Grandmothering and natural selection

Humans are unique among primates in that women regularly outlive their reproductive period by decades. The grandmother hypothesis proposes that natural selection increased the length of the human post-menopausal period—and, thus, extended longevity—as a result of the inclusive fitness benefits of grandmothering. However, it has yet to be demonstrated that the inclusive fitness benefits associated with grandmothering are large enough to warrant this explanation. Here, we show that the inclusive fitness benefits are too small to affect the evolution of longevity under a wide range of conditions in simulated populations. This is due in large part to the relatively weak selection that applies to women near or beyond the end of their reproductive period. However, we find that grandmothers can facilitate the evolution of a shorter reproductive period when their help decreases the weaning age of their matrilineal grandchildren. Because selection favours a shorter reproductive period in the presence of shorter interbirth intervals, this finding holds true for any form of allocare that helps mothers resume cycling more quickly. We conclude that while grandmothering is unlikely to explain human-like longevity, allocare could have played an important role in shaping other unique aspects of human life history, such as a later age at first birth and a shorter female reproductive period.     

The meaning of natural selection revisited at the molecular level

Various molecular interaction mechanisms cause biased transmission of genes. Meiotic drive is an example of strong bias, and compartmentalization of mammalian chromosomes reflects weak bias. Such biases are the results of interaction between DNA and proteins, and should be distinguished from natural selection. Separating the effects of molecular interaction from those of natural selection, however, is often very difficult. Natural selection and molecular mechanisms interact, and our understanding of how selection works requires revision.     

From survivors to replicators: evolution by natural selection revisited

For evolution by natural selection to occur it is classically admitted that the three ingredients of variation, difference in fitness and heredity are necessary and sufficient. In this paper, I show using simple individual-based models, that evolution by natural selection can occur in populations of entities in which neither heredity nor reproduction are present. Furthermore, I demonstrate by complexifying these models that both reproduction and heredity are predictable Darwinian products (i.e. complex adaptations) of populations initially lacking these two properties but in which new variation is introduced via mutations. Later on, I show that replicators are not necessary for evolution by natural selection, but rather the ultimate product of such processes of adaptation. Finally, I assess the value of these models in three relevant domains for Darwinian evolution.     

Maladaptation and natural selection

The transformations George Williams initiated in evolutionary biology seem so blindingly obvious in retrospect that they spur the question of why he saw what no one else did. While most humans are prone to see only what theory predicts, Williams sees in bold relief whatever does not fit. Not an adaptationist or an anti-adaptationist, Williams is better described as a maladaptionist. The challenge of finding evolutionary explanations for apparent maladaptations has been overlooked with casualness akin to that once typical for group selection. Suboptimal traits tend to be dismissed as illustrations of the weakness and stochastic nature of selection compared with mutation and drift. A closer look suggests that such constraints are only one of six possible kinds of explanations for apparently suboptimal designs: mismatch, coevolution, tradeoffs, constraints, reproductive advantage at the expense of the individual, and defenses that are aversive but useful Medicine has asked proximate questions at every possible level but has only begun to ask evolutionary questions about why bodies are vulnerable to disease. Considering all six possible evolutionary reasons for apparently suboptimal traits will spur progress not only in medicine but also more generally in biology. 'Williams Vision" may not yield a net benefit to the possessor, but it is invaluable for the species.     

Humanity and natural selection

In some modern biological images of the human species human action is reduced to a consequence of natural selection, that is, to a tendency to maximize fitness. The precise nature and scope of the theory of natural selection are, however, undecided; yet in evolutionary interpretations of human society neodarwinism is often treated as a dogma, and natural selection sometimes becomes a transcendental force. There are instances of changes in gene frequencies, in human as well as other populations, that conform with neodarwinian (“socio-biological”) presumptions; but it is not known to what extent the evolution of the human (or other) species has been due to such changes. There is still less ground for explaining the diversity of human societies in this way.The image-makers also make illegitimate use of the comparative method: familiar features of human conduct, such as favoring kin and reciprocation, are used in accounts of animal behavior, and are then rediscovered among human beings; during this two-way transfer, the meanings of words are changed; altruism, egoism, and deceit lose their moral content, and the complex human idea of kin is reduced to a measure of genetical similarity.The intention of neodarwinists is to reveal a human nature determined by evolutionary processes, but one of the most important distinctive features of our species is the plasticity of our behavior, attitudes, and intentions. Moreover, if neodarwinian premises are accepted, to speak of intention is misleading, and there are no independent criteria by which neodarwinian (or any other) arguments may be judged: all one does is regulated by the need to maximise one's inclusive fitness. Hence much writing in this field wavers between an uncompromising reduction of human action to considerations of population genetics, on the one hand, and a recognition that there are other kinds of authentic knowledge about human beings, on the other. Among the latter is historical knowledge.The neodarwinian images of humanity emphasize human depravity. In their misanthropy they reflect the outlook of conservative pessimists who have influenced European thought for two and a half millennia, and whose views imply that most attempts to improve the human condition are against nature and so must fail.An alternative, which corresponds to the facts of everyday life and of history, is that human beings are capable of rejecting what is conventionally held to be inevitable, and of determining their destiny by conscious, deliberate action.     

