Wikipedia book of Sociobiology

The Wikipedia Book of Sociobiology

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Contents Articles Sociobiology

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Biocultural anthropology

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Biosocial theory

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Cultural selection theory

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Darwinian anthropology

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Dual inheritance theory

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Evolutionary ethics

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Evolutionary anthropology

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Evolutionary biology

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Evolutionary developmental psychology

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Evolutionary neuroscience

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Evolutionary psychology

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Human behavioral ecology

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Prisoner's dilemma

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Kin selection

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Social evolution

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Social neuroscience

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Sociobiological theories of rape

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Sociophysiology

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E. O. Wilson

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W. D. Hamilton

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Robert Trivers

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George C. Williams

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Sarah Blaffer Hrdy

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Richard Machalek

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Steven Pinker

136

Frans de Waal

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Richard Lewontin

148

Steven Rose

153

Leon Kamin

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Stephen Jay Gould

157

References Article Sources and Contributors

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Image Sources, Licenses and Contributors

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Article Licenses License

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Sociobiology

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Sociobiology Part of a series on

Evolutionary biology

Diagrammatic representation of the divergence of modern taxonomic groups from their common ancestor.

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Sociobiology is a field of scientific study which is based on the assumption that social behavior has resulted from evolution and attempts to explain and examine social behavior within that context. Often considered a branch of biology and sociology, it also draws from ethology, anthropology, evolution, zoology, archaeology, population genetics, and other disciplines. Within the study of human societies, sociobiology is very closely allied to the fields of Darwinian anthropology, human behavioral ecology and evolutionary psychology. Sociobiology investigates social behaviors, such as mating patterns, territorial fights, pack hunting, and the hive society of social insects. It argues that just as selection pressure led to animals evolving useful ways of interacting with the natural environment, it led to the genetic evolution of advantageous social behavior. While the term "sociobiology" can be traced to the 1940s, the concept didn't gain major recognition until 1975 with the publication of Edward O. Wilson's book, Sociobiology: The New Synthesis. The new field quickly became the subject of heated controversy. Criticism, most notably made by Richard Lewontin and Stephen Jay Gould, centered on sociobiology's contention that genes play an ultimate role in human behavior and that traits such as aggressiveness can be explained by biology rather than a person's social environment. Sociobiologists generally responded to the criticism by pointing to the complex relationship between nature and nurture. In response to some of the potentially factious implications sociobiology had for human biodiversity[citation needed], anthropologist John Tooby and psychologist Leda Cosmides founded the field of evolutionary psychology.

Sociobiology

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Definition E.O Wilson defines sociobiology as: “The extension of population biology and evolutionary theory to social organization”[1] Sociobiology is based on the premise that some behaviors (both social and individual) are at least partly inherited and can be affected by natural selection. It begins with the idea that behaviors have evolved over time, similar to the way that physical traits are thought to have evolved. It predicts, therefore, that animals will act in ways that have proven to be evolutionarily successful over time. This can, among other things, result in the formation of complex social processes conducive to evolutionary fitness. The discipline seeks to explain behavior as a product of natural selection. Behavior is therefore seen as an effort to preserve one's genes in the population. Inherent in sociobiological reasoning is the idea that certain genes or gene combinations that influence particular behavioral traits can be inherited from generation to generation.[citation needed]

Introductory example For example, newly dominant male lions often will kill cubs in the pride that were not sired by them. This behaviour is adaptive in evolutionary terms because killing the cubs eliminates competition for their own offspring and causes the nursing females to come into heat faster, thus allowing more of his genes to enter into the population. Sociobiologists would view this instinctual cub-killing behavior as being inherited through the genes of successfully reproducing male lions, whereas non-killing behaviour may have "died out" as those lions were less successful in reproducing.

Support for premise Genetic mouse mutants have now been harnessed to illustrate the power that genes exert on behaviour. For example, the transcription factor FEV (aka Pet1) has been shown, through its role in maintaining the serotonergic system in the brain, to be required for normal aggressive and anxiety-like behavior.[2] Thus, when FEV is genetically deleted from the mouse genome, male mice will instantly attack other males, whereas their wild-type counterparts take significantly longer to initiate violent behaviour. In addition, FEV has been shown to be required for correct maternal behaviour in mice, such that their offspring do not survive unless cross-fostered to other wild-type female mice.[3] A genetic basis for instinctive behavioural traits among non-human species, such as in the above example, is commonly accepted among many biologists; however, attempting to use a genetic basis to explain complex behaviours in human societies has remained extremely controversial.[citation needed]

History According to the OED, E. O. Wilson coined the word "sociobiology" at a 1946 conference on genetics and social behaviour, and it became widely used after it was popularized by Edward O. Wilson in his 1975 book, Sociobiology: The New Synthesis. However, the influence of evolution on behavior has been of interest to biologists and philosophers since soon after the discovery of evolution itself. Peter Kropotkin's Mutual Aid: A Factor of Evolution, written in the early 1890s, is a popular example. Antecedents of modern sociobiological thinking can be traced to the 1960s and the work of such biologists as Richard D. Alexander, Robert Trivers and William D. Hamilton. The idea of the inheritance of

E. O. Wilson, a central figure in the history of sociobiology.

Sociobiology

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behaviour arose from J B S Haldane's idea about how "altruistic behaviour" (see Altruism) could be passed from generation to generation Nonetheless, it was Wilson's book that pioneered and popularized the attempt to explain the evolutionary mechanics behind social behaviors such as altruism, aggression, and nurturence, primarily in ants (Wilson's own research specialty) but also in other animals (bees, wasps and termites). The final chapter of the book is devoted to sociobiological explanations of human behavior, and Wilson later wrote a Pulitzer Prize winning book, On Human Nature, that addressed human behavior specifically. Edward H. Hagen writes in The Handbook of Evolutionary Psychology that sociobiology is, despite the public controversy regarding the applications to humans, "one of the scientific triumphs of the twentieth century." "Sociobiology is now part of the core research and curriculum of virtually all biology departments, and it is a foundation of the work of almost all field biologists" Sociobiological research on nonhuman organisms has increased dramatically and appears continuously in the world's top scientific journals such as Nature and Science.The more general term behavioral ecology is commonly used as substitute for the term sociobiology in order to avoid the public controversy.[4]

Sociobiological theory

Sociology

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Theory History

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Positivism

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Antipositivism

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Functionalism

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Conflict theories

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Middle-range

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Mathematical

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Critical theory

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Socialization

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Structure and agency Research methods

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Quantitative

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Qualitative

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Historical

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Computational

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Ethnographic

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Network-analytic

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Topics Subfields

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Change

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Cities

Sociobiology

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Class

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Crime

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Culture

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Development

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Deviance

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Demography

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Education

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Economy

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Environment

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Family

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Gender

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Health

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Industry

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Internet

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Knowledge

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Law

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Literature

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Medicine

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Mobility

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Movements

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Networks

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Organizations

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Politics

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Race & ethnicity

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Religion

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Rural

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Science

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Soc. psychology

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Stratification

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Technology Browse

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Portal Sociologists Journals Lists

Part of a series on

Ethology

Sociobiology

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Animals portal Biology portal Category

Sociobiologists believe that human behavior, as well as nonhuman animal behavior, can be partly explained as the outcome of natural selection. They contend that in order to fully understand behavior, it must be analyzed in terms of evolutionary considerations. Natural selection is fundamental to evolutionary theory. Variants of hereditary traits which increase an organism's ability to survive and reproduce will be more greatly represented in subsequent generations, i.e., they will be "selected for". Thus, inherited behavioral mechanisms that allowed an organism a greater chance of surviving and/or reproducing in the past are more likely to survive in present organisms. That inherited adaptive behaviors are present in nonhuman animal species has been multiply demonstrated by biologists, and it has become a foundation of evolutionary biology. However, there is continued resistance by some researchers over the application of evolutionary models to humans, particularly from within the social sciences, where culture has long been assumed to be the predominant driver of behavior. Sociobiology is based upon two fundamental premises: • Certain behavioral traits are inherited, • Inherited behavioral traits have been honed by natural selection. Therefore, these traits were probably "adaptive" in the species` evolutionarily evolved environment. Sociobiology uses Nikolaas Tinbergen's four categories of questions and explanations of animal behavior. Two categories are at the species level; two, at the individual level. The species-level categories (often called “ultimate explanations”) are • the function (i.e., adaptation) that a behavior serves and • the evolutionary process (i.e., phylogeny) that resulted in this functionality. The individual-level categories (often called “proximate explanations”) are • the development of the individual (i.e., ontogeny) and • the proximate mechanism (e.g., brain anatomy and hormones). Sociobiologists are interested in how behavior can be explained logically as a result of selective pressures in the history of a species. Thus, they are often interested in instinctive, or intuitive behavior, and in explaining the similarities, rather than the differences, between cultures. For example, mothers within many species of mammals – including humans – are very protective of their offspring. Sociobiologists reason that this protective behavior likely

Sociobiology evolved over time because it helped those individuals which had the characteristic to survive and reproduce. Over time, individuals who exhibited such protective behaviours would have had more surviving offspring than those who did not display such behaviours, such that this parental protection would increase in frequency in the population. In this way, the social behavior is believed to have evolved in a fashion similar to other types of nonbehavioral adaptations, such as (for example) fur or the sense of smell. Individual genetic advantage often fails to explain certain social behaviors as a result of gene-centred selection, and evolution may also act upon groups.[citation needed] The mechanisms responsible for group selection employ paradigms and population statistics borrowed from evolutionary game theory. E.O. Wilson argued that altruistic individuals must reproduce their own altruistic genetic traits for altruism to survive. When altruists lavish their resources on non-altruists at the expense of their own kind, the altruists tend to die out and the others tend to grow. In other words, altruism is more likely to survive if altruists practice the ethic that "charity begins at home". Altruism is defined as "a concern for the welfare of others". An extreme example of altruism involves a soldier risking his life to help a fellow soldier. This example raises questions about how altruistic genes can be passed on if this soldier dies without having any children to exhibit the same altruistic traits. Within sociobiology, a social behavior is first explained as a sociobiological hypothesis by finding an evolutionarily stable strategy that matches the observed behavior. Stability of a strategy can be difficult to prove, but usually, a well-formed strategy will predict gene frequencies. The hypothesis can be supported by establishing a correlation between the gene frequencies predicted by the strategy, and those expressed in a population. Altruism between social insects and littermates has been explained in such a way. Altruistic behavior, behavior that increases the reproductive fitness of others at the apparent expense of the altruist, in some animals has been correlated to the degree of genome shared between altruistic individuals. A quantitative description of infanticide by male harem-mating animals when the alpha male is displaced as well as rodent female infanticide and fetal resorption are active areas of study. In general, females with more bearing opportunities may value offspring less, and may also arrange bearing opportunities to maximize the food and protection from mates. An important concept in sociobiology is that temperamental traits within a gene pool and between gene pools exist in an ecological balance. Just as an expansion of a sheep population might encourage the expansion of a wolf population, an expansion of altruistic traits within a gene pool may also encourage the expansion of individuals with dependent traits. Sociobiology is sometimes associated with arguments over the "genetic" basis of intelligence. While sociobiology is predicated on the observation that genes do affect behavior, it is perfectly consistent to be a sociobiologist while arguing that measured IQ variations between individuals reflect mainly cultural or economic rather than genetic factors. However, many critics point out that the usefulness of sociobiology as an explanatory tool breaks down once a trait is so variable as to no longer be exposed to selective pressures. In order to explain aspects of human intelligence as the outcome of selective pressures, it must be demonstrated that those aspects are inherited, or genetic, but this does not necessarily imply differences among individuals: a common genetic inheritance could be shared by all humans, just as the genes responsible for number of limbs are shared by all individuals. Studies of human behavior genetics have generally found behavioral traits such as creativity, extroversion, aggressiveness, and IQ have high heritability. The researchers who carry out those studies are careful to point out that heritability does not constrain the influence that environmental or cultural factors may have on those traits. Criminality is actively under study, but extremely controversial. There are arguments that in some environments criminal behavior might be adaptive.[5] The novelist Elias Canetti also has noted applications of sociobiological theory to cultural practices such as slavery and autocracy. [6]

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Sociobiology

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Differences from evolutionary psychology Sociobiology differs in important ways from evolutionary psychology. Evolutionary psychology studies the animal nervous system from an evolutionary perspective, including aspects such as vision and navigation that are not necessarily related to social behavior. Sociobiology is restricted to the biology of social behavior but also studies organisms like plants. Evolutionary psychologists focus on the neural mechanisms that cause behavior whereas sociobiologists usually study only behavior. Evolutionary psychology emphasizes that, for humans, neural mechanisms evolved in an ancestral environment that differed from the current environment whereas animal sociobiologists look at animal adaptions to the current environment.

A true friend

Criticism Many critics draw an intellectual link between sociobiology and biological determinism, the belief that most human differences can be traced to specific genes rather than differences in culture or social environments. Critics also see parallels between sociobiology and biological determinism as a philosophy underlying the social Darwinian and eugenics movements of the early 20th century, and controversies in the history of intelligence testing. Steven Pinker argues that critics have been overly swayed by politics and a fear of biological determinism.[7] However, all these critics have claimed that sociobiology fails on In the decades after World War II, eugenics became increasingly unpopular within scientific grounds, independent of their academic science. Many organizations and journals that had their origins in the political critiques. In particular, Lewontin, eugenics movement began to distance themselves from the philosophy, as when Rose & Kamin drew a detailed distinction Eugenics Quarterly became Social Biology in 1969. between the politics and history of an idea and its scientific validity, as has Stephen Jay Gould.[8] Wilson and his supporters counter the intellectual link by denying that Wilson had a political agenda, still less a right-wing one. They pointed out that Wilson had personally adopted a number of liberal political stances and had attracted progressive sympathy for his outspoken environmentalism. They argued that as scientists they had a duty to uncover the truth whether that was politically correct or not. They argued that sociobiology does not necessarily lead to any particular political ideology, as many critics implied. Many subsequent sociobiologists, including Robert Wright, Anne Campbell, Frans de Waal and Sarah Blaffer Hrdy, have used sociobiology to argue quite separate points. It is often and incorrectly argued that Noam Chomsky is a critic of sociobiology. During a 1976 meeting of the Sociobiology Study Group, as reported by Ullica Segerstrale, Chomsky argued for the importance of a sociobiologically informed notion of human nature: "Chomsky even stated that he thought it was important for political radicals to postulate a relatively fixed human nature in order to be able to struggle for a better society. We

Sociobiology need a clear view of human needs in order to know the kind of society we want, Chomsky proclaimed. Not surprisingly ... no Chomsky critique of sociobiology emerged."[9] The argument that human beings are biological organisms and ought to be studied as such is a deeply entrenched theme in Chomsky's work and he has been the foremost critic of the doctrine of the "blank slate" in the social sciences (which would inspire a great deal of Steven Pinker's and others' work in evolutionary psychology), sentiments that are well articulated in the following passage: The development of personality, behavior patterns, and cognitive structures in higher organisms has often been approached in a very different way. It is generally assumed that in these domains, social environment is the dominant factor. The structures of mind that develop over time are taken to be arbitrary and accidental; there is no “human nature” apart from what develops as a specific historical product. According to this view, typical of empiricist speculation, certain general principles of learning that are common in their essentials to all (or some large class of) organisms suffice to account for the cognitive structures attained by humans, structures which incorporate the principles by which human behavior is planned, organized, and controlled. But human cognitive systems, when seriously investigated, prove to be no less marvelous and intricate than the physical structures that develop in the life of the organism. Why, then, should we not study the acquisition of a cognitive structure such as language more or less as we study some complex bodily organ?"[10] Chomsky has also hinted at the possible reconciliation of his anarchist political views and sociobiology in a discussion of Peter Kropotkin's Mutual Aid: A Factor of Evolution, which focused more on altruism than aggression, suggesting that anarchist societies were feasible because of an innate human tendency to cooperate.[11] Wilson's claims that he had never meant to imply what ought to be, only what is the case are supported by his writings, which are descriptive, not prescriptive. However, some critics have argued that the language of sociobiology sometimes slips from "is" to "ought", leading sociobiologists to make arguments against social reform on the basis that socially progressive societies are at odds with our innermost nature.[citation needed] Views such as this, however, are often criticized as examples of the naturalistic fallacy, when reasoning jumps from descriptions about what is to prescriptions about what ought to be. (A common example is the justification of militarism if scientific evidence showed warfare was part of human nature.) It has also been argued that opposition to stances considered anti-social, such as ethnic nepotism, are based on moral assumptions, not bioscientific assumptions, meaning that it is not vulnerable to being disproved by bioscientific advances.:145 The history of this debate, and others related to it, are covered in detail by Cronin (1992), Segerstråle (2000), and Alcock (2001).

