Weak and Strong Altruism in Trait Groups: Reproductive Suicide, Personal Fitness, and Expected Value

A simple variant of trait group selection, employing predators as the mechanism underlying group selection, supports contingent reproductive suicide as altruism (i.e., behavior lowering personal fitness while augmenting that of another) without kin assortment. The contingent suicidal type may either saturate the population or be polymorphic with a type avoiding suicide, depending on parameters. In addition to contingent suicide, this randomly assorting morph may also exhibit continuously expressed strong altruism (sensu Wilson 1979) usually thought restricted to kin selection. The model will not, however, support a sterile worker caste as such, where sterility occurs before life history events associated with effective altruism; reproductive suicide must remain fundamentally contingent (facultative sensu West Eberhard 1987; Myles 1988) under random assortment. The continuously expressed strong altruism supported by the model may be reinterpreted as probability of arbitrarily committing reproductive suicide, without benefit for another; such arbitrary suicide (a "load" on "adaptive" suicide) is viable only under a more restricted parameter space relative to the necessarily concomitant adaptive contingent suicide.

Group selection and kin selection: two concepts but one process.

In a recent paper, Traulsen and Nowak use a multilevel selection model to show that cooperation can be favored by group selection in finite populations [Traulsen A, Nowak M (2006) Proc Natl Acad Sci USA 103:10952-10955]. The authors challenge the view that kin selection may be an appropriate interpretation of their results and state that group selection is a distinctive process "that permeates evolutionary processes from the emergence of the first cells to eusociality and the economics of nations." In this paper, we start by addressing Traulsen and Nowak's challenge and demonstrate that all their results can be obtained by an application of kin selection theory. We then extend Traulsen and Nowak's model to life history conditions that have been previously studied. This allows us to highlight the differences and similarities between Traulsen and Nowak's model and typical kin selection models and also to broaden the scope of their results. Our retrospective analyses of Traulsen and Nowak's model illustrate that it is possible to convert group selection models to kin selection models without disturbing the mathematics describing the net effect of selection on cooperation.

Species-level selection reduces selfishness through competitive exclusion

Adaptation does not necessarily lead to traits which are optimal for the population. This is because selection is often the strongest at the individual or gene level. The evolution of selfishness can lead to a 'tragedy of the commons', where traits such as aggression or social cheating reduce population size and may lead to extinction. This suggests that species-level selection will result whenever species differ in the incentive to be selfish. We explore this idea in a simple model that combines individual-level selection with ecology in two interacting species. Our model is not influenced by kin or trait-group selection. We find that individual selection in combination with competitive exclusion greatly increases the likelihood that selfish species go extinct. A simple example of this would be a vertebrate species that invests heavily into squabbles over breeding sites, which is then excluded by a species that invests more into direct reproduction. A multispecies simulation shows that these extinctions result in communities containing species that are much less selfish. Our results suggest that species-level selection and community dynamics play an important role in regulating the intensity of conflicts in natural populations.

Weak and strong altruism in traitgGroups: Reproductive suicide, personal fitness and expected value

A simple variant of trait group selection, employing predators as themechanism underlying group selection, supports contingent reproductivesuicide as altruism (i.e., behavior lowering personal fitness whileaugmenting that of another) without kin assortment. The contingentsuicidal type may either saturate the population or be polymorphicwith a type avoiding suicide, depending on parameters. In addition tocontingent suicide, this randomly assorting morph may also exhibitcontinuously expressed strong altruism (sensu Wilson 1979) usuallythought restricted to kin selection. The model will not, however,support a sterile worker caste as such, where sterility occurs beforelife history events associated with effective altruism; reproductivesuicide must remain fundamentally contingent (facultative sensu WestEberhard 1987; Myles 1988) under random assortment. The continuouslyexpressed strong altruism supported by the model may be reinterpretedas probability of arbitrarily committing reproductive suicide, withoutbenefit for another; such arbitrary suicide (a "load" on "adaptive"suicide) is viable only under a more restricted parameter spacerelative to the necessarily concomitant adaptive contingent suicide.

Effects of selection and drift on G matrix evolution in a heterogeneous environment: a multivariate Qst-Fst Test with the freshwater snail Galba truncatula.

