availability of such data means that it is now possible (and necessary) to proceed from simple qualitative models of viral evolution, to more quantitative frameworks based on statistical inference, notably population genetics and molecular phylogenetics. We argue that these approaches are invaluable tools to the virologist and are essential for understanding the dynamics of viral infection and the outcome of therapeutic strategies. We use several recent HCV data-sets to illustrate the methods.

Genetic Variation↗

Functional morphology of beta cells in the area centralis of the cat's retina: a model for the evolution of central retinal specializations.

The dendritic morphology of beta cells in and around the area centralis of the retinae of normally pigmented and Siamese cats is described. Individual central beta cells in the Siamese cat do not differ morphologically from central beta cells in normally pigmented cats, and in both groups of animals, there is a clear morphological continuity between central and peripheral beta cells. On the basis of systematic patterns of beta cell dendritic orientation, ther area centralis of the normal cat can be divided into a central region, approximately 200 micrometers in diameter, and a pericentral region, approximately 1,400 micrometers in diameter. In the central region, nearly all beta cells have a single large primary dendrite which descends perpendicular to the plane of inner plexiform layer, and gives rise to a dendritic tree which is vertically aligned with the cell's soma. In the pericentral region, the single primary dendrite of most cells descends obliquely through the inner plexiform layer and gives rise to a dendritic tree which is displaced laterally from the position of the soma. For most of the cells the trajectory of the dendrite is systematically related to the location of the cell relative to the area centralis such that the somas are displaced away from its center, presumably in order to minimize the thickness of the ganglion cell layer in the high acuity region. Many beta cells outside the pericentral region also have oriented single primary dendrites, but their orientation seems fairly random with respect to the location of the area centralis. In the Siamese area centralis, this systematic pattern of beta cell dendritic orientation is markedly reduced, suggesting that the pattern is under genetic control. On the basis of these observations, a model for the evolution of the area centralis and fovea is presented which involves selection for systematic for systematic patterns of dendritic orientation in regions of high ganglion cell density.

Animals↗