References Notes [1] Wilson, E.O. (1978) On Human Nature Page x, Cambridge, Ma: Harvard [2] Hendricks TJ, Fyodorov DV, Wegman LJ, Lelutiu NB, Pehek EA, Yamamoto B, Silver J, Weeber EJ, Sweatt JD, Deneris ES. Pet-1 ETS gene plays a critical role in 5-HT neuron development and is required for normal anxiety-like and aggressive behaviour. Neuron. 2003 Jan 23;37(2):233-47 [3] Lerch-Haner JK, Frierson D, Crawford LK, Beck SG, Deneris ES. Serotonergic transcriptional programming determines maternal behavior and offspring survival. Nat Neurosci. 2008 Sep;11(9):1001-3. [4] The Handbook of Evolutionary Psychology, edited by David M. Buss, John Wiley & Sons, Inc., 2005. Chapter 5 by Edward H. Hagen . [5] The Sociobiology Of Sociopathy: An Integrated (http:/ / www. bbsonline. org/ Preprints/ OldArchive/ bbs. mealey. html) [6] Elias Canetti, Crowds and Power. Harmondsworth: Penguin, 1981, p. 444-445. Marx noted that in Roman times, the slave was described as the "speaking implement", the working animal as the "semi-mute implement" and the plough as the "mute implement" – Capital, Volume I, Penguin, p. 303 note. "The term nigger... implies part-animal status." – Kenneth Neill Cameron, Marxism, the science of society: an introduction. South Hadley, Mass: Bergin & Garvey Publishers, 1985. p. 141. Paul Laurence Dunbar (1872–1906), one of the first black poets to gain national recognition, famously wrote the poem "We wear the mask". (http:/ / www. cummingsstudyguides. net/ Guides4/ Dunbar. html)

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Sociobiology [7] Pinker, Steven (2002). The Blank Slate: The Modern Denial of Human Nature. New York: Viking. [8] Gould, S.J. (1996) "The Mismeasure of Man", Introduction to the Revised Edition [9] Segerstrale 2001:205 [10] Chomsky 1975:10 [11] Chomsky, Noam (1995). "Rollback, Part II." (http:/ / www. chomsky. info/ articles/ 199505--. htm#TXT2. 23) Z Magazine 8 (Feb.): 20-31.

Bibliography • Alcock, John (2001). The Triumph of Sociobiology. Oxford: Oxford University Press. Directly rebuts several of the above criticisms and misconceptions listed above. • Barkow, Jerome (Ed.). (2006) Missing the Revolution: Darwinism for Social Scientists. Oxford: Oxford University Press. • Cronin, H. (1992). The Ant and the Peacock: Altruism and Sexual Selection from Darwin to Today. Cambridge: Cambridge University Press. • Nancy Etcoff (1999). Survival of the Prettiest: The Science of Beauty. Anchor Books. ISBN 0-385-47942-5. • Haugan, Gørill (2006) Nursing home patients’ spirituality. Interaction of the spiritual, physical, emotional and social dimensions (Faculty of Nursing, Sør-Trøndelag University College Norwegian University of Science and Technology) • Richard M. Lerner (1992). Final Solutions: Biology, Prejudice, and Genocide. Pennsylvania State University Press. ISBN 0-271-00793-1. • Richards, Janet Radcliffe (2000). Human Nature After Darwin: A Philosophical Introduction. London: Routledge. • Segerstråle, Ullica (2000). Defenders of the Truth: The Battle for Science in the Sociobiology Debate and Beyond. Oxford: Oxford University Press. • Gisela Kaplan, Lesley J Rogers (2003). Gene Worship: Moving Beyond the Nature/Nurture Debate over Genes, Brain, and Gender. Other Press. ISBN 1-59051-034-8. • F. H. Schmidt: Verhaltensforschung und Recht, Berlin, Duncker & Humblot, 1982, ISBN 3 428 05099 1

External links • Sociobiology (http://plato.stanford.edu/entries/sociobiology/) (Stanford Encyclopedia of Philosophy) Harmon Holcomb (http://www.uky.edu/AS/Philosophy/HarmonHolcomb.htm) & Jason Byron (http://www. pitt.edu/~jmb165) • The Sociobiology of Sociopathy, Mealey, 1995 (http://www.bbsonline.org/Preprints/OldArchive/bbs.mealey. html) • Speak, Darwinists! (http://www.froes.dds.nl) Interviews with leading sociobiologists. • Race and Creation (http://www.prospect-magazine.co.uk/article_details.php?id=6467) - Richard Dawkins • Genetic Similarity and Ethnic Nationalism (http://www.psychology.uwo.ca/faculty/rushtonpdfs/N&N 2005-1.pdf) - An Attempted Sociobiological Explanation of the scientific basis for Political Group Formation. • A brief history on sociobiology (http://www.nytimes.com/2008/07/15/science/15wils.html?pagewanted=1)

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Biocultural anthropology

Biocultural anthropology Biocultural anthropology is the scientific exploration of the relationships between human biology and culture. Physical anthropologists throughout the first half of the 20th century viewed this relationship from a racial perspective; that is, from the assumption that typological human biological differences lead to cultural differences. After World War II the emphasis began to shift toward an effort to explore the role culture plays in shaping human biology. Contemporary biocultural anthropologists view culture as having several key roles in human biological variation: • Culture is a major human adaptation, permitting individuals and populations to adapt to widely varying local ecologies. • Characteristic human biological or biobehavioral features, such as a large frontal cortex and intensive parenting compared to other primates, are viewed in part as an adaption to the complex social relations created by culture. • Culture shapes the political economy, thereby influencing what resources are available to individuals to feed and shelter themselves, protect themselves from disease, and otherwise maintain their health. • Culture shapes the way people think about the world, altering their biology by influencing their behavior (e.g., food choice) or more directly through psychosomatic effects (e.g., the biological effects of psychological stress). While biocultural anthropologists are found in many academic anthropology departments, usually as a minority of the faculty, certain departments have placed considerable emphasis on the "biocultural synthesis." Historically, this has included Emory University, the University of Alabama, and the University of Washington [1], each of which built Ph.D. programs around biocultural anthropology; Binghamton University, which has a M.S. program in biomedical anthropology; UMass Amherst, University of Kentucky and others. Paul Baker, an anthropologist at Penn State whose work focused upon human adaptation to environmental variations, is credited with having popularized the concept of "biocultural" anthropology as a distinct subcategory of anthropology in general. Many anthropologists consider biocultural anthropology as the future of anthropology because it serves as a guiding force towards greater integration of the subdisciplines.

Controversy Other anthropologists, both biological and cultural, have criticized the biocultural synthesis, generally as part of a broader critique of "four-field holism" in U.S. anthropology (see anthropology main article). Typically such criticisms rest on the belief that biocultural anthropology imposes holism upon the biological and cultural subfields without adding value, or even destructively. For instance, contributors in the edited volume Unwrapping the Sacred Bundle: Reflections on the Disciplining of Anthropology[2] argued that the biocultural synthesis, and anthropological holism more generally, are artifacts from 19th century social evolutionary thought that inappropriately impose scientific positivism upon cultural anthropology. Some departments of anthropology have fully split, usually dividing scientific from humanistic anthropologists, such as Stanford's highly publicized 1998 division into departments of "Cultural and Social Anthropology" and "Anthropological Sciences." Underscoring the continuing controversy, this split is now being reversed over the objections of some faculty.[3] Other departments, such as at Harvard, have distinct biological and sociocultural anthropology "wings" not designed to foster cross subdisciplinary interchange.

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Biocultural anthropology

References [1] http:/ / depts. washington. edu/ anthweb/ programs/ biocultural. php [2] introduction: (http:/ / pzacad. pitzer. edu/ ~dsegal/ theory/ yanasegal. pdf) reviews: (http:/ / www. anthrosource. net/ doi/ abs/ 10. 1525/ jlat. 2006. 11. 1. 235) (http:/ / www. anthrosource. net/ doi/ abs/ 10. 1525/ an. 2006. 47. 1. 8. 2?journalCode=an) (http:/ / www. blackwell-synergy. com/ doi/ abs/ 10. 1111/ j. 1467-9655. 2006. 00372_39. x) (http:/ / muse. jhu. edu/ login?uri=/ journals/ anthropological_quarterly/ v078/ 78. 4song. pdf) [3] Anthropology departments instructed to form combined unit (http:/ / news-service. stanford. edu/ news/ 2007/ february14/ anthsci-021407. html)

External links • Essays (http://spot.colorado.edu/~kelso/) (http://bioculturalanthropology.blogspot.com/) by Prof. Jack Kelso

Biosocial theory Biosocial Theory is a theory in behavioral and social science that reduces personality disorders and mental illnesses and disabilities to biologically-determined personality traits reacting to environmental stimuli.

Biosocial theory in DBT It is common for therapists using a Dialectical Behavioral Therapy (DBT) model in the treatment of Borderline personality disorder to stress to clients that causes for their condition come both from a biological propensity to their emotional state, and an invalidating environment, that, by its negative reactions, reinforces their dysfunctional behavior. A traumatic event can start the emotional or interpersonal disregulation that spawns a vicious cycle of increased negative behavior as the person continues to be false information that people are receiving react to the environment's invalidation and the environment increasingly devalues them. "DBT is based on a biosocial theory of personality functioning in which BPD is seen as a biological disorder of emotional regulation. The disorder is characterized by heightened sensitivity to emotion, increased emotional in-tensity and a slow return to emotional baseline. Characteristic behaviors and emotional experiences associated with BPD theoretically result from the expression of this biological dysfunction in a social environment experienced as invalidating by the borderline patient."[1] The importance of stressing the biosocial theory to the client in therapy is that the information becomes a tool of validation in itself, offering the client the option of seeing their problems as no fault of their own while also offering them the possibility if taking responsibility for future change. "The biosocial theory suggests that BPD is a disorder of self-regulation, and particularly of emotional regulation, which results from biological irregularities combined with certain dysfunctional environments, as well as from their interaction and transaction over time"[2]

Biosocial theory of creativity This theory suggests that creativity is genetic, and thus, geniuses are indeed born, not made. The first part of the Biosocial Theory of Creativity is that there is a definitive link between madness, also known as irrationality, and creativity. Many of the greatest creative thinkers, Van Gogh as an example, went mad later in their lifetime. Also, mad people who are held in psychotic wards have been found to create masterpieces of art, especially in the avant-garde style. It has been proposed that the suffering of mentally ill people is compensated when they perform great works of art, as an opposite end of their mental spectrum.

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Biosocial theory The second part is that creativity is just an outlet to deal with the madness within the patients. There is also the opposite of this being put forth, in that, madness is just a form of creativity that is misunderstood by the general populace. Both of these forms are extremely controversial and are being debated. A conclusion for this may not be available for many years to come. The third part is that madness is ultimately just a result of some imbalance or defect within the brain. Some examples of these defects are brains that have unusual EEG (Electroencephalography) readings, an unbalanced neurochemistry, abnormal brain structures, or unusual hemisphere lateralization. The fourth and last part is that the creativity associated with madness is inherited, but where the inheritance comes from is not so clear. Genes and DNA is a possibility, but parental trauma that caused a defect within the womb is also possible. Then, there are environmental factors that could create such a madness. Drugs could affect the brain, which then would become an inherited defect. All of these things are possible within the bounds of modern science.

References [1] Murphy, Elizabeth T., PhD, and Gunderson, John, MD. A Promising Treatment (http:/ / web. archive. org/ web/ 19991014032825/ http:/ / www. mcleanhospital. org/ psychupdate/ psyupI-3. htm) for Borderline Personality Disorder, McLean Hospital Psychiatic Update, January 1999. [2] Linehan, M. M. (1993a) Cognitive–Behavioral Treatment of Borderline Personality Disorder. New York: Guilford Press.

External links • An exposition on the drawbacks of viewing human creativity through the lens of the Biosocial Theory (http:// creativestuff.net/Creativity_Theories/Creativity_Theories/Biosocial_Theory_of_Creativity_2006112386.html) by Steven Mizrachs. • Classical Conditioning, Arousal, and Crime: a Biosocial Perspective (http://www-rcf.usc.edu/~raine/Classical. Conditioning.Arousal.Crime.pdf) (1997) by A. Raine.

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Cultural selection theory

Cultural selection theory Cultural selection theory is a scientific discipline that explores sociological and cultural evolution the same way that Darwinian selection theory is used to explain biological evolution. There are three obvious concepts to Cultural Selection. The three concepts are social contagion theory, evolutionary epistemology, and memetics.[1] This theory is an extension of memetics. In memetics, memes, much like biology's genes, are informational units passed through generations of culture. However, unlike memetics, cultural selection theory moves past these isolated "memes" to encompass selection processes, including continuous and quantitative parameters. Two other approaches to cultural selection theory are social contagion and evolutionary epistemology.[2][3] A good example of this theory is found by looking to the reason large businesses tend to grow larger. The answer includes the benefits of mass production and distribution, international advertising, and more funds for product development. These self-amplifying effects, known as the economies of scale, give rise to selection effects which have a quantitative nature, unlike the qualitative effects described by the theory of memetics. On the whole, cultural selection theory embraces the inherent complexity of cultural change and vouches for a systemic, rather than deconstructionist, approach to analyzing the way a society's norms and values change. The Cultural Selection theory faces many objections due to the lack of evidence to support the adaption of natural selection in the structural mechanisms of cultural systems. Major objections against the Cultural Selection Theory stem from Lamarckianism, Genotype-phenotype distinction, Common Hereditary Architecture, Biological Analogue for Cultural Units, and Environmental Interactions.[4] The Biological Analogue for Cultural Units breaks down into 3 subunits. The first is regarding strict analogues. This means that a biological unit (traits etc.) should be related to a cultural unit. This is a way for the old biological model and the modern cultural model to correlate and solidify the point. The second is regarding trait analogues. This means that some analogues are viewed the wrong way. Sometimes, one analogue is mistaken for another and often, the line between the two analogues is unclear and the distinction isn't as evident. The third is regarding virus analogue. This clarifies the point that the ability of the virus is different from the organism and the ability of both the virus and organisms should be looked at independently.[5] Some have argued that in order for the Cultural Selection Theory to stand strong against objections, conclusive and explicit case studies are required. There needs to be empirical support to clarify the interaction between cultural systems and their environments. Crozier conducted a study on the acoustic adaptation of bird songs. This research study provided empirical evidence to support and strengthen the Cultural Selection Theory.[6] Like Darwin's natural selection theory, cultural selection theory has three phases too; variation, reproduction and selection. Variation gives rise to a subject, reproduction is responsible for the spread and selection is based on the factors that control the spread.

References [7]

• Fog, Agner (1999). Cultural selection [8]. Boston: Kluwer Academic Publishers. ISBN 0-7923-5579-2. OCLC 40595346 [9].Wikipedia:Citing sources • Crozier, G. K. D. (2008). "Reconsidering Cultural Selection Theory" [10]. British Journal for the Philosophy of Science 59 (2008): 455–479. doi:10.1093/bjps/axn018 [11]. [1] Reconsidering Cultural Selection TheoryCrozier, G. K. DTHE BRITISH JOURNAL FOR THE PHILOSOPHY OF SCIENCE, V. 59 (3), 09/2008, p. 455-479 [2] (Crozier, 2008) [3] Social contagion theory’s epidemiological approach construes social entities as analogous to parasites that are transmitted virally through a population of biological organisms. Evolutionary epistemology's focus lies in causally connecting evolutionary biology and rationality by generating explanations for why traits for rational behaviour or thought patterns would have been selected for in a species’ evolutionary history. Memetics models cultural change after population genetics, taking cultural units to be analogous to genes.http:/ / bf4dv7zn3u. search.

