Evolutionary ethics,Darwinian ethics and ethical naturalism

The relevance of evolutionary theory to ethics goes back to Darwin but until recently discussion employed evolutionary theory to justify ethical, social and political positions. Recently, evolutionary theory has been used to explain the existence of moral systems and moral propensities and, thereby, to provide a naturalistic basis for ethics. I argue that this approach has advanced our understanding of the basis of moral systems and moral propensities but does not as yet adequately incorporate the role of cognition in its account. Cognition has the effect of decoupling to some extent — though, of course, far from fully — human moral systems from their evolutionary origins. In an adequate account, evolutionary theory will play a crucial role but so also will our evolved cognitive abilities.     

Integrating the Multiple Biological Causes of Human Behavior

I introduce a range of examples of different causal hypotheses about human mate selection. The hypotheses I focus on come from evolutionary psychology, fluctuating asymmetry research and chemical signaling research. I argue that a major obstacle facing an integrated biology of human behavior is the lack of a causal framework that shows how multiple proximate causal mechanisms can act together to produce components of our behavior.     

Ecological frames of mind: the role of cognition in behavioral ecology.

Cognitive psychology is the study of how information, from the senses and from memory, is used in the production of behavior. Investigation of the specifics of behavioral adaptation has already led some behavioral ecologists into the domain of animal cognition. I make several arguments for the benefits and the necessity of a sophisticated assessment by ecologists of the cognitive aspects of behavioral adaptation. First, because cognition typically serves to produce adaptive behavior, cognitive structure and function should reflect ecological demands; studies of cognition in ecological contexts are opportunities to understand adaptation. Furthermore, constraints on cognitive properties may help determine how behavior meets the environment. Studies of spatial memory in food-caching corvids exemplify how cognitive aspects of behavior may both reflect and determine specifics of adaptation. Second, many models in behavioral ecology assume certain cognitive abilities, such as timing or counting. Cognitive theory and methodology should be used to determine whether animals possess these abilities. I have provided examples. Third, consideration of cognitive function can lead to original ideas about the details of behavioral adaptation. Without a thorough integration of cognitive psychology with behavioral ecology, our understanding of the relation between behavior and selective pressures will be compromised.     

Mapping Genetically Controlled Neural Circuits of Social Behavior and Visuo-Motor Integration by a Preliminary Examination of Atypical Deletions with Williams Syndrome

In this study of eight rare atypical deletion cases with Williams-Beuren syndrome (WS; also known as 7q11.23 deletion syndrome) consisting of three different patterns of deletions, compared to typical WS and typically developing (TD) individuals, we show preliminary evidence of dissociable genetic contributions to brain structure and human cognition. Univariate and multivariate pattern classification results of morphometric brain patterns complemented by behavior implicate a possible role for the chromosomal region that includes: 1) GTF2I/GTF2IRD1 in visuo-spatial/motor integration, intraparietal as well as overall gray matter structures, 2) the region spanning ABHD11 through RFC2 including LIMK1, in social cognition, in particular approachability, as well as orbitofrontal, amygdala and fusiform anatomy, and 3) the regions including STX1A, and/or CYLN2 in overall white matter structure. This knowledge contributes to our understanding of the role of genetics on human brain structure, cognition and pathophysiology of altered cognition in WS. The current study builds on ongoing research designed to characterize the impact of multiple genes, gene-gene interactions and changes in gene expression on the human brain.     

Mental content and evolutionary explanation

Cognitive ethology is the comparative study of animal cognition from an evolutionary perspective. As a sub-discipline of biology it shares interest in questions concerning the immediate causes and development of behavior. As a part of ethology it is also concerned with questions about the function and evolution of behavior. I examine some recent work in cognitive ethology, and I argue that the notions of mental content and representation are important to enable researchers to answer questions and state generalizations about the function and volution of behavior.I would like to thank Dorothy Cheney, Keith Donnellan, Alan Nelson, and Robert Seyfarth for their help.     

