Losing behavior (yielding subroutine) and human depression: Proximate and selective mechanisms

In this article, we present what we believe to be some of the proximate and selective mechanisms that allowed certain types of human depression to have evolved. Certain adult depressive patients, in losing situations, have an inability to “switch off” their submissive or yielding behavior with its concomitant cognitive and emotional components. Thus, although normal individuals accept defeat as an inevitable aspect of ritual agonistic encounters, certain depressed individuals do not.We believe the failure to terminate the yielding subroutine of ritual agonistic behavior (YS/RAB) is the proximate mechanism that underlies certain types of human depression. Also, we show that through the selective mechanisms of difference amplification and kin selection, certain types of depressive illness may be adaptive to kin. Thus, selection could have operated at the family as well as the individual level, permitting certain types of depression to have evolved. The notion that depression is the result of selective mechanisms is supported by the prevalence of depression in human populations.     

Evolutionary psychology and the generation of culture,part I: Theoretical considerations

Culture is the ongoing product of the evolved psyches of individual humans living in groups. Progress in our understanding of culture as a phenomenon depends on progress in uncovering the nature of the evolved mechanisms that comprise the human psyche, including but not limited to those responsible for learning. Actual attemps to specify information processing mechanisms that could, in fact, perform tasks humans routinely perform have demonstrated that the human psyche cannot, even in principle, be comprised only of a general purpose learning mechanism or any other general purpose mechanism, such as an inclusive fitness maximizer. Instead, the human psyche appears to consist of a large number of mechanisms, many or most of which are special purpose and domain-specific. The output of these mechanisms taken together constitutes the “private culture” of each individual, and the interactions of these private cultures lead to the cross-individual patterns of similarity that have led anthropologists to think typologically of social groups as having “a” culture. The construction of a scientific theory of culture requires as its building blocks specific models of these psychological mechanisms, and so evolutionary anthropology depends on the forging of an evolutionary psychology. The most productive application of evolutionary biology is, therefore, in the study of the psychological mechanisms that generate and shape culture, rather than in the attempt to impose on cultural change too close a parallel to population genetics and organic evolution.     

Endocrinology of human female sexuality,mating, and reproductive behavior

Hormones orchestrate and coordinate human female sexual development, sexuality, and reproduction in relation to three types of phenotypic changes: life history transitions such as puberty and childbirth, responses to contextual factors such as caloric intake and stress, and cyclical patterns such as the ovulatory cycle. Here, we review the endocrinology underlying women's reproductive phenotypes, including sexual orientation and gender identity, mate preferences, competition for mates, sex drive, and maternal behavior. We highlight distinctive aspects of women's sexuality such as the possession of sexual ornaments, relatively cryptic fertile windows, extended sexual behavior across the ovulatory cycle, and a period of midlife reproductive senescence—and we focus on how hormonal mechanisms were shaped by selection to produce adaptive outcomes. We conclude with suggestions for future research to elucidate how hormonal mechanisms subserve women's reproductive phenotypes.     

Evolutionary biology and psychiatry: The case of anorexia nervosa

A review of the literature and an analysis of 19 case histories suggest that evolutionary mechanisms such as reproductive suppression, kin selection, and parental manipulation are involved in the development of anorexia nervosa. From this perspective, anorexia nervosa is viewed as—in some cases—an adaptive feature of human reproductive strategies and may be best understood as an emergency strategy that is pursued whenever the sovereign handling of one's own reproductive potential is not possible because of socioecological or ontogenetic constraints.     

Human evolution: humanistic selection and looking to the future

Cultural evolution has predominated over biological evolution in modern man (Homo sapiens sapiens). Cultural evolution differs from biological evolution not only by inheritance of acquired characteristics but also, as is proposed in the present essay, by another kind of selection mechanism. Whereas selection in biological evolution is executed according to a criterion of reproductive success (the natural selection), selection in cultural evolution appears to be carried out according to human and humanistic criteria (success or fitness in meeting human needs, interests and humanistic values--"humanistic selection"). Many humanistic needs or values do not seem to be prerequisite for reproductive success, yet some of them (e.g. a need for freedom) seem to be inborn. Innateness, humanistic selection (decisive at a community level) and hierarchy of some human needs, interests and values appear to give cultural evolution a generally upward trend although long periods of stagnation or even regression may occur. Modern humans appear to be still at the early stage of their cultural evolution. A further cultural evolution of man appears to be, in contrast to biological evolution, predictable (with an optimistic outlook) and testable. The problem is that the hopeful result of this test will probably be known only in the fairly remote future provided that this species will not become extinct before that.     