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Cultural selection theory serialssolutions. com. myaccess. library. utoronto. ca/ ?ctx_ver=Z39. 88-2004& ctx_enc=info%3Aofi%2Fenc%3AUTF-8& rfr_id=info:sid/ summon. serialssolutions. com& rft_val_fmt=info:ofi/ fmt:kev:mtx:journal& rft. genre=article& rft. atitle=Reconsidering+ Cultural+ Selection+ Theory& rft. jtitle=The+ British+ Journal+ for+ the+ Philosophy+ of+ Science& rft. au=Crozier%2C+ G. + K. + D& rft. date=2008-09-01& rft. pub=OXFORD+ UNIV+ PRESS& rft. issn=0007-0882& rft. volume=59& rft. issue=3& rft. spage=455& rft. epage=479& rft_id=info:doi/ 10. 1093%2Fbjps%2Faxn018& rft. externalDBID=PBJP& rft. externalDocID=000258864600011 [4] Crozier GKD (2008) Reconsidering Cultural Selection Theory. Br J Philos Sci 59(3):455–479 Retrieved from: http:/ / simplelink. library. utoronto. ca/ url. cfm/ 323901 [5] Crozier, "Reconsidering Cultural Selection Theory" (http:/ / journals1. scholarsportal. info. myaccess. library. utoronto. ca/ tmp/ 5325867759933723437. pdf), The British Journal for the Philosophy of Science, September 2008 [6] Crozier, G.K.D., 2010. A formal investigation of cultural selection theory: acoustic adaptation in bird song. Biology and Philosophy 25, 781–801. Retrieved from: http:/ / journals1. scholarsportal. info. myaccess. library. utoronto. ca/ tmp/ 129825896213834992. pdf [7] my.access — University of Toronto Libraries Portal (http:/ / bf4dv7zn3u. search. serialssolutions. com. myaccess. library. utoronto. ca/ ?ctx_ver=Z39. 88-2004& ctx_enc=info:ofi/ enc:UTF-8& rfr_id=info:sid/ summon. serialssolutions. com& rft_val_fmt=info:ofi/ fmt:kev:mtx:journal& rft. genre=article& rft. atitle=Reconsidering+ Cultural+ Selection+ Theory& rft. jtitle=The+ British+ Journal+ for+ the+ Philosophy+ of+ Science& rft. au=Crozier,+ G. + K. + D& rft. date=2008-09-01& rft. pub=OXFORD+ UNIV+ PRESS& rft. issn=0007-0882& rft. volume=59& rft. issue=3& rft. spage=455& rft. epage=479& rft_id=info:doi/ 10. 1093/ bjps/ axn018& rft. externalDBID=PBJP& rft. externalDocID=000258864600011) [8] http:/ / www. agner. org/ cultsel/ toc. php [9] http:/ / www. worldcat. org/ oclc/ 40595346 [10] http:/ / bjps. oxfordjournals. org/ content/ 59/ 3/ 455. short [11] http:/ / dx. doi. org/ 10. 1093%2Fbjps%2Faxn018

Darwinian anthropology Darwinian anthropology describes an approach to anthropological analysis which employs various theories from darwinian evolutionary biology. Whilst there are a number of areas of research that can come under this broad description (Marks, 2004) some specific research projects have been closely associated with the label. A prominent example is the project that developed in the mid 1970s with the goal of applying sociobiological perspectives to explain patterns of human social relationships, particularly kinship patterns across human cultures. This kinship-focused Darwinian anthropology was a significant intellectual forebear of evolutionary psychology, and both draw on biological theories of the evolution of social behavior (in particular inclusive fitness theory) upon which the field of sociobiology was founded.

Overview In 1974 the biologist Richard D. Alexander published an article The Evolution of Social Behavior which drew upon W.D.Hamilton's work on inclusive fitness and kin selection and noted that: Although ten years have passed since Hamilton's landmark papers, apparently only a single social scientist (Campbell, 31) has made a distinct effort to incorporate kin selection into theories of human altruism.[...] But so have the biologists, for one reason or another, failed to consider the enormous literature on topics like kinship systems and reciprocity in human behavior.(Alexander 1974, 326) Amongst other suggestions, Alexander suggested that certain patterns of social cooperation documented by ethnographers, in particular the avunculate ('mother's brother') relationship, could be explained in reference to individuals pursuing a strategy of individual inclusive fitness maximization under conditions of low certainty-of-paternity. This hypothesis was subsequently taken up and elaborated in a series of studies by other darwinian anthropologists: The hypothesis follows that matrilineal inheritance is a cultural trait that evolved in response to low probability of paternity. [...] The paternity hypothesis was rescued and made explicit in the context of modern evolutionary theory by Alexander in 1974. Over the next 10 years this provided the impetus leading to numerous theoretical refinements and scholarly and empirical investigations (Flinn 1981;

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Darwinian anthropology Gaulin & Schlegel 1980; Greene 1978; Hartung 1981b; Kurland 1979).(Hartung 1985,661-663) Ultimately these analyses were considered unsuccessful, and were specifically criticized by other sociobiologists on a number of grounds. One problem was said to be that interpreting inclusive fitness theory to imply that individuals have evolved the characteristic of pursuing strategies to maximize their own 'inclusive fitness' is erroneous; the theory should instead be interpreted to describe selection pressures on genes: Alexander’s argument... erred through looking at things from the point of view of an individual... I believe this kind of error is all too easy to make when we use the technical term ‘fitness’ [of individuals]. This is why I have avoided using the term in this book. There is really only one entity whose point of view matters in evolution, and that entity is the selfish gene.(Dawkins 1989 (1976), 137 emphasis in original) A related problem was that, in assuming that individuals simply operate as inclusive fitness maximizing agents, any investigation of the proximate psychological mechanisms that mediate social behaviors was ignored. Symons made this observation in his 1989 Critique of Darwinian Anthropology: DA’s central hypothesis is that “evolved behavioral tendencies” cause human “behavior to assume the form that maximizes inclusive fitness”(Irons 1979b, 33). Turke and Betzig (1985) state this hypothesis as a formal prediction: “Modern Darwinian theory predicts that human behavior will be adaptive, that is, designed to promote maximum reproductive success through available descendant and nondescendant relatives.”(p 79)… [T]he key terms in [this] quotation are used in DA to bypass the question of phenotypic design in characterizations of adaptation.(Symons 1989, 131-132) Symons, along with Tooby and Cosmides, were amongst the founders of the emerging school of evolutionary psychology, whose aim has been to focus more on these hypothesized proximate psychological mechanisms.

Theoretical background Darwinian anthropology was critiqued by Symonds for its agnosticism as to the psychological mechanisms governing how social behavior is actually expressed in the human species, and its reliance on interpreting inclusive fitness theory to simply imply that humans have evolved to be inclusive fitness maximizers. This section will review some of the relevant background discussion in inclusive fitness theory to clarify why this position was considered untenable.

Evolutionary versus proximate explanations Inclusive Fitness theory has often been interpreted to mean that social behavior per se is a goal of evolution, and also that genes (or individual organisms) are selected to find ways of actively distinguishing the identity of close genetic relatives ‘in order to’ engage in social behaviors with them. [M]any misunderstandings persist. In many cases, they result from conflating “coefficient of relatedness” and “proportion of shared genes,” which is a short step from the intuitively appealing—but incorrect—interpretation that “animals tend to be altruistic toward those with whom they share a lot of genes.” These misunderstandings don’t just crop up occasionally; they are repeated in many writings, including undergraduate psychology textbooks—most of them in the field of social psychology, within sections describing evolutionary approaches to altruism. (Park 2007, p860)[1] The apparent rationale for this common mis-interpretation is that organisms would thereby benefit the “Inclusive Fitness of the individuals (and genes) involved”. This approach overlooks the point that evolution is not a teleological process, but a passive, consequential and undirected biological process, where environmental variations and drift effects are present alongside random gene mutations and natural selection. Inclusive fitness theory takes the form of an ultimate explanation, specifically a criterion (br>c), for the evolution of social behaviors, not a proximate mechanism governing the expression of social behaviors. What forms of social

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Darwinian anthropology behavior might meet this criterion are cannot be a priori specified by the theory, nor can it shed light on whether the life history of a species provides opportunities for social interactions to occur. Thus, strictly speaking, the interpretation that organisms ‘have evolved to’ direct social behavior towards genetic relatives is not implied by the theory (see also inclusive fitness). Investigating how inclusive fitness theory might apply to the potential emergence of social traits in any given species must begin with an analysis of the evolutionarily typical ecological niche, demographics, and patterns of interaction of that species. Where significant interaction between individuals is not present in the life history of a species, the theory is necessarily null regarding social behaviors between individuals. As Silk (2001) put it; The role of kinship in the daily lives of animals depends on the demographic composition of the groups in which they live. Kin selection will only be an important force in the evolution of social behavior if animals find themselves in situations where they have an opportunity to fulfill the predictions of Hamilton’s rule. At a minimum, kin must be available. The number, availability, and degree of relatedness among kin will depend on how groups are constructed in nature.” (Silk 2001, 77) Consideration must thus be given to whether the ecological niche leads to the clustering of individuals in groups or whether individuals are typically solitary. Socioecology research, for example, suggests that fundamental influences on demographic patterns are the distribution/fecundity of primary food sources as well as patterns of predation. When considering social behavior traits of a given species, consideration of these influences is in a sense, logically prior to analyses of inclusive fitness pressures on the species.

Selection pressure on genes or strategy of individuals Darwinian anthropology, following R. D. Alexander, used the notion of the inclusive fitness of individuals rather than the inclusive fitness of genes. Dawkins (above) pointed to this as an error. The source of the confusion can be traced to discussions in Hamilton's early papers on inclusive fitness. In his 1963 paper Hamilton refers, unambiguously, to selection pressures on genes; [T]he ultimate criterion which determines whether G [a gene] will spread is not whether the behaviour is to the benefit of the behaver but whether it is to the benefit of the gene G; and this will be the case if the average net result of the behaviour is to add to the gene pool a handful of genes containing G in higher concentration than does the gene pool itself.” (1996 [1963], 7) However, in his paper published in 1964, actually written before the 1963 paper (Hamilton 1996), Hamilton had included a subsidiary discussion on what the genetic theory might imply for how we look at the fitness of individuals: Actually, in the preceding mathematical account we were not concerned with the inclusive fitness of individuals as described here but rather with certain averages of them which we call the inclusive fitness of types. But the idea of the inclusive fitness of an individual is nevertheless a useful one. Just as in the sense of classical selection we may consider whether a given character expressed in an individual is adaptive in the sense of being in the interest of his personal fitness or not, so in the present sense of selection we may consider whether the character or trait of behaviour is or is not adaptive in the sense if being in the interest of his inclusive fitness.” (Hamilton 1996 [1964], 38) It is clear here that the formal treatment is of the selection pressures on types (genes or traits), whilst the notion of individual inclusive fitness may serve as a guide to the adaptiveness of the trait; just as consideration of effects of a trait on an individual's fitness can be instructive when considering classical selection on traits. At the same time, it is understandable that Alexander took the inclusive fitness of individuals as a heuristic device.

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Darwinian anthropology

Context-based or discrimination-based expression mechanisms In his 1964 paper, Hamilton 'hazards' “the following unrigorous statement of the main principle that has emerged from the model”; The Social behaviour of a species evolves in such a way that in each distinct behaviour-evoking situation the individual will seem to value his neighbours’ fitness against his own according to the coefficients of relationship appropriate to that situation.”(1964 [1996], 49) He uses the terms 'hazards', 'unrigorous', and 'will seem' deliberately, since his formal analysis makes clear that the model specifies the evolutionary selection pressure, rather than specifying what mechanisms govern the proximate expression of social behaviors. He also clearly points to social behaviors being evoked in distinct situations, and that individuals may encounter potential social recipients of different degree of relationship in different situations. If one ignores the cautious qualifying words however, the passage might readily be interpreted to imply that individuals are indeed expected to make an active assessment of the degree of relatedness of others they interact with in different situations. Later in the paper, Hamilton again discusses the issue of whether the performance (or expression) of social behaviors might be conditional on; (a) discriminating factors which correlate with close relationship with the recipient, or (b) actually discriminating which individuals 'really are' in close relationship with the recipient: The selective advantage which makes behaviour conditional in the right sense on the discrimination of factors which correlate with the relationship of the individual concerned is therefore obvious. It may be, for instance, that in respect of a certain social action performed towards neighbours indiscriminately, an individual is only just breaking even in terms of inclusive fitness. If he could learn to recognise those of his neighbours who really were close relatives and could devote his beneficial actions to them alone an advantage to inclusive fitness would at once appear. Thus a mutation causing such discriminatory behaviour itself benefits inclusive fitness and would be selected. In fact, the individual may not need to perform any discrimination so sophisticated as we suggest here; a difference in the generosity of his behaviour according to whether the situations evoking it were encountered near to, or far from, his own home might occasion an advantage of a similar kind.” (1996 [1964], 51) For certain social behaviors, Hamilton suggests there may be selection pressure for more discerning discrimination of genetic relatedness, were the mutation to occur. But 'in fact' the same net result of accurately targeting social behaviors towards genetic relatives could be achieved via a simpler mechanism of being expressed in proximity to the actor's 'home'. Hamilton is thus agnostic as to whether evolved social behaviors might be expressed via straightforward proximate mechanisms such as location-based cues, or whether more specific discriminatory powers might govern their expression. He does suggest that the distinct social contexts within which various social behaviors are expressed are factors to consider. Other theorists have discussed these questions of whether proximity, context or more discriminatory expression may govern behaviors: Animals cannot, of course, be expected to know, in a cognitive sense, who their relatives are, and in practice the behaviour that is favoured by natural selection will be equivalent to a rough rule of thumb such as ‘share food with anything that moves in the nest in which you are sitting.’ If families happen to go around in groups, this fact provides a useful rule of thumb for kin selection: ‘care for any individual you often see’.” (Dawkins 1979, 187) If, for example, animals behave with an equal degree of altruism to all their “neighbours”… and if on average animals are related to their neighbours, then I would regard this as an example of kin selection. It is not a necessary feature of kin selection that an animal should distinguish different degrees of relationship among its neighbours, and behave with greater altruism to the more closely related…”(Maynard Smith 1976, 282 emphasis in original) Dawkins believes social behaviors will in practice be governed by context-based expression. Maynard Smith is, like Hamilton, agnostic, but reiterates the point that context-based cues might well govern their expression and that actively distinguishing relatives is not necessarily expected for the expression of those social traits whose evolution

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Darwinian anthropology is governed by inclusive fitness criteria. In sum, inclusive fitness theory does imply that; the evolutionary emergence of social behavior can occur where there is statistical association of genes between social actors and recipients; but that the expression of such evolved social behaviors is not necessarily governed by actual genetic relatedness between participants. The evolutionary criterion and the proximate mechanism must thus not be confused: the first does require genetic association (of the form br>c), the second does not. Darwinian anthropology's central premise that human behavior has evolved to maximize the inclusive fitness of individuals is thus not a logical derivative of the theory. Also, the notion that humans will discriminate social behaviors towards genetic relatives is again not entailed by the theory.

Reception by anthropologists Before the questions raised within anthropology about the study of ‘kinship’ by Schneider and others from the 1960s onwards, anthropology itself had paid very little attention to the notion that social bonds were anything other than connected to consanguinal (or genetic) relatedness (or its local cultural conceptions). The social bonding associated with provision of and sharing of food was one important exception, particularly in the work of Richards, but this was largely ignored by descriptions of ‘kinship’ till more recently. Although questioning the means by which ‘kinship bonds’ form, few of these early accounts questioned the fundamental role of ‘procreative ties’ in social bonding (Schneider, 1984). From the 1950s onwards, reports on kinship patterns in the New Guinea Highlands added some momentum to what had until then been only occasional fleeting suggestions that living together (co-residence) might underlie social bonding, and eventually contributed to the general shift away from a genealogical approach. For example, on the basis of his observations, Barnes suggested: [C]learly, genealogical connexion of some sort is one criterion for membership of many social groups. But it may not be the only criterion; birth, or residence, or a parent’s former residence, or utilization of garden land, or participation in exchange and feasting activities or in house-building or raiding, may be other relevant criteria for group membership.”(Barnes 1962,6) Similarly, Langness' ethnography of the Bena Bena also emphasized a break with the genealogical perspective: The sheer fact of residence in a Bena Bena group can and does determine kinship. People do not necessarily reside where they do because they are kinsmen: rather they become kinsmen because they reside there.” (Langness 1964, 172 emphasis in original) By 1972, Schneider had raised deep problems with the notion that human social bonds and 'kinship' was a natural category built upon genealogical ties (for more information, see kinship), and especially in the wake of his 1984 critique this has become broadly accepted by most, if not all, anthropologists. The darwinian anthropology (and other sociobiological) perspectives, arising in the early 1970s, had not unreasonably assumed that the genealogical conceptions of human kinship, in place since Morgan's early work in the 1870s, were still valid as a universal feature of humans. But they emerged at precisely the time that anthropology, being particularly sensitive about its own apparent 'ethnocentric' generalizations about kinship (from cultural particulars to human universals) was seeking to distance itself from these conceptions. The vehemence of Sahlins' rebuttal of sociobiology's genetic relatedness perspective in his 1976 The use and abuse of Biology, which underlined the non-genealogical nature of human kinship, can be understood as part of this 'distancing' trend.