Oxytocin,testosterone, and human social cognition

I describe an integrative social‐evolutionary model for the adaptive significance of the human oxytocinergic system. The model is based on a role for this hormone in the generation and maintenance of social familiarity and affiliation across five homologous, functionally similar, and sequentially co‐opted contexts: mothers with offspring, female and male mates, kin groups, individuals with reciprocity partners, and individuals within cooperating and competing social groups defined by culture. In each situation, oxytocin motivates, mediates and rewards the cognitive and behavioural processes that underlie the formation and dynamics of a more or less stable social group, and promotes a relationship between two or more individuals. Such relationships may be positive (eliciting neurological reward, reducing anxiety and thus indicating fitness‐enhancing effects), or negative (increasing anxiety and distress, and thus motivating attempts to alleviate a problematic, fitness‐reducing social situation). I also present evidence that testosterone exhibits opposite effects from oxytocin on diverse aspects of cognition and behaviour, most generally by favouring self‐oriented, asocial and antisocial behaviours. I apply this model for effects of oxytocin and testosterone to understanding human psychological disorders centrally involving social behaviour. Reduced oxytocin and higher testosterone levels have been associated with under‐developed social cognition, especially in autism. By contrast, some combination of oxytocin increased above normal levels, and lower testosterone, has been reported in a notable number of studies of schizophrenia, bipolar disorder and depression, and, in some cases, higher oxytocin involves maladaptively ‘hyper‐developed’ social cognition in these conditions. This pattern of findings suggests that human social cognition and behaviour are structured, in part, by joint and opposing effects of oxytocin and testosterone, and that extremes of such joint effects partially mediate risks and phenotypes of autism and psychotic‐affective conditions. These considerations have direct implications for the development of therapies for alleviating disorders of social cognition, and for understanding how such disorders are associated with the evolution of human cognitive‐affective architecture.     

Why there are no objective values: A critique of ethical intuitionism from an evolutionary point of view

Using concepts of evolutionary game theory, this paper presents a critique of ethical intuitionism, or non-naturalism, in its cognitivist and objectivist interpretation. While epistemological considerations suggest that human rational learning through experience provides no basis for objective moral knowledge, it is argued below that modern evolutionary theory explains why this is so, i.e., why biological organisms do not evolve so as to experience objective preferences and obligations. The difference between the modes of the cognition of objective and of valuative environmental attributes is explained with reference to different modes of natural selection acting on the cognitive apparatus of the organism. The negative implications are pointed out which the observable diversity of intraspecific behavioural adaptations and of cultural values has for the cognitivist, objectivist foundation of ethics. Eventually a non-cognitivist alternative to ethical intuitionism is outlined in terms of empirical authority relations, with the ritualisation of dominance-submission patterns as the evolutionary origin of human charismatic authority.     

The biological roots of morality

The question whether ethical behavior is biologically determined may refer either to thecapacity for ethics (e.i., the proclivity to judge human actions as either right or wrong), or to the moralnorms accepted by human beings for guiding their actions. My theses are: (1) that the capacity for ethics is a necessary attribute of human nature; and (2) that moral norms are products of cultural evolution, not of biological evolution.Humans exhibits ethical behavior by nature because their biological makeup determines the presence of the three necessary, and jointly sufficient, conditions for ethical behavior: (i) the ability to anticipate the consequences of one's own actions; (ii) the ability to make value judgements; and (iii) the ability to choose between alternative courses of action. Ethical behavior came about in evolution not because it is adaptive in itself, but as a necessary consequece of man's eminent intellectual abilities, which are an attribute directly promoted by natural selection.Since Darwin's time there have been evolutionists proposing that the norms of morality are derived from biological evolution. Sociobiologists represent the most recent and most subtle version of that proposal. The sociobiologists' argument is that human ethical norms are sociocultural correlates of behaviors fostered by biological evolution. I argue that such proposals are misguided and do not escape the naturalistic fallacy. The isomorphism between the behaviors promoted by natural selection and those sanctioned by moral norms exist only with respect to the consequences of the behaviors; the underlying causations are completely disparate.This article is based on a paper presented at the International Symposium onBiological Models of Human Action, Palma de Mallorca, Spain, 16–18 December 1985.     

Human Perceptual and Phobic Biases for Snakes: A Review of the Experimental Evidence

In this paper I review the literature on the evolutionary origins of phobias and describe the current state of research on the neurobiology and developmental origins of ophidiophobia—fear of snakes. In doing so I compare experimental evidence related to evolutionary explanations for snake fears and phobias which are outlined in Seligman’s Preparedness Theory and Isbell’s Snake Detection Theory. These theories have been tested extensively using a variety of experimental paradigms aimed at determining the “innateness” of snake fears, the neural pathways involved in fear responses to snakes, and the perceptual biases associated with snake stimuli. However, in the vast majority of these experiments, the stimuli presented are photographs of snakes rather than the real thing. I argue that this point of methodology, while ironically supportive of the findings, is based on some assumptions about cognition and consciousness which run counter to neuroscience. In understanding human responses to snakes, we need to understand better the interplay between cognition and consciousness and how these represent a pluralism of mind in which perception is much more than we think.     