Unraveling the effect of selection vs. drift on the evolution of quantitative traits is commonly achieved by one of two methods. Either one contrasts population differentiation estimates for genetic markers and quantitative traits (the Q(st)-F(st) contrast) or multivariate methods are used to study the covariance between sets of traits. In particular, many studies have focused on the genetic variance-covariance matrix (the G matrix). However, both drift and selection can cause changes in G. To understand their joint effects, we recently combined the two methods into a single test (accompanying article by Martin et al.), which we apply here to a network of 16 natural populations of the freshwater snail Galba truncatula. Using this new neutrality test, extended to hierarchical population structures, we studied the multivariate equivalent of the Q(st)-F(st) contrast for several life-history traits of G. truncatula. We found strong evidence of selection acting on multivariate phenotypes. Selection was homogeneous among populations within each habitat and heterogeneous between habitats. We found that the G matrices were relatively stable within each habitat, with proportionality between the among-populations (D) and the within-populations (G) covariance matrices. The effect of habitat heterogeneity is to break this proportionality because of selection for habitat-dependent optima. Individual-based simulations mimicking our empirical system confirmed that these patterns are expected under the selective regime inferred. We show that homogenizing selection can mimic some effect of drift on the G matrix (G and D almost proportional), but that incorporating information from molecular markers (multivariate Q(st)-F(st)) allows disentangling the two effects.

Cultural transmission can inhibit the evolution of altruistic helping.

The study of culturally inherited traits has led to the suggestion that the evolution of helping behaviors is more likely with cultural transmission than without. Here we evaluate this idea through a comparative analysis of selection on helping under both genetic and cultural inheritance. We develop two simple models for the evolution of helping through cultural group selection: one in which selection on the trait depends solely on Darwinian fitness effects and one in which selection is driven by nonreproductive factors, specifically imitation of strategies achieving higher payoffs. We show that when cultural variants affect Darwinian fitness, the selection pressure on helping can be markedly increased relative to that under genetic transmission. By contrast, when variants are driven by nonreproductive factors, the selection pressure on helping may be reduced relative to that under genetic inheritance. This occurs because, unlike biological offspring, the spread of cultural variants from one group to another through imitation does not reduce the number of these variants in the source group. As a consequence, there is increased within-group competition associated with traits increasing group productivity, which reduces the benefits of helping. In these cases, selection for harming behavior (decreasing the payoff to neighbors) may occur rather than selection for helping.

Selection on age at first and at last reproduction in the long-lived Alpine Swift Apus melba

The way an organism spreads its reproduction over time is defined as a life-history trait, and selection is expected to favour life-history traits associated with the highest fitness return. We use a long-term dataset of 277 life histories to investigate the shape and strength of selection acting on the age at first reproduction and at last reproduction in the long-lived Alpine Swift. Both traits were under strong directional selection, but in opposite directions, with selection favouring birds starting their reproductive career early and being able to reproduce for longer. There was also evidence for stabilising selection acting on both traits, suggesting that individuals should nonetheless refrain from reproducing in their first 2 years of life (i.e. when inexperienced), and that reproducing after 7 years of age had little effect on lifetime fitness, probably due to senescence.

Long-run selection and the work ethic

That individuals contribute in social dilemma interactions even when contributing is costly is a well-established observation in the experimental literature. Since a contributor is always strictly worse off than a non-contributor the question is raised if an intrinsic motivation to contribute can survive in an evolutionary setting. Using recent results on deterministic approximation of stochastic evolutionary dynamics we give conditions for equilibria with a positive number of contributors to be selected in the long run.

Les origines biologiques de la musique.

La musique est un comportement humain incontestablement universel, elle demeure néanmoins peu abordée par l’anthropologie. Si les connaissances empiriques accumulées à ce jour ont permis de bien la caractériser à des niveaux proximaux d’analyse, la question de son origine évolutionniste est, en contrepartie, souvent délaissée. Or, toute tentative sérieuse de comprendre ce phénomène requiert une investigation de sa fonction adaptative et de sa phylogénèse. Le projet entrepris ici consiste en une tentative de définition du concept de musique en terme d’universaux, d’une comparaison interspécifique du phénomène et d’un résumé de l’histoire phylogénétique des comportements musicaux, ainsi que d’une analyse de deux modèles portant sur les origines de la musique (Miller, 2000; Mithen, 2006). De ces modèles sont extraites des prévisions qui sont confrontées à des données empiriques provenant de disciplines diverses afin d’évaluer leur valeur scientifique. L’analyse des données disponibles permet de produire un inventaire des universaux musicaux aux plans cognitif, structurel, émotionnel, fonctionnel et symbolique et d’identifier ainsi certaines des bases biologiques du phénomène. Plusieurs mécanismes évolutionnistes, dont la sélection naturelle, la sélection sexuelle, la sélection de groupe et la sélection parentale sont employés par les divers auteurs afin d’expliquer l’apparition du phénomène musical. Il appert que la musique a joué un rôle important dans la relation parent-enfant au cours de l’évolution humaine, de même que dans la cohésion sociale, la coordination des activités et la formation de l’identité de groupe. En ce qui a trait aux deux modèles analysés ici, chacun ne traite que d’une partie des invariants musicaux et leur comparaison permet d’établir qu’ils sont mutuellement exclusifs. En guise de conclusion, nous tentons de formuler un scénario évolutif qui concilie les différentes hypothèses abordées.