Alternative approaches The lack of success of darwinian anthropology created space for alternative approaches to analyzing human social behaviors from a biological perspective. Alexander's initial point (above) that the inclusive fitness framework had been scarcely applied to human kinship and social patterns has remained largely valid. But the move away from genealogical kinship in anthropology has continued to be a major barrier to any potential resolution. This section reviews a range of approaches to synthesizing ideas from evolutionary biology to observations and data about human

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Darwinian anthropology social behaviour across contemporary human populations. Whilst some of these approaches include the inclusive fitness approach, others may seek to demonstrate fit to other theories from evolutionary biology, or to demonstrate that certain proximate mechanisms of social behaviour are both compatible with the inclusive fitness approach, and also with the broad variety of ethnographic data on human kinship patterns.

Evolutionary psychology main article: evolutionary psychology

Human behavioural ecology main article: Human behavioral ecology

Nurture kinship Main article: Nurture kinship

Criticism Theories in evolutionary biology relvant to understanding social behavior may not be limited to frameworks such as inclusive fitness theory. The theory of reciprocal altruism may have equal or greater explanatory power for some forms of human social behavior, and perhaps kinship patterns. Other approaches may maintain that human behavior is less amenable to biological analysis due to the prominent influence of social learning and cultural transmission in the human species, and instead advance ideas based on the role of e.g. culture, historical contingencies or economic/environmental conditions. All or any of these may or may not contribute valuable insights to our understanding of social behavior and social patterns in humans.

References [1] Park, J.H. (2007) Persistent Misunderstandings of Inclusive Fitness and Kin Selection: Their Ubiquitous Appearance in Social Psychology Textbooks. Evolutionary Psychology 5(4): 860-873.

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Dual inheritance theory

Dual inheritance theory Dual inheritance theory (DIT), also known as gene–culture coevolution or biocultural evolution, was developed in the late 1970s and early 1980s to explain how human behavior is a product of two different and interacting evolutionary processes: genetic evolution and cultural evolution. In DIT, culture is defined as information and behavior acquired through social learning. One of the theory's central claims is that culture evolves partly through a Darwinian selection process, which dual inheritance theorists often describe by analogy to genetic evolution. Because genetic evolution is relatively well understood, most of DIT examines cultural evolution and the interactions between cultural evolution and genetic evolution.

Theoretical basis DIT holds that genetic and cultural evolution interacted in the evolution of Homo sapiens. DIT recognizes that the natural selection of genotypes is an important component of the evolution of human behavior and that cultural traits can be constrained by genetic imperatives. However, DIT also recognizes that genetic evolution has endowed the human species with a parallel evolutionary process of cultural evolution. DIT makes three main claims:[1]

Culture capacities are adaptations The human capacity to store and transmit culture arose from genetically evolved psychological mechanisms. This implies that at some point during the evolution of the human species a type of social learning leading to cumulative cultural evolution was evolutionarily advantageous.

Culture evolves Social learning processes give rise to cultural evolution. Cultural traits are transmitted differently than genetic traits and, therefore, result in different population-level effects on behavioral variation.

Genes and culture coevolve Cultural traits alter the social and physical environments under which genetic selection operates. For example, the cultural adoptions of agriculture and dairying have, in humans, caused genetic selection for the traits to digest starch and lactose, respectively.[2][3][4][5][6][7] As another example, it is likely that once culture became adaptive, genetic selection caused a refinement of the cognitive architecture that stores and transmits cultural information. This refinement may have further influenced the way culture is stored and the biases that govern its transmission. DIT also predicts that, under certain situations, cultural evolution may select for traits that are genetically maladaptive. An example of this is the demographic transition, which describes the fall of birth rates within industrialized societies. Dual inheritance theorists hypothesize that the demographic transition may be a result of a prestige bias, where individuals that forgo reproduction to gain more influence in industrial societies are more likely to be chosen as cultural models.[8][9]

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Dual inheritance theory

View of culture People have defined the word "culture" to describe a large set of different phenomena.[10][11] A definition that sums up what is meant by "culture" in DIT is: Culture is information stored in individuals' brains that is capable of affecting behavior and that got there through social learning.[12][13] This view of culture emphasizes population thinking by focusing on the process by which culture is generated and maintained. It also views culture as a dynamic property of individuals, as opposed to a view of culture as a superorganic entity to which individuals must conform.[14] This view's main advantage is that it connects individual-level processes to population-level outcomes.[15]

Genetic influence on cultural evolution Genes have an impact on cultural evolution via psychological predispositions on cultural learning.[16] Genes encode much of the information needed to form the human brain. Genes constrain the brain's structure and, hence, the ability of the brain to acquire and store culture. Genes may also endow individuals with certain types of transmission bias (described below).

Cultural influences on genetic evolution Culture can profoundly influence gene frequencies in a population. One of the best known examples is the prevalence of the genotype for adult lactose absorption in human populations, such as Northern Europeans and some African societies, with a long history of raising cattle for milk. Other societies such as East Asians and Amerindians, retain the typical mammalian genotype in which the body shuts down lactase production shortly after the normal age of weaning. This implies that the cultural practice of raising cattle for milk led to selection for genetic traits for lactose digestion.[17] Recently, analysis of natural selection on the human genome suggests that civilization has accelerated genetic change in humans over the past 10,000 years.[18]

Mechanisms of cultural evolution In DIT, the evolution and maintenance of cultures is described by five major mechanisms: natural selection of cultural variants, random variation, cultural drift, guided variation and transmission bias.

Natural selection Cultural differences among individuals can lead to differential survival of individuals. The patterns of this selective process depend on transmission biases and can result in behavior that is more adaptive to a given environment.

Random variation Random variation arises from errors in the learning, display or recall of cultural information, and is roughly analogous to the process of mutation in genetic evolution.

Cultural drift Cultural drift is a process roughly analogous to genetic drift in evolutionary biology.[19][20][21] In cultural drift, the frequency of cultural traits in a population may be subject to random fluctuations due to chance variations in which traits are observed and transmitted (sometimes called "sampling error").[22] These fluctuations might cause cultural variants to disappear from a population. This effect should be especially strong in small populations.[23] A model by Hahn and Bentley shows that cultural drift gives a reasonably good approximation to changes in the popularity of American baby names. Drift processes have also been suggested to explain changes in archaeological pottery and

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Dual inheritance theory technology patent applications. Changes in the songs of song birds are also thought to arise from drift processes, where distinct dialects in different groups occur due to errors in songbird singing and acquisition by successive generations.[24] Cultural drift is also observed in an early computer model of cultural evolution.[25]

Guided variation Cultural traits may be gained in a population through the process of individual learning. Once an individual learns a novel trait, it can be transmitted to other members of the population. The process of guided variation depends on an adaptive standard that determines what cultural variants are learned.

Biased transmission Understanding the different ways that culture traits can be transmitted between individuals has been an important part of DIT research since the 1970s.[26][27] Transmission biases occur when some cultural variants are favored over others during the process of cultural transmission.[28] Boyd and Richerson (1985) defined and analytically modeled a number of possible transmission biases. The list of biases has been refined over the years, especially by Henrich and McElreath. Content bias Content biases result from situations where some aspect of a cultural variant's content makes them more likely to be adopted.[29] Content biases can result from genetic preferences, preferences determined by existing cultural traits, or a combination of the two. For example, food preferences can result from genetic preferences for sugary or fatty foods and socially-learned eating practices and taboos. Content biases are sometimes called "direct biases." Context bias Context biases result from individuals using clues about the social structure of their population to determine what cultural variants to adopt. This determination is made without reference to the content of the variant. There are two major categories of context biases: (1) model-based biases, and (2) frequency-dependent biases. Model-based biases Model-based biases result when an individual is biased to choose a particular "cultural model" to imitate. There are four major categories of model-based biases: (1) prestige bias, (2) skill bias, (3) success bias, (4) similarity bias.[30] A "prestige bias" results when individuals are more likely to imitate cultural models that are seen as having more prestige. A measure of prestige could be the amount of deference shown to a potential cultural model by other individuals. A "skill bias" results when individuals can directly observe different cultural models performing a learned skill and are more likely to imitate cultural models that perform better at the specific skill. A "success bias" results from individuals preferentially imitating cultural models that they determine are most generally successful (as opposed to successful at a specific skill as in the skill bias.) A "similarity bias" results when individuals are more likely to imitate cultural models that are perceived as being similar to the individual based on specific traits. Frequency-dependent biases Frequency-dependent biases result when an individual is biased to choose particular cultural variants based on their perceived frequency in the population. The most explored frequency-dependent bias is the "conformity bias." Conformity biases result when individuals attempt to copy the mean or the mode cultural variant in the population. Another possible frequency dependent bias is the "rarity bias." The rarity bias results when individuals preferentially choose cultural variants that are less common in the population. The rarity bias is also sometimes called a "nonconformist" or "anti-conformist" bias.

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Dual inheritance theory

Social learning and cumulative cultural evolution In DIT, the evolution of culture is dependent on the evolution of social learning. Analytic models show that social learning becomes evolutionarily beneficial when the environment changes with enough frequency that genetic inheritance can not track the changes, but not fast enough that individual learning is more efficient.[31] While other species have social learning, and thus some level of culture, only humans, some birds and chimpanzees are known to have cumulative culture.[32] Boyd and Richerson argue that the evolution of cumulative culture depends on observational learning and is uncommon in other species because it is ineffective when it is rare in a population. They propose that the environmental changes occurring in the Pleistocene may have provided the right environmental conditions.[33] Michael Tomasello argues that cumulative cultural evolution results from a "ratchet effect" that began when humans developed the cognitive architecture to understand others as mental agents.[34] Furthermore Tomasello proposed in the 80s that there are some disparities between the observational learning mechanisms found in humans and great apes - which go some way to explain the observable difference between great ape traditions and human types of culture (see Emulation (observational learning)).

Cultural group selection Although group selection is commonly thought to be nonexistent or unimportant in genetic evolution,[35][36][37] DIT predicts that, due to the nature of cultural inheritance, it may be an important force in cultural evolution. The reason group selection is thought to operate in cultural evolution is because of conformist biases (see above section on transmission biases). Conformist biases make it difficult for novel cultural traits to spread through a population. Conformist bias also helps maintain variation between groups. These two properties, rare in genetic transmission, are necessary for group selection to operate.[38] Based on an earlier model by Cavalli-Sforza and Feldman,[39] Boyd and Richerson show that conformist biases are almost inevitable when traits spread through social learning,[40] implying that group selection is common in cultural evolution. Analysis of small groups in New Guinea imply that cultural group selection might be a good explanation for slowly changing aspects of social structure, but not for rapidly changing fads.[41] The ability of cultural evolution to maintain intergroup diversity is what allows for the study of cultural phylogenetics.[42]

Historical development The idea that human cultures undergo a similar evolutionary process as genetic evolution goes back at least to Darwin[43] In the 1960s, Donald T. Campbell published some of the first theoretical work that adapted principles of evolutionary theory to the evolution of cultures.[44] In 1976, two developments in cultural evolutionary theory set the stage for DIT. In that year Richard Dawkins's The Selfish Gene introduced ideas of cultural evolution to a popular audience. Although one of the best-selling science books of all time, because of its lack of mathematical rigor, it had little impact on the development of DIT. Also in 1976, geneticists Marcus Feldman and Luigi Luca Cavalli-Sforza published the first dynamic models of gene–culture coevolution.[45] These models were to form the basis for subsequent work on DIT, heralded by the publication of three seminal books in the 1980s. The first was Charles Lumsden and E.O. Wilson's Genes, Mind and Culture.[46] This book outlined a series of mathematical models of how genetic evolution might favor the selection of cultural traits and how cultural traits might, in turn, affect the speed of genetic evolution. While it was the first book published describing how genes and culture might coevolve, it had relatively little impact on the further development of DIT.[47] Some critics felt that their models depended too heavily on genetic mechanisms at the expense of cultural mechanisms.[48] Controversy surrounding Wilson's sociobiological theories may also have decreased the lasting impact of this book. The second 1981 book was Cavalli-Sforza and Feldman's Cultural Transmission and Evolution: A Quantitative Approach. Borrowing heavily from population genetics and epidemiology, this book built a mathematical theory concerning the spread of cultural traits. It describes the evolutionary implications of vertical transmission, passing

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Dual inheritance theory cultural traits from parents to offspring; oblique transmission, passing cultural traits from any member of an older generation to a younger generation; and horizontal transmission, passing traits between members of the same population. The next significant DIT publication was Robert Boyd and Peter Richerson's 1985 Culture and the Evolutionary Process. This book presents the now-standard mathematical models of the evolution of social learning under different environmental conditions, the population effects of social learning, various forces of selection on cultural learning rules, different forms of biased transmission and their population-level effects, and conflicts between cultural and genetic evolution. The book's conclusion also outlined areas for future research that are still relevant today.

Current and future research In their 1985 book, Boyd and Richerson outlined an agenda for future DIT research. This agenda, outlined below, called for the development of both theoretical models and empirical research. DIT has since built a rich tradition of theoretical models over the past two decades.[49] However, there has not been a comparable level of empirical work. In a 2006 interview Harvard biologist E. O. Wilson expressed disappointment at the little attention afforded to DIT: "...for some reason I haven't fully fathomed, this most promising frontier of scientific research has attracted very few people and very little effort." [50] Kevin Laland and Gillian Brown attribute this lack of attention to DIT's heavy reliance on formal modeling. "In many ways the most complex and potentially rewarding of all approaches, [DIT], with its multiple processes and cerebral onslaught of sigmas and deltas, may appear too abstract to all but the most enthusiastic reader. Until such a time as the theoretical hieroglyphics can be translated into a respectable empirical science most observers will remain immune to its message."[51] Economist Herbert Gintis disagrees with this critique, citing empirical work as well as more recent work using techniques from behavioral economics.[52] These behavioral economic techniques have been adapted to test predictions of cultural evolutionary models in laboratory settings [53][54] as well as studying differences in cooperation in fifteen small-scale societies in the field.[55] Since one of the goals of DIT is to explain the distribution of human cultural traits, ethnographic and ethnologic techniques may also be useful for testing hypothesis stemming from DIT. Although findings from traditional ethnologic studies have been used to buttress DIT arguments,[56][57] thus far there have been little ethnographic fieldwork designed to explicitly test these hypotheses.[58] Herb Gintis has named DIT one of the two major conceptual theories with potential for unifying the behavioral sciences, including economics, biology, anthropology, sociology, psychology and political science. Because it addresses both the genetic and cultural components of human inheritance, Gintis sees DIT models as providing the best explanations for the ultimate cause of human behavior and the best paradigm for integrating those disciplines with evolutionary theory.[59] In a review of competing evolutionary perspectives on human behavior, Laland and Brown see DIT as the best candidate for uniting the other evolutionary perspectives under one theoretical umbrella.[60]

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Relation to other fields Sociology and cultural anthropology Two major topics of study in both sociology and cultural anthropology are human cultures and cultural variation. However, Dual Inheritance theorists charge that both disciplines too often treat culture as a static superorganic entity that dictates human behavior.[61][62] Cultures are defined by a suite of common traits shared by a large group of people. DIT theoriests argue that this doesn't sufficiently explain variation in cultural traits at the individual level. By contrast, DIT models human culture at the individual level and views culture as the result of a dynamic evolutionary process at the population level.[63]

Human sociobiology and evolutionary psychology Human sociobiologists and evolutionary psychologists try to understand how maximizing genetic fitness, in either the modern era or past environments, can explain human behavior. When faced with a common and seemingly maladaptive trait, practitioners from these disciplines try to determine how the trait actually increases genetic fitness (maybe through kin selection or by speculating about early evolutionary environments). Dual Inheritance theorists, in contrast, will consider a variety of genetic and cultural processes in addition to natural selection on genes.