Which humans behave adaptively,and why does it matter?

There has long been debate about the relevance of evolutionary theory to the study of humans. To many of us, however, the debate has shifted from whether to proceed with an evolutionary approach to how to proceed. Increasingly, it has been argued that studies of the current reproductive function of human traits make little or no contribution to the understanding of the psyche (e.g., Symons 1989). Here, on the basis of arguments about the relationship between an adaptation and an adaptive outcome, and a review of studies that assess current adaptiveness, I argue to the contrary that knowledge of the contexts in which people do or do not behave adaptively provides important information about the nature of the mechanisms that comprise the human psyche. In particular, studies that indicate that people behave adaptively in at least some contemporary environments cast doubt on many nonevolutionary constructions of human nature, and can be used now to distinguish alternative evolutionary constructions that are at odds over many issues pentaining to the human psyche's ontogeny and evolutionary background, especially the extent to which the human psyche is general purpose.     

Interactions between the sexes: new perspectives on sexual selection and reproductive isolation

Understanding the processes underlying the origin of new species is a fundamental problem in evolutionary research. Whilst it has long been recognised that closely related taxa often differ markedly in reproductive characteristics, only relatively recently has sexual selection been evoked as a key promoter of speciation through its ability to generate reproductive isolation (RI). Sexual selection potentially can influence the probability that individuals from the same or different populations will reproduce successfully since it shapes precisely those traits involved in mating and reproduction. If reproductive characters diverge along different trajectories, then sexual selection can impact on the evolution of reproductive barriers operating both before and after mating. In this perspective, we consider some new advances in our understanding of the coevolution of male and female sexual signals and receptors and suggest how these developments may provide heretofore neglected insights into the mechanisms by which isolating barriers may emerge. Specifically, we explore how selfish genetic elements (SGEs) can mediate pre- and post-copulatory mate choice, thereby influencing gene flow and ultimately population divergence; we examine evidence from studies of intracellular sperm–egg interactions and propose that intracellular gametic incompatibilities may arise after sperm entry into the egg, and thus contribute to RI; we review findings from genomic studies demonstrating rapid, adaptive evolution of reproductive genes in both sexes and discuss whether such changes are causal in determining RI or simply associated with it; and finally, we consider genetic, developmental and functional mechanisms that might constrain reproductive trait diversification, thereby limiting the scope for reproductive barriers to arise via sexual selection. We hope to stimulate work that will further the understanding of the role sexual selection plays in generating RI and ultimately speciation.     

The adaptive significance of cultural behavior

In this article, I argue that human social behavior is a product of the coevolution of human biology and culture. While critical of attempts by anthropologists to explain cultural practices as if they were independent of the ability of individual human beings to survive and reproduce, I am also leery of attempts by biologists to explain the consistencies between neo-Darwinian theory and cultural behavior as the result of natural selection for that behavior. Instead, I propose that both biological and cultural attributes of human beings result to a large degree from the selective retention of traits that enhance the inclusive fitnesses of individuals in their environments. Aspects of human biology and culture may be adaptive in the same sense despite differences between the mechanisms of selection and regardless of their relative importance in the evolution of a trait. The old idea that organic and cultural evolution are complementary can thus be used to provide new explanations for why people do what they do.This article was prepared from remarks I presented at the Smithsonian Conference on Human Biogeography in April 1974. Help with phenotypic costs came from an NSF Pre-doctoral Fellowship and the University of Michigan Society of Fellows.     