Grandmothering and the evolution of homo erectus

Despite recent, compelling challenge, the evolution of Homo erectus is still commonly attributed to big game hunting and/or scavenging and family provisioning by men. Here we use a version of the "grandmother" hypothesis to develop an alternative scenario, that climate-driven adjustments in female foraging and food sharing practices, possibly involving tubers, favored significant changes in ancestral life history, morphology, and ecology leading to the appearance, spread and persistence of H. erectus. Available paleoclimatic, environmental, fossil and archaeological data are consistent with this proposition; avenues for further critical research are readily identified. This argument has important implications for widely-held ideas about the recent evolution of long human lifespans, the prevalence of male philopatry among ancestral hominids, and the catalytic role of big game hunting and scavenging in early human evolution.     

Sexual selection and natural selection in bird speciation

The role of sexual selection in speciation is investigated, addressing two main issues. First, how do sexually selected traits become species recognition traits? Theory and empirical evidence suggest that female preferences often do not evolve as a correlated response to evolution of male traits. This implies that, contrary to runaway (Fisherian) models of sexual selection, premating isolation will not arise as an automatic side effect of divergence between populations in sexually selected traits. I evaluate premating isolating mechanisms in one group, the birds. In this group premating isolation is often a consequence of sexual imprinting, whereby young birds learn features of their parents and use these features in mate choice. Song, morphology and plumage are known recognition cues. I conclude that perhaps the main role for sexual selection in speciation is in generating differences between populations in traits. Sexual imprinting then leads to these traits being used as species recognition mechanisms. The second issue addressed in this paper is the role of sexual selection in adaptive radiation, again concentrating on birds. Ecological differences between species include large differences in size, which may in themselves be sufficient for species recognition, and differences in habitat, which seem to evolve frequently and at all stages of an adaptive radiation. Differences in habitat often cause song and plumage patterns to evolve as a result of sexual selection for efficient communication. Therefore sexual selection is likely to have an important role in generating premating isolating mechanisms throughout an adaptive radiation. It is also possible that sexual selection, by creating more allopatric species, creates more opportunity for ecological divergence to occur. The limited available evidence does not support this idea. A role for sexual selection in accelerating ecological diversification has yet to be demonstrated.     

Visualizing and quantifying natural selection

Modern methods of analysis are enabling researchers to study natural selection at a new level of detail. Multivariate statistical techniques can Identify specific targets of selection and provide parameter estimates that fit into equations for evolutionary change. A more Intuitive understanding of the form of selection can be provided through graphical representation of selection surfaces. Combinations of quantitative and visual analyses are providing researchers with new insights into the details of natural selection in the wild.     

Genetic linkage and natural selection

The prevalence of recombination in eukaryotes poses one of the most puzzling questions in biology. The most compelling general explanation is that recombination facilitates selection by breaking down the negative associations generated by random drift (i.e. Hill–Robertson interference, HRI). I classify the effects of HRI owing to: deleterious mutation, balancing selection and selective sweeps on: neutral diversity, rates of adaptation and the mutation load. These effects are mediated primarily by the density of deleterious mutations and of selective sweeps. Sequence polymorphism and divergence suggest that these rates may be high enough to cause significant interference even in genomic regions of high recombination. However, neither seems able to generate enough variance in fitness to select strongly for high rates of recombination. It is plausible that spatial and temporal fluctuations in selection generate much more fitness variance, and hence selection for recombination, than can be explained by uniformly deleterious mutations or species-wide selective sweeps.     

Sexual selection and speciation: the comparative evidence revisited

The spectacular diversity in sexually selected traits in the animal kingdom has inspired the hypothesis that sexual selection can promote species divergence. In recent years, several studies have attempted to test this idea by correlating species richness with estimates of sexual selection across phylogenies. These studies have yielded mixed results and it remains unclear whether the comparative evidence can be taken as generally supportive. Here, we conduct a meta‐analysis of the comparative evidence and find a small but significant positive overall correlation between sexual selection and speciation rate. However, we also find that effect size estimates are influenced by methodological choices. Analyses that included deeper phylogenetic nodes yielded weaker correlations, and different proxies for sexual selection showed different relationships with species richness. We discuss the biological and methodological implications of these findings. We argue that progress requires more representative sampling and justification of chosen proxies for sexual selection and speciation rate, as well as more mechanistic approaches.     

Limits to natural selection

We review the various factors that limit adaptation by natural selection. Recent discussion of constraints on selection and, conversely, of the factors that enhance "evolvability", have concentrated on the kinds of variation that can be produced. Here, we emphasise that adaptation depends on how the various evolutionary processes shape variation in populations. We survey the limits that population genetics places on adaptive evolution, and discuss the relationship between disparate literatures.     

Between beanbag genetics and natural selection

The encounter between the Darwinian theory of evolution and Mendelism could be resolved only when reductionist tools could be applied to the analysis of complex systems. The instrumental reductionist interpretation of the hereditary basis of continuously varying traits provided mathematical tools which eventually allowed the construction of the Modern Synthesis of the theory of evolution.When genotypic as well as environmental variance allow the isolation of parts of the system, it is possible to apply Mendelian reductionism, that is , to treat the phenotypic trait as if ti causally determined by discrete genes for the trait. howeverm such a beanbag genetics approach obscures the system's eye-view. The concept of heritability, defined as the proportion of the total phenotypic variance due to (additive) hereditary variation, asserts that genetic elements have discrete effects; but by relating to the genotypic variance, it avoids the trap of reffering to genes for characters.