Human behavioral ecology Human behavioral ecology (HBE) and DIT have a similar relationship to what ecology and evolutionary biology have in the biological sciences. HBE is more concerned about ecological process and DIT more focused on historical process. One difference is that human behavioral ecologists often assume that culture is a system that produces the most adaptive outcome in a given environment. This implies that similar behavioral traditions should be found in similar environments. However, this is not always the case. A study of African cultures showed that cultural history was a better predictor of cultural traits than local ecological conditions.[64]

Memetics Memetics, which comes from the meme idea described in Dawkins's The Selfish Gene, is similar to DIT in that it treats culture as an evolutionary process that is distinct from genetic transmission. However, there are some philosophical differences between memetics and DIT.[65] One difference is that memetics' focus is on the selection potential of discrete replicators (memes), where DIT allows for transmission of both non-replicators and non-discrete cultural variants. DIT does not assume that replicators are necessary for cumulative adaptive evolution. DIT also more strongly emphasizes the role of genetic inheritance in shaping the capacity for cultural evolution. But perhaps the biggest difference is a difference in academic lineage. Memetics as a label is more influential in popular culture than in academia. Critics of memetics argue that it is lacking in empirical support or is conceptually ill-founded, and question whether there is hope for the memetic research program succeeding. Proponents point out that many cultural traits are discrete, and that many existing models of cultural inheritance assume discrete cultural units, and hence involve memes.[66]

Criticisms A number of criticisms of DIT have been put forward.[67][68][69] From some points of view, use of the term ‘dual inheritance’ to refer to both what is transmitted genetically and what is transmitted culturally is technically misleading.[citation needed] Many opponents cite horizontal transmission of ideas to be so "different" from the typical vertical transmission (reproduction) in genetic evolution that it is not evolution. However, 1)even genetic evolution uses non-vertical transmission through the environmental alteration of the genome during life by acquired circumstance: epigenetics, and 2) genetic evolution is also affected by direct horizontal transmission between separate species of plants and strains of bacteria: horizontal gene transfer. Other critics argue that there can be no

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Dual inheritance theory "dual" inheritance without cultural inheritance being "sequestered" by the biotic genome [citation needed]. Evidence for this process is scarce and controversial. Why this is a demand of critics, however, can be considered unclear as it refutes none of the central claims laid down by proponents of DIT. More serious criticisms of DIT arise from the choice of Darwinian selection as an explanatory framework for culture. Some argue, cultural evolution does not possess the algorithmic structure of a process that can be modeled in a Darwinian framework as characterized by John von Neumann[70] and used by John Holland to design the genetic algorithm.[71] Forcing culture into a Darwinian framework gives a distorted picture of the process for several reasons. First, some argue Darwinian selection only works as an explanatory framework when variation is randomly generated.[citation needed] To the extent that transmission biases are operative in culture, they mitigate the effect of Darwinian change, i.e. change in the distribution of variants over generations of exposure to selective pressures.[citation needed] Second, since acquired change can accumulate orders of magnitude faster than inherited change, if it is not getting regularly discarded each generation, it quickly overwhelms the population-level mechanism of change identified by Darwin; it ‘swamps the phylogenetic signal’. [citation needed] However, DIT proponents might reply that, 1) biotic evolution does not function only on randomly generated phenotypes either, since the phenotypes present in a population are the combined result of random and selective effects during the last generation; and 2)that transmission bias would quite often also reinforce "Darwinian change" since it is widely evidenced that Culture has adaptive value in increasing human fitness. Another discord in opinion stems from DIT opponents' assertion that there exists some "creative force" that is applied to each idea as it is received and before it is passed on, and that this agency is so powerful that it can be stronger than the selective system of other individuals assessing what to teach and whether your idea has merit [citation needed] . But if this criticism was valid then it would be comparatively much easier to argue an unpopular or incorrect concepts than it actually is. In addition, nothing about DIT runs counter to the idea that an internally selective process (some would call creativity) also determines the fitness of ideas received and sent. In fact this decision making is a large part of the territory embraced by DIT proponents but is poorly understood due to limitations in neurobiology (for more information see Neural Darwinism). Related criticisms of the effort to frame culture in Darwinian terms have been leveled by Richard Lewontin,[72] Niles Eldredge,[73] and Stuart Kauffman.[74]

References [1] McElreath, R. & Henrich, J. 2007. Dual inheritance theory: the evolution of human cultural capacities and cultural evolution. (http:/ / arbeit. ucdavis. edu/ mcelreath/ files/ Henrich and McElreath final. pdf) In R. Dunbar and L. Barrett, (Eds.), Oxford Handbook of Evolutionary Psychology Oxford: Oxford University Press. [2] Simoons, F. 1969. Primary adult lactose intolerance and the milking habit: A problem in biologic and cultural interrelations: I. Review of the medical research. The American Journal of Digestive Diseases 14:819-836. [3] Simoons, F. 1970. Primary adult lactose intolerance and the milking habit: A problem in biologic and cultural interrelations: II. A culture historical hypothesis. The American Journal of Digestive Diseases 15:695-710. [4] Cavalli-Sforza, L., P. Menozzi and A. Piazza. 1994. The history and geography of human genes Princeton: Princeton University Press [5] Holden, C. and R. Mace. 1997. Phylogenetic analysis of the evolution of lactose digestion in adults. Human Biology 69:605-628. [6] Durham, W. 1991. Coevolution: Genes, culture and human diversity. Stanford: Stanford University Press. Chapter 5 [7] Perry, G., N. Dominy, K. Claw, A. Lee, H> Fiegler, R. Redon, J. Werner, F. Villanea, J. Mountain, R. Misra, N. Carter, C. Lee, A. Stone. Diet and the evolution of human amylase gene copy number variation. Nature Genetics 39:1256-1260. [8] Boyd, R. and P. J. Richerson. 1985. Culture and the Evolutionary Process. Chicago: University of Chicago Press. pp. 199-202. [9] Richerson, P. J. and R. Boyd. 2005. Not By Genes Alone: How Culture Transformed Human Evolution. Chicago: University of Chicago Press. pp. 169-182. [10] Kroeberm A. and C. Kluckhohn. 1952. Culture; A Critical Review of Concepts and Definitions. Cambridge, MA: Harvard University. [11] Fox, R. and B. King. 2002. Anthropology Beyond Culture Oxford: Berg. [12] Richerson, P. and R. Boyd. 2005. Not By Genes Alone: How Culture Transformed Human Evolution Chicago: University of Chicago Press. pg 6. [13] Boyd, R. and P. Richerson. 2000. Memes: Universal Acid or a Better Mouse Trap? In R. Aunger, Ed. Darwinizing Culture: The Status of Memetics as a Science. Oxford: Oxford University Press. pp. 143-162. Full text (http:/ / www. sscnet. ucla. edu/ anthro/ faculty/ boyd/ CambMeme. PDF)

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Dual inheritance theory [14] Richerson, P.J. and R. Boyd. 2001. Culture is Part of Human Biology: Why the Superorganic Concept Serves the Human Sciences Badly. In Science Studies: Probing the Dynamics of Scientific Knowledge, In S. Maasen and M. Winterhager, Ed. Bielefeld: Verlag. (http:/ / xcelab. net/ rm/ wp-content/ uploads/ 2008/ 08/ cultureisbiology. pdf) [15] Richerson, P. and R. Boyd. 2005. Not By Genes Alone: How Culture Transformed Human Evolution Chicago: University of Chicago Press. pg 7. [16] Sasaki, J. Y. (2013). Promise and Challenges Surrounding Culture–Gene Coevolution and Gene–Culture Interactions. (http:/ / www. tandfonline. com/ doi/ full/ 10. 1080/ 1047840X. 2013. 764814) Psychological Inquiry, 24(1), 64-70. [17] Laland, K. N. and G. R. Brown. 2002. Sense & Nonsense: Evolutionary Perspectives on Human Behavior. Oxford: Oxford University Press. pg. 260 [18] Cochran, G. and H. Harpending. 2009. The 10,000 Year Explosion: How Civilization Accelerated Human Evolution. Basic Books. [19] Koerper, H. and E. Stickel. 1980. Cultural Drift: A Primary Process of Culture Change Journal of Anthropological Research. 36: 463-469. [20] Cavalli-Sfornza, L. and M. Feldman. 1981. Cultural Transmission and Evolution: A Quantitative Approach. Princeton, New Jersey: Princeton University Press. [21] Bentley, R.A., M.W. Hahn and S.J. Shennan. 2004. Random drift and culture change. Proceedings of the Royal Society B 271: 1443-1450. Full text (http:/ / artsci. wustl. edu/ ~pboyer/ CEwebsite/ Archive/ ShennanRandomDrift. pdf) [22] Hahn, M.W. and R. A. Bentley. 2003. Drift as a mechanism for cultural change: An example from baby names. Proceedings of the Royal Society B 271: S353-S356. Full text (http:/ / www. duke. edu/ ~mwh3/ BabyNames. pdf) [23] Boyd, R. and P. J. Richerson. 1985. Culture and the Evolutionary Process. Chicago: University of Chicago Press. pp.9, 69 [24] P.J.B. Slater, V.M. Janik. Vocal Learning. In "Encyclopedia of Animal Behavior", 2010, Pages 551-557. http:/ / www. sciencedirect. com/ science/ referenceworks/ 9780080453378 [25] Gabora, L. (1995). Meme and variations: A computer model of cultural evolution (http:/ / www. vub. ac. be/ CLEA/ liane/ papers/ mav. htm). In (L. Nadel & D. Stein, Eds.) 1993 Lectures in Complex Systems, Addison-Wesley, 471-486. [26] Feldman, M. and L. Cavalli-Sfornza. 1976. Cultural and biological evolutionary processes, selection for a trait under complex transmission. Theoretical Population Biology 9: 238-259. [27] Feldman, M and L. Cavalli-Sfornza. 1977. The evolution of continuous variation: II, complex transmission and assortive mating. Theoretical Population Biology 11:161-181. [28] Boyd, R., and P. Richerson. 1985. Culture and the Evolutionary Process. Chicago: The University of Chicago Press. [29] Henrich, J. and R. McElreath. 2007. Dual inheritance theory: the evolution of human cultural capacities and cultural evolution. Oxford Handbook of Evolutionary Psychology, R. Dunbar and L. Barrett, eds., Ch. 38. Oxford: Oxford Univ Press. [30] Henrich, J. and R. McElreath. 2003. The evolution of cultural evolution. (http:/ / arbeit. ucdavis. edu/ mcelreath/ files/ henrich mcelreath EA 2003. pdf) Evolutionary Anthropology 12:123-135. [31] Richerson, P.J. and R. Boyd. 2000. Climate, culture, and the evolution of cognition. In C.M. Heyes and L. Huber, (Eds), The Evolution of Cognition. Massachusetts: MIT Press. [32] Tomasello, M. 1999. The Cultural Origins of Human Cognition. Cambridge, Massachusetts: Cambridge University Press. [33] Richerson, P. J. and R. Boyd. 2000. The Pleistocene and the origins of human culture: built for speed. Perspectives in Ethology 13:1-45, 2000. [34] Tomasello, M. 1999. The Cultural Origins of Human Cognition. Cambridge, MA: Harvard University Press. [35] Williams, G.C. 1972. Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Princeton: Princeton University Press. ISBN 0-691-02357-3 [36] Williams, G.C.. 1986. Evolution Through Group Selection. Blackwell. ISBN 0-632-01541-1 [37] Maynard Smith, J.. 1964. Group selection and kin selection 'Nature' 201:1145–1147 [38] Uyenoyama, M. and M. W. Feldman. 1980. Theories of kin and group selection: a population genetics perspective. Theoretical Population Biology 17:380-414. [39] Cavalli-Sforza, L. L. and M. W. Feldman. 1973. Models for cultural inheritance. I. Group mean and within group variation. Theoretical Population Biology 4:42-44. [40] Boyd, R. and P. J. Richerson. 1985. Culture and the Evolutionary Process. Chicago: University of Chicago Press. pg. 227-240. [41] Soltis, J., Boyd, R. and P. J. Richerson. 1995. Can group-functional behaviors evolve by cultural group selection? An empirical test (http:/ / www. des. ucdavis. edu/ faculty/ richerson/ SoltisBoydRichersonCA95. pdf) Current Anthropology 36:473-494 [42] Mace, R., C. Holden, and S. Shennan (Eds.) 2005. The evolution of cultural diversity: a phylogenetic approach. London:University College London Press. [43] Darwin, C. 1874. The descent of man and selection in relation to sex. 2nd ed. 2 vols. New York: American Home Library. [44] Campbell, D. 1965. Variation and selective retention in socio-cultural evolution. In Social change in developing areas: A reinterpretation of evolutionary theory, ed. H. Barringer, G. Blanksten, and R. Mack, 19-49. Cambridge, MA: Schenkman Publishing Company. [45] Feldman, M. and Cavalli-Sforna L. 1976. Cultural and biological evolutionary processes, selection for a trait under complex transmission. Theoretical Population Biology, 9:238-59. [46] Lumsden C., and E. Wilson. 1981. Genes, Mind and Culture: The Coevolutionary Process. Cambridge, MA: Harvard University Press. [47] Laland K. and G. Brown. 2002. Sense and Nonsense: Evolutionary Perspectives on Human Behavior. Oxford: Oxford University Press. [48] Boyd, R. and Richerson, P. 1983. The cultural transmission of acquired variation: effects on genetic fitness. Journal of Theoretical Biology 100:567-96.

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Dual inheritance theory [49] Boyd, R. and P. J. Richerson. 2005. The Origin and Evolution of Cultures. Oxford: Oxford University Press. pg. 294-299. [50] Haag, Allison. 2006. The synthesizer (http:/ / seedmagazine. com/ news/ 2007/ 01/ the_synthesizer. php). SEED, 2(7): 46. [51] Laland, K. N. and G. R. Brown. 2002. Sense & Nonsense: Evolutionary Perspectives on Human Behavior. Oxford: Oxford University Press. pg 290. [52] Herb Gintis Amazon.com review: http:/ / www. amazon. com/ review/ product/ 0198508840/ [53] McElreath, R., M. Lubell, P. J. Richerson, T. M. Waring, W. Baum, E. Edsten, C. Efferson, and B. Paciotti. 2005. Applying formal models to the laboratory study of social learning: The impact of task difficulty and environmental fluctuation. (http:/ / www. des. ucdavis. edu/ faculty/ Richerson/ WheatPotatoes. pdf) Evolution and Human Behavior 26: 483-508. [54] Baum, W. M., P. J. Richerson, C. M. Efferson, B. M. Paciotti. 2004. Cultural evolution in laboratory micro-societies including traditions of rule-giving and rule-following (http:/ / www. des. ucdavis. edu/ faculty/ Richerson/ Baumetalprinted. pdf). Evolution and Human Behavior 25: 305-326. 2004. [55] Henrich, J., R. Boyd, S. Bowles, C. Camerer, E. Fehr, H. Gintis (Eds). 2004. Foundations of Human Sociality: Economic Experiments and Ethnographic Evidence from Fifteen Small-Scale Societies Oxford: Oxford University Press. [56] Cavalli-Sfornza, L. L. and M. Feldman. 1981. Cultural Transmission and Evolution: A Quantitative Approach. Princeton, New Jersey: Princeton University Press. [57] Boyd, R. and P. J. Richerson. 1985. Culture and the Evolutionary Process. Chicago: University of Chicago Press. [58] McElreath, R. 2004. Social learning and the maintenance of cultural variation: An evolutionary model and data from East Africa (http:/ / arbeit. ucdavis. edu/ mcelreath/ files/ mcelreath AA 2004. pdf). American Anthropologist 106:308-321. [59] Gintis, H. 2006. Behavioral and Brain Sciences A framework for the integration of the behavioral sciences (http:/ / www. umass. edu/ preferen/ gintis/ Unity-BBS Print Version. pdf) Behavioral and Brain Sciences 30:1-61 [60] Laland, K. N. and G. R. Brown. 2002. Sense & Nonsense: Evolutionary Perspectives on Human Behavior. Oxford: Oxford University Press. pg. 287-319. [61] Richerson, P. and R. Boyd. 2001. Culture is part of human biology: Why the superorganic concept serves the human sciences badly (http:/ / www. des. ucdavis. edu/ faculty/ Richerson/ CultureIsBiology. pdf). In M. Goodman and A. S. Moffat(Eds.) Probing Human Origins. Cambridge, Massachusetts: The American Academy of Arts & Sciences. [62] Gintis, H. 2007. A framework for the unification of the behavioral sciences (http:/ / www. umass. edu/ preferen/ gintis/ Unity-BBS Print Version. pdf). Behavioral and Brain Sciences. 30:1-61. [63] Richerson, P. J. and R. Boyd. 2005. Not By Genes Alone: How Culture Transformed Human Evolution. Chicago: University of Chicago Press. pg. 5-8 [64] Guglielmino, C. R., Viganotti, C., Hewlett, B., and Cavalli-Sforza, L.L. 1995. Cultural variation in Africa: role of mechanism of transmission and adaptation. Proceedings of the National Academy of Sciences USA 92:7585-7589. [65] Boyd, R. and P.J. Richerson. 2000. Memes: universal acid or better mouse trap (http:/ / www. des. ucdavis. edu/ faculty/ Richerson/ CambMeme. PDF). In R. Aunger (Ed), Darwinizing Culture: The Status of Memetics as a Science. Oxford: Oxford University Press. pg 143–162. [66] Laland, K. N. and G. R. Brown. 2002. Sense & Nonsense: Evolutionary Perspectives on Human Behavior. Oxford: Oxford University Press. pg. 289-290. [67] Gabora, L. (2008). The cultural evolution of socially situated cognition (http:/ / www. vub. ac. be/ CLEA/ liane/ papers/ ssc. htm). Cognitive Systems Research, 9(1-2), 104-113. [68] Gabora, L. (2011). Five clarifications about cultural evolution (https:/ / people. ok. ubc. ca/ lgabora/ papers/ gabora-five-JOCC2011. pdf). Journal of Cognition and Culture, 11, 61-83. [69] Gabora, L. (2011). How a generation was misled about natural selection (http:/ / www. psychologytoday. com/ blog/ mindbloggling/ 201105/ how-generation-was-misled-about-natural-selection). Psychology Today, Mindbloggling. [70] von Neumann, J. (1966). The theory of self-reproducing automata. University of Illinois Press, Chicago. [71] Holland, J. (1975). Adaptation in natural and artificial systems. Cambridge: MIT Press. [72] Fracchia, J., & Lewontin, R. C. (1999). Does culture evolve? History and Theory, 38, 52−78. [73] Temkin, I. & Eldredge, N. (2007). Phylogenetics and material cultural evolution. Current Anthropology, 48(1), 146-153. [74] Kauffman, S. (1999). Darwinism, neoDarwinism, and the autocatalytic model of culture: Commentary on Origin of Culture (http:/ / www. cogsci. ecs. soton. ac. uk/ cgi/ psyc/ newpsy?10. 022), Psycoloquy, 10(22), 1−4.