The evolution of partial reproductive isolation as an adaptive optimum

Decades of theoretical work on the evolution of adaptive prezygotic isolation have led to an interesting finding—namely that stable partial reproductive isolation is a relatively common outcome. This conclusion is generally lost, however, in the desire to pinpoint when exactly speciation occurs. Here, we argue that the evolution of partial reproductive isolation is of great interest in its own right and matches empirical findings that ongoing hybridization is taxonomically widespread. We present the mechanisms by which partial reproductive isolation can be a stable evolutionary endpoint, concentrating on insights from theoretical studies. We focus not on cases in which hybridization results from constraints imposed by ongoing migration or mutation, but on the intriguing idea that partial reproductive isolation may instead be an adaptive optimum. We identify three general categories of selective mechanisms that can lead to partial reproductive isolation: context-dependent hybrid advantage, indirect selection due to the varying actions of sexual selection in different geographic contexts, and a balance of costs of choosiness with indirect selection for stronger mating preferences. By any of these mechanisms, stable partial reproductive isolation can potentially provide a robust evolutionary alternative to either complete speciation or population fusion.     

Does Biology Constrain Culture?

Most social scientists would agree that the capacity for human culture was probably fashioned by natural selection, but they disagree about the implications of this supposition. Some believe that natural selection imposes important constraints on the ways in which culture can vary, while others believe that any such constraints must be negligible. This article employs a "thought experiment" to demonstrate that neither of these positions can be justified by appeal to general properties of culture or of evolution. Natural selection can produce mechanisms of cultural transmission that are neither adaptive nor consistent with the predictions of acultural evolutionary models (those ignoring cultural evolution). On the other hand, natural selection can also produce mechanisms of cultural transmission that are highly consistent with acultural models. Thus, neither side of the sociobiology debate is justified in dismissing the arguments of the other. Natural selection may impose significant constraints on some human behaviors, but negligible constraints on others. Models of simultaneous genetic/cultural evolution will be useful in identifying domains in which acultural evolutionary models are, and are not, likely to be useful.     

Genetic architecture and balancing selection: the life and death of differentiated variants

Balancing selection describes any form of natural selection, which results in the persistence of multiple variants of a trait at intermediate frequencies within populations. By offering up a snapshot of multiple co‐occurring functional variants and their interactions, systems under balancing selection can reveal the evolutionary mechanisms favouring the emergence and persistence of adaptive variation in natural populations. We here focus on the mechanisms by which several functional variants for a given trait can arise, a process typically requiring multiple epistatic mutations. We highlight how balancing selection can favour specific features in the genetic architecture and review the evolutionary and molecular mechanisms shaping this architecture. First, balancing selection affects the number of loci underlying differentiated traits and their respective effects. Control by one or few loci favours the persistence of differentiated functional variants by limiting intergenic recombination, or its impact, and may sometimes lead to the evolution of supergenes. Chromosomal rearrangements, particularly inversions, preventing adaptive combinations from being dissociated are increasingly being noted as features of such systems. Similarly, due to the frequency of heterozygotes maintained by balancing selection, dominance may be a key property of adaptive variants. High heterozygosity and limited recombination also influence associated genetic load, as linked recessive deleterious mutations may be sheltered. The capture of deleterious elements in a locus under balancing selection may reinforce polymorphism by further promoting heterozygotes. Finally, according to recent genomewide scans, balanced polymorphism might be more pervasive than generally thought. We stress the need for both functional and ecological studies to characterize the evolutionary mechanisms operating in these systems.     

A critique of Darwinian anthropology

Darwinian anthropology holds that human behavior is adaptive in the sense of being designed to maximize reproductive success and that measurement of reproductive differentials typically illuminates adaptation. I argue to the contrary, that adaptive design is usually manifested at the psychological rather than at the behavioral level, that measuring reproductive differentials is at best an inefficient and ambiguous way to illuminate adaptation, and that Darwin's theory of natural selection sheds light on human affairs only insofar as it promotes understanding of the psychology that underpins these affairs.     