Estudo da habilidade de permanência de fêmeas da raça Canchim aos 76 meses de idade

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Change and Aging Senescence as an Adaptation

Understanding why we age is a long-lived open problem in evolutionary biology. Aging is prejudicial to the individual, and evolutionary forces should prevent it, but many species show signs of senescence as individuals age. Here, I will propose a model for aging based on assumptions that are compatible with evolutionary theory: i) competition is between individuals; ii) there is some degree of locality, so quite often competition will be between parents and their progeny; iii) optimal conditions are not stationary, and mutation helps each species to keep competitive. When conditions change, a senescent species can drive immortal competitors to extinction. This counter-intuitive result arises from the pruning caused by the death of elder individuals. When there is change and mutation, each generation is slightly better adapted to the new conditions, but some older individuals survive by chance. Senescence can eliminate those from the genetic pool. Even though individual selection forces can sometimes win over group selection ones, it is not exactly the individual that is selected but its lineage. While senescence damages the individuals and has an evolutionary cost, it has a benefit of its own. It allows each lineage to adapt faster to changing conditions. We age because the world changes.

Studies on the genetics of artemisinin resistance in malaria

A existência de estirpes de Plasmodium falciparum resistentes a multiplos fármacos é um dos problemas mais graves no controlo da malária. Novos fármacos, como a artemisinina (ART) e seus derivados são cada vez mais utilizados no tratamento da malaria e muito embora até ao momento não haja registos de fármaco-resistência estável à ART o seu surgimento seria desastroso devido á falta de alternativas. A investigação apresentada nesta tese descreve a selecção de resistência estável à ART e ao artesunato (ATN) utilizando um modelo roedor de malária, o parasita Plasmodium chabaudi chabaudi (Plasmodium chabaudi). Dois clones de Plasmodium chabaudi diferentes, AS-15CQ e AS-30CQ, foram inoculados em murganhos que por sua vez foram tratados na presença de concentrações sucessivamente crescentes de ATN e ART, sendo que no final do processo de seleção de resistência, os parasitas obtidos apresentavam uma resistência de 6 e 15 vezes superior ao ATN e à ART, respectivamente, em relação aos parasitas iniciais. Os clones obtidos foram nomeados respectivamente AS-ATN (obtido a partir de AS-15CQ por seleção com pressão de ATN) e AS-ART (obtido a partir de AS-30CQ por seleção com pressão de ART). A resistência obtida durante o processo de seleção é estável após clonagem, congelamento/descongelamento, passagem sanguínea na ausência de pressão de fármaco e transmissão natural através do mosquito vector. A sequência nucleotídica e o número de cópias dos genes previamente descritos na literatura como moduladores putativos de resistência à ART e seus derivados: mdr1, cg10, tctp e atp6; foi comparada entre parasitas resistentes e sensíveis, não tendo sido encontradas nenhumas alterações, quer na sequência quer no número de cópias destes genes. Posteriormente, numa tentativa de identificar os genes envolvidos na resistância à ART e ao ATN a técnica de Linkage Group Selection (LGS) foi utilizada. Para tal dois cruzamentos genéticos foram realizados. Estes cruzamentos foram realizados entre os clones fármaco-resistentes; AS-ART e AS-ATN e um clone geneticamente distinto dos anteriores e sensível aos fármacos em estudos, AJ. Após realização do LGS quatro loci genéticos; nos cromossomas de P. chabaudi 1, 2, 6 e 8 foram encontrados associados à resistência. Atendendo a que, a selecção no cromossoma 2 era a mais forte, este locus foi submetido a subsequentes análises genéticas, tendo sido encontradas duas mutações diferentes (V739F e V770F) num gene que codifica para um enzima de desubiquitinação (gene ubp-1).