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Further reading Books • Lumsden, C. J. and E. O. Wilson. 1981. Genes, Mind, and Culture: The Coevolutionary Process. Cambridge, Massachusetts: Harvard University Press. • Cavalli-Sforza, L. L. and M. Feldman. 1981. Cultural Transmission and Evolution: A Quantitative Approach. Princeton, New Jersey: Princeton University Press. • Boyd, R. and P. J. Richerson. 1985. Culture and the Evolutionary Process. Chicago: University of Chicago Press. • Durham, W. H. 1991. Coevolution: Genes, Culture and Human Diversity. Stanford, California: Stanford University Press. ISBN 0-8047-1537-8 • Tomasello, M. 1999. The Cultural Origins of Human Cognition. Cambridge, Massachusetts: Cambridge University Press. • Shennan, S. J. 2002. Genes, Memes and Human History: Darwinian Archaeology and Cultural Evolution. London: Thames and Hudson. • Laland, K. N. and G. R. Brown. 2002. Sense & Nonsense: Evolutionary Perspectives on Human Behavior. Oxford: Oxford University Press. • Boyd, R. and P. J. Richerson. 2005. The Origin and Evolution of Cultures. Oxford: Oxford University Press. • Richerson, P. J. and R. Boyd. 2005. Not By Genes Alone: How Culture Transformed Human Evolution. Chicago: University of Chicago Press.

Reviews • Smith, E. A. 1999. Three styles in the evolutionary analysis of human behavior. (http://faculty.washington.edu/ easmith/ThreeStyles.pdf) In L. Cronk, N. Chagnon, and W. Irons, (Eds.) Adaptation and Human Behavior: An Anthropological Perspective New York: Aldine de Gruyter. • Henrich, J. and R. McElreath. 2003. The evolution of cultural evolution. (http://arbeit.ucdavis.edu/mcelreath/ files/henrich mcelreath EA 2003.pdf) Evolutionary Anthropology 12:123-135. • Mesoudi, A., A. Whiten, and K. N. Laland. 2006. Towards a unified science of cultural evolution. (http:// amesoudi.googlepages.com/Mesoudi_Whiten_Laland_BBS_2006.pdf) Behavioral and Brain Sciences 29:329-383 • Gintis, H. 2006. A framework for the integration of the behavioral sciences (http://www.umass.edu/preferen/ gintis/Unity-BBS Print Version.pdf) Behavioral and Brain Sciences 30:1-61 • Bentley, R.A., C. Lipo, H.D.G. Maschner and B. Marler 2007. Darwinian Archaeologies. In R.A. Bentley, H.D.G. Maschner & C. Chippendale (Eds.) Handbook of Archaeological Theories. Lanham (MD): AltaMira Press. • McElreath, R. & Henrich, J. 2007. Modeling cultural evolution. (http://arbeit.ucdavis.edu/mcelreath/files/ mcelreath_henrich_mce_final.pdf) In R. Dunbar and L. Barrett, (Eds.), Oxford Handbook of Evolutionary Psychology Oxford: Oxford University Press. • McElreath, R. & Henrich, J. 2007. Dual inheritance theory: the evolution of human cultural capacities and cultural evolution. (http://arbeit.ucdavis.edu/mcelreath/files/Henrich and McElreath final.pdf) In R. Dunbar and L. Barrett, (Eds.), Oxford Handbook of Evolutionary Psychology Oxford: Oxford University Press. • Sterelny, Kim (2002). Review Genes, Memes and Human History (http://www.vuw.ac.nz/phil/staff/ documents/sterelny-papers/tripod.pdf#search='dual inheritance theorypdf'). Stephen Shennan. London: Thames and Hudson. p. 304. • Laland, K.N., Odling-Smee, J. & Myles, S. 2010. How culture shaped the human genome: bringing genetics and the human sciences. Nature Reviews: Genetics 11:137-148

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Journal articles • "Culture, Adaptation, and Innateness" (http://www.sscnet.ucla.edu/anthro/faculty/boyd/Innateness ver 4.1. pdf). The Innate Mind: Culture and Cognition. |coauthors= requires |author= (help) • "Built for Speed, Not for Comfort: Darwinian Theory and Human Culture" (http://www.sscnet.ucla.edu/ anthro/faculty/boyd/Built for Speed, Not for Comfort.pdf). History and Philosophy of the Life Sciences (23): 425–465. 2001. |coauthors= requires |author= (help)

External links Current DIT researchers • Rob Boyd (http://www.sscnet.ucla.edu/anthro/faculty/boyd/), Department of Anthropology, UCLA • Marcus Feldman (http://www-evo.stanford.edu/marc.html), Department of Biological Sciences, Stanford • Joe Henrich (http://www.psych.ubc.ca/~henrich/home.html), Departments of Psychology and Economics, University of British Columbia • Richard McElreath (http://arbeit.ucdavis.edu/mcelreath/), Anthropology Department, UC Davis • Peter J. Richerson (http://www.des.ucdavis.edu/faculty/Richerson/Richerson.htm), Department of Environmental Science and Policy, UC Davis

Related researchers • Liane Gabora (https://people.ok.ubc.ca/lgabora/), Department of Psychology, University of British Columbia • Herb Gintis (http://people.umass.edu/gintis/), Emeritus Professor of Economics, University of Massachusetts & Santa Fe Institute • Kevin Laland (http://lalandlab.st-andrews.ac.uk/), School of Biology, University of St. Andrews • Ruth Mace (http://www.ucl.ac.uk/heeg/), Department of Anthropology, University College London • Michael Tomasello (http://email.eva.mpg.de/~tomas/), Department of Developmental and Comparative Psychology, Max Planck Institute for Evolutionary Anthropology

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Evolutionary ethics is a term referring equally to a form of descriptive ethics or normative ethics. Descriptive evolutionary ethics consists of biological approaches to ethics (morality) based on the role of evolution in shaping human psychology and behavior. Such approaches may be based in scientific fields such as evolutionary psychology, sociobiology, or ethology with a focus on understanding and explaining observed ethical preferences or choices and their origins. On the other hand, normative evolutionary ethics may represent a more independent attempt to use evolution, alone or partially, to justify an ethical system. This project has not, according to one view, been especially successful; for example, Richard Dawkins describes how we must rise above our selfish genes to behave morally (that is, evolution has endowed us with various instincts, but we need some other moral system to decide which ones to empower or control). Dawkins has since expressed interest in what Sam Harris calls a science of morality, which starts with the assumption that "morality" refers to "facts about the flourishing of conscious creatures".

History In the chapter On the Development of the Intellectual and Moral Faculties During Primeval and Civilised Times of The descent of man (1871) Charles Darwin set out to explain the origin of human morality in order to show that there was no absolute gap between man and animals. For Darwin, morality was a problem of natural history. He believed that a moral sense (altruism) would have little selective advantage for the individual, but it would be adaptive for the group. He did not construct a new system of Evolutionary Ethics.[1] David Hume first described what is now known as the is-ought problem: making unjustified claims about what ought to be on the basis of statements about what is. The problem is the justification of an ethical system. The problem is not what we ought to do, but why. Thomas Huxley allows that ethical sentiments have evolved but denies that this

Evolutionary ethics provides a basis for morality (Evolution and Ethics,1893): The propounders of what are called the "ethics of evolution," when the "evolution of ethics" would usually better express the object of their speculations, adduce a number of more or less interesting facts and more or less sound arguments, in favour of the origin of the moral sentiments, in the same way as other natural phenomena, by a process of evolution. I have little doubt, for my part, that they are on the right track; but as the immoral sentiments have no less been evolved, there is, so far, as much natural sanction for the one as the other. The thief and the murderer follow nature just as much as the philanthropist. Cosmic evolution may teach us how the good and the evil tendencies of man may have come about; but, in itself, it is incompetent to furnish any better reason why what we call good is preferable to what we call evil than we had before.[2] Huxley's criticism alluded to the is-ought problem developed earlier by David Hume and the related naturalistic fallacy developed later by G. E. Moore. The moral philosopher Henry Sidgwick (1838–1900) claimed that evolution was irrelevant for ethics because it could not be used as a justification for ethics. British philosopher G. E. Moore (Principia Ethica) demonstrated that all systems of naturalistic ethics, including evolutionary ethics, are flawed. He first pointed out that even if evolution is progress, it cannot be concluded that the more advanced organisms are more advanced in every respect. So, it is impossible to infer particular moral judgements from that fact. Furthermore, the view that "we ought to move in the direction of evolution simply because it is the direction of evolution" was invalid because it was an example of the naturalistic fallacy, that is the fallacy of defining 'the good' by reference to some other thing. American philosopher William James wrote about natural selection: "The entire modern deification of survival per se, survival returning into itself, survival naked and abstract, with the denial of any substantive excellence in what survives, except for more survival still, is surely the strangest intellectual stopping-place ever proposed by one man to another".[3] John Dewey was also a critic of evolutionary ethics, although both philosophers accepted the fact of evolution. Dewey added that the discovery of the evolutionary origin of particular moral sentiments is not identical with the discovery of the foundation of an ethical system.[4] Evolutionary biologist and geneticist Theodosius Dobzhansky was highly critical of evolutionary ethics: "No theory of evolutionary ethics can be acceptable unless it gives a satisfactory explanation of just why the promotion of evolutionary development must be regarded as the summum bonum" and "even if the direction of evolution were demonstrated to be "good", man is likely to prefer to be free rather than to be reasonable".[5]

Analytic philosophy Logical positivist philosopher A. J. Ayer stated in Language, Truth and Logic (1936) that moral judgements are pure expressions of feeling. They are unverifiable and cannot be true or false. In 1986, Michael Ruse summarized the role of evolution as the source of ethical feelings: Our moral sense, our altruistic nature, is an adaptation—a feature helping us in the struggle for existence and reproduction—no less than hands and eyes, teeth and feet. It is a cost-effective way of getting us to cooperate, which avoids both the pitfalls of blind action and the expense of a superbrain of pure rationality.[6] In applying science to metaethics, Ruse writes: In a sense … the evolutionist's case is that ethics is a collective illusion of the human race, fashioned and maintained by natural selection in order to promote individual reproduction. … ethics is illusory inasmuch as it persuades us that it has an objective reference. This is the crux of the biological position.[7]Wikipedia:Verifiability

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Descriptive evolutionary ethics Descriptive evolutionary ethics is empirical research into moral attitudes and beliefs (humans) or moral behaviour (animals) in an evolutionary framework. Examples can be found in the field of evolutionary psychology. Evolutionary psychology attempts to explain major features of psychology in terms of species-wide evolved (via natural selection) predispositions. Ethical topics addressed include altruistic behaviors, deceptive or harmful behaviors, an innate sense of fairness or unfairness, feelings of kindness or love, self-sacrifice, feelings related to competitiveness and moral punishment or retribution, moral "cheating" or hypocrisy, and inclinations for a wide variety of actions judged morally good or bad by (at least some within) a given society. A key issue of evolutionary psychology has been how altruistic feelings and behaviors could have evolved when the process of natural selection is based on the multiplication over time only of those genes that adapt better to changes in the environment of the species. Theories addressing this have included kin selection and reciprocal altruism (both direct and indirect, and on a society-wide scale). Group selection theories have also been advanced.

Normative evolutionary ethics Normative evolutionary ethics aims at defining which acts are right or wrong, and which things are good or bad in an evolutionary context. It is not merely describing, but it is prescribing goals, values and obligations. For example eugenics is a form of normative evolutionary ethics, because it defines what is "good" on the basis of genetics and the theory of evolution. Social Darwinism is a more wide ranging topic. However, to the extent it promotes ethical values and policies based on the theory of evolution, it can also be classified as a normative evolutionary ethics. According to philosopher G. E. Moore (see above) all systems of naturalistic ethics, including normative evolutionary ethics, do commit the naturalistic fallacy. The naturalistic fallacy does not apply to descriptive evolutionary ethics because no ethical statements are inferred from facts. Also, the naturalistic fallacy does not apply to weaker forms of normative evolutionary ethics, namely those which are consistent with evolution, but not derivable from evolution.[citation needed]

Criticisms P. G. Woolcock argues[8] that all normative evolutionary ethics are invalid. For example the argument 1. The human species can survive only if we let severely physically and mentally handicapped infants and children die. 2. Therefore: we ought to let severely physically and mentally handicapped infants die. is a fallacy because the first statement is a purely descriptive premise containing no values, and a value pops up in the conclusion. It is the famous naturalistic fallacy (G. E. Moore). Additionally, the first premise is almost certainly false. We could make the argument valid by adding a second premise, namely: 1b We ought to do whatever is necessary to ensure the survival of the human species but then we would no longer be deducing a value conclusion from a purely factual premise, because 1b has a value component. This can also be explained in this way: if the definition of "good" is "whatever furthers human survival", then it should be nonsensical to ask "Is human survival itself good?", but it seems a perfectly meaningful question. This is Moore's open-question argument. Another fallacy according to Woolcock is the confusion between an instrumental and a categorical justification. Consider the argument "We ought to be altruistic because evolution has selected altruism over millions of years as a reliable guide to what is good". Evolutionary theory, however, tell us only that altruism is good for the survival of our species, not that the survival of the human species is good. A categorical justification would justify people's actions regardless of what their goals are.

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The future Given the current state of knowledge, Huxley's statement with regards to "why what we call good is preferable to what we call evil" is still accurate with regards to individual human tastes and predispositions. Yet research in the fields of evolutionary psychology and primatology is beginning to reveal, in the general case, what is good and bad for our species in order for it to thrive and, in turn, more likely be happy. Even so, it can never say (in principle) why the species should prefer happiness to misery, but simply that it does (or does not) so prefer. Hence, evolutionary psychology's primary focus is to derive, especially through the deep analysis of hunter-gatherer culture and primate models, what is the most accurate description of general human predispositions (i.e. our innate "hard-wiring"). And as this understanding grows, it will become more and more feasible to redesign culture itself to be more "user friendly" to its human members, according to some standard . After all, in the ultimate sense, culture (like a computer) is a tool to serve its users. Noted primatologist Frans De Waal asserts, "In the words of Edward Wilson, biology holds us "on a leash" and will let us stray only so far from who we are. We can design our life any way we want, but whether we will thrive depends on how well the life fits human predispositions" [9] Thus, the goals of evolutionary psychology overlap with the science of morality.