The acquisition of conscience and Developmental Systems Theory

Summary Proponents of Developmental Systems Theory (DST) argue that it offers an alternative to current research programs in biology that are built on the historic disjunction between evolutionary and developmental biology. In this paper I illustrate how DST can be used to account for the acquisition of an important component of moral agency, conscience. Susan Oyama, a major proponent of DST, has set moral issues outside the compass of DST. Thus, I examine her reasons for restricting DST to non-moral matters, and argue that they are not decisive. On the positive side, I argue that DST not only is compatible with attempts to describe and explain moral agency but also aids us in understanding it. In particular, I show how DST can provide a fruitful perspective for viewing some significant current findings and theories in moral developmental psychology about the acquisition of conscience. The familiar dichotomies resisted by DST, those between genes and environment, inherited and acquired, innate and learned, and biological and cultural, have also plagued human developmental psychology, including moral development. By bringing a DST perspective to the study of moral development, I illustrate how a DST perspective might offer a promising way to reconceive that phenomenon, and provide some insights into how further work in understanding the development of moral agency might proceed. Thus, I hope to contribute to the current efforts of proponents of DST to integrate developmental and evolutionary considerations.     

Spatial memory tasks in rodents: what do they model?

The analysis of spatial learning and memory in rodents is commonly used to investigate the mechanisms underlying certain forms of human cognition and to model their dysfunction in neuropsychiatric and neurodegenerative diseases. Proper interpretation of rodent behavior in terms of spatial memory and as a model of human cognitive functions is only possible if various navigation strategies and factors controlling the performance of the animal in a spatial task are taken into consideration. The aim of this review is to describe the experimental approaches that are being used for the study of spatial memory in rats and mice and the way that they can be interpreted in terms of general memory functions. After an introduction to the classification of memory into various categories and respective underlying neuroanatomical substrates, I explain the concept of spatial memory and its measurement in rats and mice by analysis of their navigation strategies. Subsequently, I describe the most common paradigms for spatial memory assessment with specific focus on methodological issues relevant for the correct interpretation of the results in terms of cognitive function. Finally, I present recent advances in the use of spatial memory tasks to investigate episodic-like memory in mice.     

The evolution of linguistic rules

Rule-like behaviour is found throughout human language, provoking a number of apparently conflicting explanations. This paper frames the topic in terms of Tinbergen’s four questions and works within the context of rule-like behaviour seen both in nature and the non-linguistic domain in humans. I argue for a minimal account of linguistic rules which relies on powerful domain-general cognition, has a communicative function allowing for multiple engineering solutions, and evolves mainly culturally, while leaving the door open for some genetic adaptation in the form of learning biases.     

Perspectives: The Looking Time Experimental Paradigm in Studies of Animal Visual Perception and Cognition

Experiments are the foundation of empirical science, and experimental paradigms that are broadly applicable across settings and species are particularly useful for comparative research. Originally developed to address questions related to perception and cognition of pre‐verbal human infants, the looking time experimental paradigm has been increasingly used to study animal behavior and cognition, particularly in non‐human primates. Looking time experiments are based on the assumption that animals direct eye gaze toward objects or scenes based on their degree of interest, and use looking behavior to infer perceptual or cognitive characteristics of subjects. This paradigm can be used in a variety of contexts and is not based on species‐typical behaviors, allowing for intra‐ and interspecific comparisons. Here, we describe the history of use of looking time measures, provide an overview of the problems and controversies related to this method, and offer recommendations on how to implement looking time tasks, focusing on the preparation of stimuli, experimental procedures, and data analysis. Our overview focuses on non‐human primates, where most work has been carried out, but the issues discussed should be applicable to a wide range of species. We conclude that despite pertinent criticism, looking time tasks are practical when executed and interpreted properly. The further implementation of these methods in studies of animal behavior and cognition is likely to be fruitful.     

Quantum-like behavior without quantum physics I

Recently there has been much interest in the possible quantum-like behavior of the human brain in such functions as cognition, the mental lexicon, memory, etc., producing a vast literature. These studies are both empirical and theoretical, the tenets of the theory in question being mainly, and apparently inevitably, those of quantum physics itself, for lack of other arenas in which quantum-like properties are presumed to obtain. However, attempts to explain this behavior on the basis of actual quantum physics going on at the atomic or molecular level within some element of brain or neuronal anatomy (other than the ordinary quantum physics that underlies everything), do not seem to survive much scrutiny. Moreover, it has been found empirically that the usual physics-like Hilbert space model seems not to apply in detail to human cognition in the large. In this paper we lay the groundwork for a theory that might explain the provenance of quantum-like behavior in complex systems whose internal structure is essentially hidden or inaccessible. The approach is via the logic obeyed by these systems which is similar to, but not identical with, the logic obeyed by actual quantum systems. The results reveal certain effects in such systems which, though quantum-like, are not identical to the kinds of quantum effects found in physics. These effects increase with the size of the system.     