Studies on the genetics of artemisinin resistance in malaria

A existência de estirpes de Plasmodium falciparum resistentes a multiplos fármacos é um dos problemas mais graves no controlo da malária. Novos fármacos, como a artemisinina (ART) e seus derivados são cada vez mais utilizados no tratamento da malaria e muito embora até ao momento não haja registos de fármaco-resistência estável à ART o seu surgimento seria desastroso devido á falta de alternativas. A investigação apresentada nesta tese descreve a selecção de resistência estável à ART e ao artesunato (ATN) utilizando um modelo roedor de malária, o parasita Plasmodium chabaudi chabaudi (Plasmodium chabaudi). Dois clones de Plasmodium chabaudi diferentes, AS-15CQ e AS-30CQ, foram inoculados em murganhos que por sua vez foram tratados na presença de concentrações sucessivamente crescentes de ATN e ART, sendo que no final do processo de seleção de resistência, os parasitas obtidos apresentavam uma resistência de 6 e 15 vezes superior ao ATN e à ART, respectivamente, em relação aos parasitas iniciais. Os clones obtidos foram nomeados respectivamente AS-ATN (obtido a partir de AS-15CQ por seleção com pressão de ATN) e AS-ART (obtido a partir de AS-30CQ por seleção com pressão de ART). A resistência obtida durante o processo de seleção é estável após clonagem, congelamento/descongelamento, passagem sanguínea na ausência de pressão de fármaco e transmissão natural através do mosquito vector. A sequência nucleotídica e o número de cópias dos genes previamente descritos na literatura como moduladores putativos de resistência à ART e seus derivados: mdr1, cg10, tctp e atp6; foi comparada entre parasitas resistentes e sensíveis, não tendo sido encontradas nenhumas alterações, quer na sequência quer no número de cópias destes genes. Posteriormente, numa tentativa de identificar os genes envolvidos na resistância à ART e ao ATN a técnica de Linkage Group Selection (LGS) foi utilizada. Para tal dois cruzamentos genéticos foram realizados. Estes cruzamentos foram realizados entre os clones fármaco-resistentes; AS-ART e AS-ATN e um clone geneticamente distinto dos anteriores e sensível aos fármacos em estudos, AJ. Após realização do LGS quatro loci genéticos; nos cromossomas de P. chabaudi 1, 2, 6 e 8 foram encontrados associados à resistência. Atendendo a que, a selecção no cromossoma 2 era a mais forte, este locus foi submetido a subsequentes análises genéticas, tendo sido encontradas duas mutações diferentes (V739F e V770F) num gene que codifica para um enzima de desubiquitinação (gene ubp-1).

HUMAN TOXOPLASMOSIS OUTBREAKS AND THE AGENT INFECTING FORM. FINDINGS FROM A SYSTEMATIC REVIEW

SUMMARY Toxoplasmosis, a worldwide highly prevalent zoonotic infection, is transmitted either by the oocysts, from water and soil, or the tissue cysts, in raw or undercooked infected meat, of Toxoplasma gondii. An ongoing debate is whether there are differences between the clinical and epidemiological characteristics of the outbreaks due to one or the other infective form of the agent. We performed a systematic review, recovering 437 reported outbreaks of which 38 were selected. They were complete reports containing ascribed Toxoplasma infecting form, and clinical and demographic data. There was no gender or age group selection in the outbreaks, which were described more often in the Americas. A large number of individuals were affected when oocysts, associated with soil and water contaminated with cat feces, were considered the transmission source. Onset of symptoms occurred early when the infection was ascribed to meat tissue cysts (11.4 ± 6.7 days) with sharpened temporal distribution of cases, while a broader and prolonged appearance of new cases was observed when oocysts in water were the source of the infection (20 ± 7 days, p < 0.001). Such information may be useful in the design and implementation of control strategies.

Multivariate QST-FST comparisons: a neutrality test for the evolution of the g matrix in structured populations.

Neutrality tests in quantitative genetics provide a statistical framework for the detection of selection on polygenic traits in wild populations. However, the existing method based on comparisons of divergence at neutral markers and quantitative traits (Q(st)-F(st)) suffers from several limitations that hinder a clear interpretation of the results with typical empirical designs. In this article, we propose a multivariate extension of this neutrality test based on empirical estimates of the among-populations (D) and within-populations (G) covariance matrices by MANOVA. A simple pattern is expected under neutrality: D = 2F(st)/(1 - F(st))G, so that neutrality implies both proportionality of the two matrices and a specific value of the proportionality coefficient. This pattern is tested using Flury's framework for matrix comparison [common principal-component (CPC) analysis], a well-known tool in G matrix evolution studies. We show the importance of using a Bartlett adjustment of the test for the small sample sizes typically found in empirical studies. We propose a dual test: (i) that the proportionality coefficient is not different from its neutral expectation [2F(st)/(1 - F(st))] and (ii) that the MANOVA estimates of mean square matrices between and among populations are proportional. These two tests combined provide a more stringent test for neutrality than the classic Q(st)-F(st) comparison and avoid several statistical problems. Extensive simulations of realistic empirical designs suggest that these tests correctly detect the expected pattern under neutrality and have enough power to efficiently detect mild to strong selection (homogeneous, heterogeneous, or mixed) when it is occurring on a set of traits. This method also provides a rigorous and quantitative framework for disentangling the effects of different selection regimes and of drift on the evolution of the G matrix. We discuss practical requirements for the proper application of our test in empirical studies and potential extensions.