Notes [1] [2] [3] [4] [5] [6]

Paul Lawrence Farber, 1994, The Temptations of Evolutionary Ethics, chapter 1 Huxley, p. 66 Quoted by Farber, 1994, p. 112 Farber, 1994, p.111-117 Theodosius Dobzhansky, The Biological Basis of Human Freedom, Columbia University Press, 1956, p. 128-129. Ruse, M. (1986). Evolutionary Ethics: A phoenix arisen. Zygon Journal of Religion and Science, 21, p. 99. Available online at http:/ / www. philoscience. unibe. ch/ documents/ educational_materials/ Ruse1986/ Ruse1986. pdf [7] Ruse, 1986, p. 235 [8] Peter G. Woolcock, 'The Case against Evolutionary Ethics Today' in: Biology and the Foundation of Ethics, Cambridge University Press, 1999, pp 276-306 [9] Frans de Waal, "The empathic ape" (http:/ / www. annular. org/ ~sdbrown/ the-empathic-ape. html), New Scientist, 8 October 2005

References • Huxley, Thomas Henry (1893). "Evolution and Ethics" (http://aleph0.clarku.edu/huxley/CE9/E-E.html). In Nitecki, Matthew H.; Nitecki, Doris V. Evolutionary Ethics. Albany: State University of New York (published 1993). ISBN 0-7914-1499-X • Ruse, Michael (1995). "Evolutionary Ethics: A Phoenix Arisen". In Thompson, Paul. Issues in Evolutionary Ethics. Albany: State University of New York. ISBN 0-7914-2027-2

Further reading • Curry, O. (2006). Who's afraid of the naturalistic fallacy? Evolutionary Psychology, 4, 234-247. Full text (http:// www.epjournal.net/filestore/ep04234247.pdf) • Dawkins, Richard (1976). The Selfish Gene. ISBN 1-155-16265-X. • Duntley, J.D., & Buss, D.M. (2004). The evolution of evil. In A. Miller (Ed.), The social psychology of good and evil. New York: Guilford. 102-123. Full text (http://homepage.psy.utexas.edu/homepage/Group/BussLAB/ pdffiles/The evolution of evil.pdf) • Hauser, Marc (2006). Moral Minds. ISBN 0-06-078070-3. • Huxley, Julian. Evolutionary Ethics 1893-1943. Pilot, London. In USA as Touchstone for ethics Harper, N.Y. (1947) [includes text from both T.H. Huxley and Julian Huxley]

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Evolutionary ethics • Katz, L. (Ed.) Evolutionary Origins of Morality: Cross-Disciplinary Perspectives (http://human-nature.com/ nibbs/02/levy.html) Imprint Academic, 2000 ISBN 0-907845-07-X • Kitcher, Philip (1995) “Four Ways of “Biologicizing” Ethics” in Elliott Sober (ed.) Conceptual Issues in Evolutionary Biology, The MIT Press • Kitcher, Philip (2005) “Biology and Ethics” in David Copp (ed.) The Oxford Handbook of Ethical Theory, Oxford University Press • Krebs, D. L. & Denton, K. (2005). Toward a more pragmatic approach to morality: A critical evaluation of Kohlberg’s model. Psychological Review, 112, 629-649. Full text (http://www.sfu.ca/psyc/faculty/krebs/ publications/Toward a More Pragmatic Approach to Morality.pdf) • Krebs, D. L. (2005). An evolutionary reconceptualization of Kohlberg’s model of moral development. In R. Burgess & K. MacDonald (Eds.) Evolutionary Perspectives on Human Development, (pp. 243–274). CA: Sage Publications. Full text (http://www.sfu.ca/psyc/faculty/krebs/publications/MacDonald.pdf) • Mascaro, S., Korb, K.B., Nicholson, A.E., Woodberry, O. (2010). Evolving Ethics: The New Science of Good and Evil. Exeter, UK: Imprint Academic. • Richerson, P.J. & Boyd, R. (2004). Darwinian Evolutionary Ethics: Between Patriotism and Sympathy. In Philip Clayton and Jeffrey Schloss, (Eds.), Evolution and Ethics: Human Morality in Biological and Religious Perspective, pp. 50–77. Full text (http://www.des.ucdavis.edu/faculty/richerson/Dethics.pdf) ISBN 0-8028-2695-4 • Ridley, Matt (1996). The Origins of Virtue. Viking. ISBN 0-14-026445-0. • Ruse, Michael (1993). "The New Evolutionary Ethics". In Nitecki, Matthew H.; Nitecki, Doris V. Evolutionary Ethics. Albany: State University of New York. ISBN 0-7914-1499-X • Shermer, Michael (2004). The Science of Good and Evil: Why People Cheat, Gossip, Care, Share, and Follow the Golden Rule. New York: Henry Holt and Company. ISBN 0-8050-7520-8. • Teehan, J. & diCarlo, C. (2004). On the Naturalistic Fallacy: A conceptual basis for evolutionary ethics. Evolutionary Psychology, 2, 32-46. Full text (http://www.epjournal.net/filestore/ep023246.pdf) • de Waal, Frans (1996). Good Natured: The Origins of Right and Wrong in Humans and Other Animals. London: Harvard University Press. ISBN 0-674-35660-8. • Walter, A. (2006). The anti-naturalistic fallacy: Evolutionary moral psychology and the insistence of brute facts. Evolutionary Psychology, 4, 33-48. Full text (http://www.epjournal.net/filestore/ep043348.pdf) • Wilson, D. S., E. Dietrich, et al. (2003). On the inappropriate use of the naturalistic fallacy in evolutionary psychology. Biology and Philosophy 18: 669-682. Full text (http://biology.binghamton.edu/dwilson/Wilson publications/DSW14.pdf) • Wilson, D. S. (2002). Evolution, morality and human potential. Evolutionary Psychology: Alternative Approaches. S. J. Scher and F. Rauscher, Kluwer Press: 55-70 Full text (http://biology.binghamton.edu/ dwilson/Wilson publications/DSW16.pdf) • Wilson, E. O. (1979). On Human Nature. ISBN 0-671-54130-7. • Wright, Robert (1995). The Moral Animal. ISBN 0-679-40773-1.

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External links • The Evolution of Ethics: An Introduction to Cybernetic Ethics (http://www.evolutionaryethics.com/) by S. E. Bromberg • Evolutionary Ethics (http://www.iep.utm.edu/e/evol-eth.htm) at the Internet Encyclopedia of Philosophy • Morality and Evolutionary Biology (http://plato.stanford.edu/entries/morality-biology) entry by William FitzPatrick in the Stanford Encyclopedia of Philosophy • Biological Altruism (http://plato.stanford.edu/entries/altruism-biological) entry by Samir Okasha in the Stanford Encyclopedia of Philosophy

Evolutionary anthropology Part of a series on

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Evolutionary anthropology is the interdisciplinary study of the evolution of human physiology and human behaviour and the relation between hominids and non-hominid primates. Evolutionary anthropology is based in natural science and social science. Various fields and disciplines are: • • • • • • •

Human evolution and anthropogeny. Paleoanthropology and paleontology of primates. Primatology and Primate Ethology and Paleontology. The sociocultural evolution of human behavior. The evolutionary psychology of humans. The archaeological study of human technology and change over time and space. Human evolutionary genetics and changes in the human genome over time.

• The cognitive neuroscience and neuroanthropology of human and primate cognition, culture and actions and abilities.

Evolutionary anthropology • Human behavioural ecology and the interaction of humans and the environment. • Studies of human anatomy, endocrinology, and neurobiology and differences and changes between species, variation between human groups, and relationships to cultural factors. Evolutionary anthropology is concerned with both biological and cultural evolution of humans, past and present. It is based on a scientific approach, and brings together fields such as archaeology, behavioral ecology, psychology, primatology, and genetics. It is a dynamic and interdisciplinary field, drawing on many lines of evidence to understand the human experience, past and present. Studies of biological evolution generally concern the evolution of the human form. Cultural evolution involves the study of cultural change over time and space and frequently incorporate Cultural transmission models. Note that cultural evolution is not the same as biological evolution, and that human culture involves the transmission of cultural information, which behaves in ways quite distinct from human biology and genetics. The study of cultural change is increasingly performed through cladistics and genetic models.

External links • Evolutionary Anthropology Society [1] • Max Planck Institute for Evolutionary Anthropology [2] • • • • • • • • • • • • •

Evolutionary Anthropology Journal [3] Institute of Human Origins at Arizona State University [4] Becoming Human Program of Institute of Human Origins at Arizona State University [5] Institute of Cognitive and Evolutionary Anthropology of the University of Oxford [6] Evolutionary Anthropology at California State University, Fullerton [7] Centre for Human Evolutionary Studies at Rutgers, State University of New Jersey [8] Graduate Program in Evolutionary Anthropology at Rutgers, State University of New Jersey [9] Evolutionary Anthropology Program at Washington State University [10] Evolutionary Anthropology at University of New Mexico [11] Evolutionary Anthropology at Duke University [12] Evolutionary Anthropology Research Group at Durham University [13] Department of Human Evolutionary Biology at Harvard University [14] Human and Evolutionary Biology Program at University of Southern California [15]

References [1] http:/ / www. anth. uconn. edu/ eas/ [2] http:/ / www. eva. mpg. de/ [3] http:/ / www3. interscience. wiley. com/ journal/ 38641/ toc [4] http:/ / iho. asu. edu/ [5] http:/ / www. becominghuman. org/ [6] http:/ / www. icea. ox. ac. uk/ [7] http:/ / anthro. fullerton. edu/ evolanth/ [8] http:/ / evolution. rutgers. edu/ [9] http:/ / anthro. rutgers. edu/ graduate-program/ prospective-students/ 196?task=view [10] http:/ / libarts. wsu. edu/ anthro/ gradevol. html [11] http:/ / www. unm. edu/ ~anthro/ programs_evolutionary_anthropology. html [12] http:/ / evolutionaryanthropology. duke. edu/ [13] http:/ / www. dur. ac. uk/ anthropology/ research/ earg/ [14] http:/ / www. heb. fas. harvard. edu/ [15] http:/ / dornsife. usc. edu/ bisc/ heb/ home/ index. cfm

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Evolutionary biology Part of a series on

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Evolutionary biology is a sub-field of biology concerned with the study of the evolutionary processes that produced the diversity of life on Earth. Someone who studies evolutionary biology is known as an evolutionary biologist. Evolutionary biologists study the descent of species, and the origin of new species.

Subfields The study of evolution is the unifying concept in evolutionary biology. Evolutionary biology is a conceptual subfield of biology that intersects with other subfields that are delimited by organizational level (e.g., cell biology, population biology), taxonomic level (e.g., zoology, Graphical representation of the modern "Tree of ornithology, herpetology) or angle of approach (e.g., field biology, Life on the Web" project. theoretical biology, experimental evolution, paleontology). Usually, these intersections are combined into specific fields such as evolutionary ecology and evolutionary developmental biology.

Evolutionary biology

History Evolutionary biology, as an academic discipline in its own right, emerged as a result of the modern evolutionary synthesis in the 1930s and 1940s.[1] It was not until the 1970s and 1980s, however, that a significant number of universities had departments that specifically included the term evolutionary biology in their titles. In the United States, as a result of the rapid growth of molecular and cell biology, many universities have split (or aggregated) their biology departments into molecular and cell biology-style departments and ecology and evolutionary biology-style departments (which often have subsumed older departments in botany, zoology and the like). The subdiscipline of paleontology is often found in earth science/geology/geoscience departments. Microbiology has recently developed into an evolutionary discipline. It was originally ignored due to the paucity of morphological traits and the lack of a species concept in microbiology. Now, evolutionary researchers are taking advantage of our extensive understanding of microbial physiology, the ease of microbial genomics, and the quick generation time of some microbes to answer evolutionary questions. Similar features have led to progress in viral evolution, particularly for bacteriophages.

Important evolutionary biologists Many biologists have contributed to our current understanding of evolution. The establishment of evolutionary biology as a professional scientific discipline only started with the development of population genetics and the formulation of the modern evolutionary synthesis. Theodosius Dobzhansky and E. B. Ford were particularly important in the establishment of an empirical research programme for evolutionary biology. Ernst Mayr, George Gaylord Simpson and G. Ledyard Stebbins were also important discipline-builders during the modern synthesis, in the fields of systematics, paleontology and botany, respectively. Through training many future evolutionary biologists, James Crow, Richard Lewontin, Dan Hartl, Marcus Feldman, and Brian Charlesworth have also made large contributions to building the discipline of evolutionary biology.

Journals Some scientific journals specialize exclusively in evolutionary biology as a whole, including the journals Evolution, Journal of Evolutionary Biology, and BMC Evolutionary Biology. Some journals cover sub-specialties within evolutionary biology, such as the journals Systematic Biology, Molecular Biology and Evolution and its sister journal Genome Biology and Evolution, and Cladistics. Other journals combine aspects of evolutionary biology with other related fields. For example, Molecular Ecology, Proceedings of the Royal Society of London Series B, The American Naturalist and Theoretical Population Biology have overlap with ecology and other aspects of organismal biology. Overlap with ecology is also prominent in the review journals Trends in Ecology and Evolution and Annual Review of Ecology, Evolution, and Systematics. The journals Genetics and PLoS Genetics overlap with molecular genetics questions that are not obviously evolutionary in nature.

Current research topics Current research in evolutionary biology covers diverse topics, as should be expected given the centrality of evolution to understanding biology. Modern evolutionary biology incorporates ideas from diverse areas of science, such as molecular genetics and even computer science. First, some fields of evolutionary research try to explain phenomena that were poorly accounted for by the work of the modern evolutionary synthesis. These phenomena include speciation, the evolution of sexual reproduction, the evolution of cooperation, the evolution of ageing, and evolvability.

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Evolutionary biology Second, biologists ask the most straightforward evolutionary question: "what happened and when?". This includes fields such as paleobiology, as well as systematics and phylogenetics. Third, the modern evolutionary synthesis was devised at a time when nobody understood the molecular basis of genes. Today, evolutionary biologists try to determine the genetic architecture of interesting evolutionary phenomena such as adaptation and speciation. They seek answers to questions such as how many genes are involved, how large are the effects of each gene, to what extent are the effects of different genes interdependent, what sort of function do the genes involved tend to have, and what sort of changes tend to happen to them (e.g., point mutations vs. gene duplication or even genome duplication). Evolutionary biologists try to reconcile the high heritability seen in twin studies with the difficulty in finding which genes are responsible for this heritability using genome-wide association studies. One challenge in studying genetic architecture is that the classical population genetics that catalyzed the modern evolutionary synthesis must be updated to take into account modern molecular knowledge. This requires a great deal of mathematical development to relate DNA sequence data to evolutionary theory as part of a theory of molecular evolution. For example, biologists try to infer which genes have been under strong selection by detecting selective sweeps. Fourth, the modern evolutionary synthesis involved agreement about which forces contribute to evolution, but not about their relative importance. Current research seeks to determine this. Evolutionary forces include natural selection, sexual selection, genetic drift, genetic draft, developmental constraints, mutation bias and biogeography. An evolutionary approach is also key to much current research in biology that does not set out to study evolution per se, especially in organismal biology and ecology. For example, evolutionary thinking is key to life history theory. Annotation of genes and their function relies heavily on comparative, i.e., evolutionary, approaches. The field of evo-devo investigates how developmental processes work by using the comparative method to determine how they evolved.

References [1] Sterelny (2009) p.314

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Evolutionary developmental psychology, (or EDP), is the application of the basic principles of Darwinian evolution, particularly natural selection, to explain contemporary human development. It involves the study of the genetic and environmental mechanisms that underlie the universal development of social and cognitive competencies and the evolved epigenetic (gene-environment interactions) processes that adapt these competencies to local conditions. It assumes that not only are behaviors and cognitions that characterize adults the product of natural selection pressures operating over the course of evolution, but so also are characteristics of children's behaviors and minds. It further proposes that an evolutionary account would provide some insight into not only predictable stages of ontogeny, but into specific differences between individuals as well. Such a perspective suggests that there are multiple alternative strategies to recurring problems that human children would have faced throughout our evolutionary past and that individual differences in developmental patterns aren’t necessarily idiosyncratic reactions, but are predictable, adaptive responses to environmental pressures.

Brief history Traditionally, evolutionary psychologists tended to focus their research and theorizing primarily on adults, especially on behaviors related to socializing and mating. There was much less of a focus on psychological development, as it relates to Darwinian evolution. Developmental psychologists, for their part, have been wary of the perceived genetic determinism of evolutionary thinking, which seemed critical of all the major theories in developmental psychology. Pioneers of EDP have worked to integrate evolutionary and developmental theories, without totally discarding the traditional theories of either. They argue that a greater understanding of the “whys” of human development will help us acquire a better understanding of the “hows” and “whats” of human development.

Some basic assumptions 1. All evolutionarily-influenced characteristics develop, and this requires examining not only the functioning of these characteristics in adults but also their ontogeny. 2. All evolved characteristics develop via continuous and bidirectional gene-environment interactions that emerge dynamically over time. 3. Development is constrained by genetic, environmental, and cultural factors. 4. An extended childhood is needed in which to learn the complexities of human social communities and economies. 5. Many aspects of childhood serve as preparations for adulthood and were selected over the course of evolution (deferred adaptations). 6. Some characteristics of infants and children were selected to serve an adaptive function at specific times in development and not as preparations for adulthood (ontogenetic adaptations). 7. Children show a high degree of plasticity, or flexibility, and the ability to adapt to different contexts.