Interethnic Interaction,Strategic Bargaining Power,and the Dynamics of Cultural Norms

Ethnic groups are universal and unique to human societies. Such groups sometimes have norms of behavior that are adaptively linked to their social and ecological circumstances, and ethnic boundaries may function to protect that variation from erosion by interethnic interaction. However, such interaction is often frequent and voluntary, suggesting that individuals may be able to strategically reduce its costs, allowing adaptive cultural variation to persist in spite of interaction with out-groups with different norms. We examine five mechanisms influencing the dynamics of ethnically distinct cultural norms, each focused on strategic individual-level choices in interethnic interaction: bargaining, interaction-frequency-biased norm adoption, assortment on norms, success-biased interethnic social learning, and childhood socialization. We use Bayesian item response models to analyze patterns of norm variation and interethnic interaction in an ethnically structured Amazonian population. We show that, among indigenous Matsigenka, interethnic education with colonial Mestizos is more strongly associated with Mestizo-typical norms than even extensive interethnic experience in commerce and wage labor is. Using ethnographic observations, we show that all five of the proposed mechanisms of norm adoption may contribute to this effect. However, of these mechanisms, we argue that changes in relative bargaining power are particularly important for ethnic minorities wishing to preserve distinctive norms while engaging in interethnic interaction in domains such as education. If this mechanism proves applicable in a range of other ethnographic contexts, it would constitute one cogent explanation for when and why ethnically structured cultural variation can either persist or erode given frequent, and often mutually beneficial, interethnic interaction.     

The elastic between genes and culture

A society's culture may come to include information tending to lead to fitness-reducing behavior on the part of some or all of its members. This phenomenon results from conflict among factions within each society, from transmitted misinformation (e.g., cupping restores health), from natural and human-caused environmental change so that previously adaptive information becomes maladaptive, and from the long and short-term negative “side effects” of information that may otherwise be fitness enhancing. Because some cultural information may be fitness reducing, we apparently have been selected for individual-level traits that often result in our revising socially transmitted information that might otherwise have maladaptive consequences. Two examples of such traits are adolescent “rebelliousness” and the tendency to learn most readily from those higher than ourselves in status. Such leading-to-culture-revision traits are very imperfect mechanisms, however, so that some likely-to-be maladaptive cultural information, such as medical cupping or denying infants the colostrum, remains part of the culture. It is doubtful, given the structure of modern human populations and the ubiquity of culture change, that such maladaptive socially transmitted information leads to natural selection for genetic “direct biases” against accepting the practices in question.     

The on‐again,off‐again relationship between mitochondrial genomes and species boundaries

The study of reproductive isolation and species barriers frequently focuses on mitochondrial genomes and has produced two alternative and almost diametrically opposed narratives. On one hand, mtDNA may be at the forefront of speciation events, with co‐evolved mitonuclear interactions responsible for some of the earliest genetic incompatibilities arising among isolated populations. On the other hand, there are numerous cases of introgression of mtDNA across species boundaries even when nuclear gene flow is restricted. We argue that these seemingly contradictory patterns can result from a single underlying cause. Specifically, the accumulation of deleterious mutations in mtDNA creates a problem with two alternative evolutionary solutions. In some cases, compensatory or epistatic changes in the nuclear genome may ameliorate the effects of mitochondrial mutations, thereby establishing coadapted mitonuclear genotypes within populations and forming the basis of reproductive incompatibilities between populations. Alternatively, populations with high mitochondrial mutation loads may be rescued by replacement with a more fit, foreign mitochondrial haplotype. Coupled with many nonadaptive mechanisms of introgression that can preferentially affect cytoplasmic genomes, this form of adaptive introgression may contribute to the widespread discordance between mitochondrial and nuclear genealogies. Here, we review recent advances related to mitochondrial introgression and mitonuclear incompatibilities, including the potential for cointrogression of mtDNA and interacting nuclear genes. We also address an emerging controversy over the classic assumption that selection on mitochondrial genomes is inefficient and discuss the mechanisms that lead lineages down alternative evolutionary paths in response to mitochondrial mutation accumulation.     

How mechanisms of habitat preference evolve and promote divergence with gene flow

Habitat preference may promote adaptive divergence and speciation, yet the conditions under which this is likely are insufficiently explored. We use individual‐based simulations to study the evolution and consequence of habitat preference during divergence with gene flow, considering four different underlying genetically based behavioural mechanisms: natal habitat imprinting, phenotype‐dependent, competition‐dependent and direct genetic habitat preference. We find that the evolution of habitat preference generally requires initially high dispersal, is facilitated by asymmetry in population sizes between habitats, and is hindered by an increasing number of underlying genetic loci. Moreover, the probability of habitat preference to emerge and promote divergence differs greatly among the underlying mechanisms. Natal habitat imprinting evolves most easily and can allow full divergence in parameter ranges where no divergence is possible in the absence of habitat preference. The reason is that imprinting represents a one‐allele mechanism of assortative mating linking dispersal behaviour very effectively to local selection. At the other extreme, direct genetic habitat preference, a two‐allele mechanism, evolves under restricted conditions only, and even then facilitates divergence weakly. Overall, our results indicate that habitat preference can be a strong reproductive barrier promoting divergence with gene flow, but that this is highly contingent on the underlying preference mechanism.     