The Tsimane Health and Life History Project: Integrating anthropology and biomedicine

The Tsimane Health and Life History Project, an integrated bio‐behavioral study of the human life course, is designed to test competing hypotheses of human life‐history evolution. One aim is to understand the bidirectional connections between life history and social behavior in a high‐fertility, kin‐based context lacking amenities of modern urban life (e.g. sanitation, banks, electricity). Another aim is to understand how a high pathogen burden influences health and well‐being during development and adulthood. A third aim addresses how modernization shapes human life histories and sociality. Here we outline the project's goals, history, and main findings since its inception in 2002. We reflect on the implications of current findings and highlight the need for more coordinated ethnographic and biomedical study of contemporary nonindustrial populations to address broad questions that can situate evolutionary anthropology in a key position within the social and life sciences.     

Genes, language, cognition, and culture: towards productive inquiry

The Queen Mary conference on “Integrating Genetic and Cultural Evolutionary Approaches to Language,” and the papers in this special issue, clearly illustrate the excitement and potential of trans-disciplinary approaches to language as an evolved biological capacity (phylogeny) and an evolving cultural entity (glossogeny). Excepting the present author, the presenters/authors are mostly young rising stars in their respective fields, and include scientists with backgrounds in linguistics, animal communication, neuroscience, evolutionary biology, anthropology, and computer science. On display was a clear willingness to engage with different approaches and terminology and a commitment to shared standards of scientific rigor, empirically driven theory, and logical argument. Because the papers assembled here, together with the introduction, speak for themselves, I will focus in this “extro-duction” on some of the terminological and conceptual difficulties which threaten to block this exciting wave of scientific progress in understanding language evolution, in both senses of that term. In particular I will first argue against the regrettably widespread practice of opposing cultural and genetic explanations of human cognition as if they were dichotomous. Second, I will unpack the debate concerning “general-purpose” and “domain-specific” mechanisms, which masquerades as a debate about nativism but is nothing of the sort. I believe that framing discussions of language in these terms has generated more heat than light, and that a modern molecular understanding of genes, development, behavior, and evolution renders many of the assumptions underlying this debate invalid.     

The computer program STRUCTURE does not reliably identify the main genetic clusters within species: simulations and implications for human population structure

One of the primary goals of population genetics is to succinctly describe genetic relationships among populations, and the computer program STRUCTURE is one of the most frequently used tools for doing so. The mathematical model used by STRUCTURE was designed to sort individuals into Hardy–Weinberg populations, but the program is also frequently used to group individuals from a large number of populations into a small number of clusters that are supposed to represent the main genetic divisions within species. In this study, I used computer simulations to examine how well STRUCTURE accomplishes this latter task. Simulations of populations that had a simple hierarchical history of fragmentation showed that when there were relatively long divergence times within evolutionary lineages, the clusters created by STRUCTURE were frequently not consistent with the evolutionary history of the populations. These difficulties can be attributed to forcing STRUCTURE to place individuals into too few clusters. Simulations also showed that the clusters produced by STRUCTURE can be strongly influenced by variation in sample size. In some circumstances, STRUCTURE simply put all of the individuals from the largest sample in the same cluster. A reanalysis of human population structure suggests that the problems I identified with STRUCTURE in simulations may have obscured relationships among human populations—particularly genetic similarity between Europeans and some African populations.     

It's all in your head: the role of quantity estimation in sperm competition

The study of animal cognition has provided valuable data throughout the years, yet its reliance on laboratory work leaves some open questions. The main question is whether animals employ cognition in daily decision-making. The following discussion uses sperm competition (SC) as a test case for demonstrating the effect of cognition on routine choices, in this case, sexual selection. Cognition is manifested here by males' ability to represent the number of rivals competing with them. I claim that response to SC is driven by quantity estimation and the ability to assess competition magnitude cognitively. Hence, cognition can determine males' response to SC, and consequentially it can be selected within this context. This supports the argument that cognition constitutes an integral part of an individual's toolbox in solving real-life problems, and shows that physical and behavioural phenomena can expose cognition to selection and facilitate its evolution.