Alternative developmental pathways and the propensity to migrate: a case study in the Atlantic salmon

Migratory behaviour with its associated phenotypic changes is generally viewed as an adaptive strategy because it incurs survival or reproductive advantages to migrants. The development of a migrant phenotype is believed to be controlled by threshold mechanisms, where individuals emigrate only after surpassing a particular body size but delay migration if below. For such a strategy to respond to natural selection, part of the phenotypic variance in the propensity to migrate must be explained by variation in additive genetic effects. Here, we use data gathered in the field and from a common rearing experiment to test for a genetic basis associated with seaward migration in Atlantic salmon (Salmo salar L.). We document a high heritability of the liability trait underlying the propensity to emigrate in juvenile salmon, and significant differences between offspring grouped according to their sires in body-size threshold values above which emigration takes place. The presence of additive genetic variance in both the liability and thresholds makes the onset of migration a process sensitive to selection and may therefore constitute an important explanatory mechanism for the interpopulation differences in the size at seaward migration observed in this species.     

Evolutionary psychology and the generation of culture,part II: Case study: A computational theory of social exchange

Models of the various adaptive specializations that have evolved in the human psyche could become the building blocks of a scientific theory of culture. The first step in creating such models is the derivation of a so-called “computational theory” of the adaptive problem each psychological specialization has evolved to solve. In Part II, as a case study, a sketch of a computational theory of social exchange (cooperation for mutual benefit) is developed. The dynamics of natural selection in Pleistocene ecological conditions define adaptive information processing problems that humans must be able to solve in order to participate in social exchange: individual recognition, memory for one's history of interaction, value communication, value modeling, and a shared grammar of social contracts that specifies representational structure and inferential procedures. The nature of these adaptive information processing problems places constraints on the class of cognitive programs capable of solving them; this allows one to make empirical predictions about how the cognitive processes involved in attention, communication, memory, learning, and reasoning are mobilized in situations of social exchange. Once the cognitive programs specialized for regulating social exchange are mapped, the variation and invariances in social exchange within and between cultures can be meaningfully discussed.     

Disruptive selection in a bimodal population of Darwin's finches

A key part of the ecological theory of adaptive radiation is disruptive selection during periods of sympatry. Some insight into this process might be gained by studying populations that are bimodal for dual-context traits, i.e. those showing adaptive divergence and also contributing to reproductive isolation. A population meeting these criteria is the medium ground finch (Geospiza fortis) of El Garrapatero, Santa Cruz Island, Galápagos. We examined patterns of selection in this population by relating individual beak sizes to interannual recaptures during a prolonged drought. Supporting the theory, disruptive selection was strong between the two beak size modes. We also found some evidence of selection against individuals with the largest and smallest beak sizes, perhaps owing to competition with other species or to gaps in the underlying resource distribution. Selection may thus simultaneously maintain the current bimodality while also constraining further divergence. Spatial and temporal variation in G. fortis bimodality suggests a dynamic tug of war among factors such as selection and assortative mating, which may alternatively promote or constrain divergence during adaptive radiation.     

社群学习对植食性鸟类和哺乳动物觅食行为的作用

社群学习是动物的一种可塑性行为表现型式。综述了社群学习对植食性鸟类和哺乳动物觅食行为的作用,并述评了其学习机制。社群同伴对动物个体觅食地点、时间和取食方式均有影响,母体摄食的食物信息可通过胎盘和乳汁显著影响幼体的食物选择。动物通过观察学习、嗅闻学习以及味觉厌恶学习,不仅能更快找到食物资源,提高觅食效率,而且能有效降低中毒与被捕食的风险,从而提高其适合度。