Evolutionary developmental psychology

Domain-specificity vs. domain-generality A fundamental issue is how best to characterize the cognitive mechanisms that afford humans such flexibility in problem-solving. Authors Leda Cosmides and John Tooby would argue that human beings simply possess a greater number of content-specific modules, each of which specializes in solving a specific type of adaptive problem. And it is the sheer number of these content-specific modules which lends humans such great problem-solving flexibility. Other authors, such as Robert Burgess and Kevin B. MacDonald, while agreeing that content-specific modules exist, favor a differing view. They would say instead that the flexibility of human problem-solving ability is owed primarily to powerful domain-generality, and that humans use the same non-specific cognitive machinery for a multitude of different tasks. It is also important to point out that this is not an either/or argument for the legitimacy of the domain-specific or the domain-general position, but is concerned simply with the importance of both in regards to our problem-solving capabilities.

Relevant journals • Evolution and Development [1] Research relevant to interface of evolutionary and developmental biology

Further reading • Bjorklund, D.F.; Pellegrini, A.D. (2002). The Origins of Human Nature: Evolutionary Developmental Psychology [2] . Washington, D.C.: American Psychological Association. • Bjorklund, D.F.; Pellegrini, A.D. (2000). "Child Development and Evolutionary Psychology" [3]. Child Development 71 (6): 1687–1708. doi:10.1111/1467-8624.00258 [4]. PMID 11194266 [5]. • Boyce, W.T. & Ellis, B.J. (2005). Biological sensitivity to context: I. An evolutionary-developmental theory of the origins and functions of stress reactivity. Development & Psychopathology, 17, 271-301. Full text [6] • Burgess, R. L. & MacDonald (Eds.) (2004). Evolutionary Perspectives on Human Development, 2nd ed [7]. Thousand Oaks, CA: Sage Publications. • Burman, J. T. (in press). Experimenting in relation to Piaget: Education is a Chaperoned Process of Adaptation. Perspectives on Science, 16(2). • Ellis, B.J., & Bjorklund, D.F. (Eds.) (2005). Origins of the social mind: Evolutionary psychology and child development [8]. New York: Guilford Press. • Ellis, B.J., Essex, M.J., & Boyce, W.T. (2005). Biological sensitivity to context: II. Empirical explorations of an evolutionary-developmental theory. Development & Psychopathology 17, 303-328. Full text [9] • Ellis, B.J. (2004). Timing of pubertal maturation in girls: An integrated life history approach. Psychological Bulletin, 130, 920-958. Full text [10] • Flinn M.V. (2004). Culture and developmental plasticity: Evolution of the social brain. In K. MacDonald and R. L. Burgess (Eds.), Evolutionary Perspectives on Human Development. Chapter 3, pp. 73-98. Thousand Oaks, CA: Sage. Full text [11] • Flinn, M.V. & Ward, C.V. (2004). Ontogeny and Evolution of the Social Child. In B. Ellis & D. Bjorklund (Eds.), Origins of the social mind: Evolutionary psychology and child development. Chapter 2, pp. 19-44. London: Guilford Press. Full text [12] • Geary, David C. (2006). "Evolutionary developmental psychology: Current status and future directions" [13]. Developmental Review 26 (2): 113–119. doi:10.1016/j.dr.2006.02.005 [14]. • Geary, D. C. (2005). Folk knowledge and academic learning. In B. J. Ellis & D. F. Bjorklund (Eds.), Origins of the social mind. (pp. 493-519). New York: Guilford Publications. Full text [15] • Geary, D. C. (2004). Evolution and cognitive development. In R. Burgess & K. MacDonald (Eds.), Evolutionary perspectives on human development (pp. 99-133). Thousand Oaks, CA: Sage Publications. Full text [16] • Geary, D. C., Byrd-Craven, J., Hoard, M. K., Vigil, J., & Numtee, C. (2003). Evolution and development of boys’ social behavior. Developmental Review, 23, 444-470. Full text [17]

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Evolutionary developmental psychology • Geary, D.C., & Bjorklund, D.F. (2000). Evolutionary Developmental Psychology. Child Development, 71, 57-65. Full text [18] • MacDonald, K. (2005). Personality, Evolution, and Development. In R. Burgess and K. MacDonald (Eds.), Evolutionary Perspectives on Human Development, 2nd edition, pp. 207–242. Thousand Oaks, CA: Sage. Full text [19] • MacDonald, K., & Hershberger, S. (2005). Theoretical Issues in the Study of Evolution and Development. In R. Burgess and K. MacDonald (Eds.), Evolutionary Perspectives on Human Development, 2nd edition, pp. 21–72. Thousand Oaks, CA: Sage. Full text [20] • Maestripieri, D. & Roney, J.R. (2006). Evolutionary developmental psychology: Contributions from comparative research with nonhuman primates. Developmental Review, 26, 120-137. Full text [21] • Medicus G. (1992). The Inapplicability of the Biogenetic Rule to Behavioral Development. Human Development 35, 1-8. Full text [22] • Robert, J. S. Taking old ideas seriously: Evolution, development, and human behavior. New Ideas in Psychology. • Vigil, J. M., Geary, D. C., & Byrd-Craven, J. (2005). A life history assessment of early childhood sexual abuse in women. Developmental Psychology, 41, 553-561. Full text [23]

References [1] http:/ / www3. interscience. wiley. com/ journal/ 118546131/ home [2] http:/ / www. apa. org/ books/ 431671A. html [3] http:/ / bernard. pitzer. edu/ ~dmoore/ psych199s03articles/ Bjorklund. pdf [4] http:/ / dx. doi. org/ 10. 1111%2F1467-8624. 00258 [5] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 11194266 [6] http:/ / ag. arizona. edu/ fcs/ fshd/ people/ ellis/ DPBoyceEllis2005. pdf [7] http:/ / www. csulb. edu/ ~kmacd/ HEB_2005_3. pdf [8] http:/ / www. guilford. com/ cgi-bin/ cartscript. cgi?page=pr/ ellis. htm& dir=pp/ dp& cart_id=208191. 21056 [9] http:/ / ag. arizona. edu/ fcs/ fshd/ people/ ellis/ DPEllisEssexBoyce2005. pdf [10] http:/ / ag. arizona. edu/ fcs/ fshd/ people/ ellis/ Psyc%20Bull%20Ellis%202004. pdf [11] http:/ / www. missouri. edu/ ~anthmark/ pdf/ Ch_3--Mark_Flinn. pdf [12] http:/ / www. missouri. edu/ ~anthmark/ pdf/ Flinn& WardE& B2004. pdf [13] http:/ / web. missouri. edu/ ~gearyd/ files/ Geary%20Final%20Evo%20Dev%20Psy%20Issue. pdf [14] http:/ / dx. doi. org/ 10. 1016%2Fj. dr. 2006. 02. 005 [15] http:/ / web. missouri. edu/ ~psycorie/ FolkKnowledgePDF. pdf [16] http:/ / web. missouri. edu/ ~psycorie/ EvoCogDev%5BChap%5D. pdf [17] http:/ / web. missouri. edu/ ~psycorie/ DevelRev03. pdf [18] http:/ / www. missouri. edu/ ~psycorie/ EvoDevPsy. pdf#search='evolutionary%20developmental%20psychology' [19] http:/ / www. csulb. edu/ ~kmacd/ BURG%2008%20ED. pdf [20] http:/ / www. csulb. edu/ ~kmacd/ BURG%2002%20ED. pdf [21] http:/ / primate. uchicago. edu/ 2006DR. pdf [22] http:/ / homepage. uibk. ac. at/ ~c720126/ humanethologie/ ws/ medicus/ block6/ HumanDevelopment. pdf#search=' [23] http:/ / web. missouri. edu/ ~psycorie/ Vigiletal%5BDP2005%5D. pdf

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Evolutionary neuroscience is an interdisciplinary scientific research field that studies the evolution of nervous systems. Evolutionary neuroscientists attempt to understand the evolution and natural history of nervous system structure and function. The field draws on concepts and findings from both neuroscience and evolutionary biology. Historically, most empirical work has been in the area of comparative neuroanatomy, and modern studies often make use of phylogenetic comparative methods. Selection experiments and experimental evolution approaches are also being used more frequently. Conceptually and theoretically, the field is related to fields as diverse as comparative psychology, neuroethology, developmental neurobiology, evo-devo, behavioral ecology, biological anthropology, sociobiology, cognitive neuroscience, sociocultural evolution and evolutionary psychology.

Evolutionary neuroscience

History The field began after the publication of Darwin's On the Origin of Species, but brain evolution was largely viewed at the time in relation to the incorrect scala naturae. The 1936 book The Comparative Anatomy of the Nervous System of Vertebrates was a landmark publication in the field. Following the Evolutionary Synthesis, the study of comparative neuroanatomy was conducted with an evolutionary view, and modern studies incorporate developmental genetics.

Researchers • • • • • • •

Glenn Northcutt Terrence Deacon Georg F. Striedter Jon Kaas John Allman William H. Calvin Merlin Donald

References External links • Brain Behavior and Evolution (http://content.karger.com/ProdukteDB/produkte. asp?Aktion=JournalGuidelines&ProduktNr=223831) - (Journal) • "Comparative Vertebrate Neuroanatomy: Evolution and Adaptation" (http://www.wiley.com/WileyCDA/ WileyTitle/productCd-0471733830.html) - Ann B. Butler, William Hodos • Sinauer.com (http://www.sinauer.com/detail.php?id=8206) - Principles of Brain Evolution Georg F. Striedter, University of California, Irvine' (book review, 2004)

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Evolutionary psychology (EP) is an approach in the social and natural sciences that examines psychological traits such as memory, perception, and language from a modern evolutionary perspective. It seeks to identify which human psychological traits are evolved adaptations – that is, the functional products of natural selection or sexual selection. Adaptationist thinking about physiological mechanisms, such as the heart, lungs, and immune system, is common in evolutionary biology. Some evolutionary psychologists apply the same thinking to psychology, arguing that the mind has a modular structure similar to that of the body, with different modular adaptations serving different functions. Evolutionary psychologists argue that much of human behavior is the output of psychological adaptations that evolved to solve recurrent problems in human ancestral environments.[1] The adaptationist approach is steadily increasing as an influence in the general field of psychology. Evolutionary psychologists suggest that EP is not simply a subdiscipline of psychology but that evolutionary theory can provide a foundational, metatheoretical framework that integrates the entire field of psychology, in the same way it has for biology.[2][3] Evolutionary psychologists hold that behaviors or traits that occur universally in all cultures are good candidates for evolutionary adaptations[] including the abilities to infer others' emotions, discern kin from non-kin, identify and prefer healthier mates, and cooperate with others. They report successful tests of theoretical predictions related to such topics as infanticide, intelligence, marriage patterns, promiscuity, perception of beauty, bride price, and parental investment.[4]

Evolutionary psychology The theories and findings of EP have applications in many fields, including economics, environment, health, law, management, psychiatry, politics, and literature.[5][6] Controversies concerning EP involve questions of testability, cognitive and evolutionary assumptions (such as modular functioning of the brain, and large uncertainty about the ancestral environment), importance of non-genetic and non-adaptive explanations, as well as political and ethical issues due to interpretations of research results.

Scope Principles Evolutionary psychology is an approach that views human nature as the product of a universal set of evolved psychological adaptations to recurring problems in the ancestral environment. Proponents of EP suggest that it seeks to integrate psychology into the other natural sciences, rooting it in the organizing theory of biology (evolutionary theory), and thus understanding psychology as a branch of biology. Anthropologist John Tooby and psychologist Leda Cosmides note: "Evolutionary psychology is the long-forestalled scientific attempt to assemble out of the disjointed, fragmentary, and mutually contradictory human disciplines a single, logically integrated research framework for the psychological, social, and behavioral sciences—a framework that not only incorporates the evolutionary sciences on a full and equal basis, but that systematically works out all of the revisions in existing belief and research practice that such a synthesis requires."[7] Just as human physiology and evolutionary physiology have worked to identify physical adaptations of the body that represent "human physiological nature," the purpose of evolutionary psychology is to identify evolved emotional and cognitive adaptations that represent "human psychological nature." According to Steven Pinker, EP is "not a single theory but a large set of hypotheses" and a term that "has also come to refer to a particular way of applying evolutionary theory to the mind, with an emphasis on adaptation, gene-level selection, and modularity." Evolutionary psychology adopts an understanding of the mind that is based on the computational theory of mind. It describes mental processes as computational operations, so that, for example, a fear response is described as arising from a neurological computation that inputs the perceptional data, e.g. a visual image of a spider, and outputs the appropriate reaction, e.g. fear of possibly dangerous animals. While philosophers have generally considered the human mind to include broad faculties, such as reason and lust, evolutionary psychologists describe evolved psychological mechanisms as narrowly focused to deal with specific issues, such as catching cheaters or choosing mates. EP views the human brain as comprising many functional mechanisms,[citation needed] called psychological adaptations or evolved cognitive mechanisms or cognitive modules, designed by the process of natural selection. Examples include language-acquisition modules, incest-avoidance mechanisms, cheater-detection mechanisms, intelligence and sex-specific mating preferences, foraging mechanisms, alliance-tracking mechanisms, agent-detection mechanisms, and others. Some mechanisms, termed domain-specific, deal with recurrent adaptive problems over the course of human evolutionary history.[citation needed] Domain-general mechanisms, on the other hand, are proposed to deal with evolutionary novelty.[citation needed] EP has roots in cognitive psychology and evolutionary biology but also draws on behavioral ecology, artificial intelligence, genetics, ethology, anthropology, archaeology, biology, and zoology. EP is closely linked to sociobiology, but there are key differences between them including the emphasis on domain-specific rather than domain-general mechanisms, the relevance of measures of current fitness, the importance of mismatch theory, and psychology rather than behavior. Most of what is now labeled as sociobiological research is now confined to the field of behavioral ecology.[citation needed] Nikolaas Tinbergen's four categories of questions can help to clarify the distinctions between several different, but complementary, types of explanations.[8] Evolutionary psychology focuses primarily on the "why?" questions, while traditional psychology focuses on the "how?" questions.[9]

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Sequential vs. Static Perspective Historical/Developmental Explanation of current form in terms of a historical sequence How vs. Why Questions

Current Form Explanation of the current form of species

Proximate Ontogeny How an individual organism's Developmental explanations for changes in structures function individuals, from DNA to their current form

Mechanism Mechanistic explanations for how an organism's structures work

Evolutionary Why a species evolved the structures (adaptations) it has

Adaptation A species trait that evolved to solve a reproductive or survival problem in the ancestral environment

Phylogeny The history of the evolution of sequential changes in a species over many generations

Premises Evolutionary psychology is founded on several core premises. 1. The brain is an information processing device, and it produces behavior in response to external and internal inputs.[10] 2. The brain's adaptive mechanisms were shaped by natural and sexual selection. 3. Different neural mechanisms are specialized for solving problems in humanity's evolutionary past. 4. The brain has evolved specialized neural mechanisms that were designed for solving problems that recurred over deep evolutionary time, giving modern humans stone-age minds. 5. Most contents and processes of the brain are unconscious; and most mental problems that seem easy to solve are actually extremely difficult problems that are solved unconsciously by complicated neural mechanisms. 6. Human psychology consists of many specialized mechanisms, each sensitive to different classes of information or inputs. These mechanisms combine to produce manifest behavior.

History Evolutionary psychology has its historical roots in Charles Darwin’s theory of natural selection. In The Origin of Species, Darwin predicted that psychology would develop an evolutionary basis: In the distant future I see open fields for far more important researches. Psychology will be based on a new foundation, that of the necessary acquirement of each mental power and capacity by gradation. —Charles Darwin, The Origin of Species, 1859, p. 449.

Nobel Laureates Nikolaas Tinbergen (left) and

Two of his later books were devoted to the study of animal emotions Konrad Lorenz (right) who were, with Karl von Frisch, acknowledged for work on animal and psychology; The Descent of Man, and Selection in Relation to Sex behavior in 1871 and The Expression of the Emotions in Man and Animals in 1872. Darwin's work inspired William James’s functionalist approach to psychology. Darwin's theories of evolution, adaptation, and natural selection have provided insight into why brains function the way they do. The content of EP has derived from, on one hand, the biological sciences (especially evolutionary theory as it relates to ancient human environments, the study of paleoanthropology and animal behavior) and, on the other, the human sciences, especially psychology.

Evolutionary psychology Evolutionary biology as an academic discipline emerged with the modern evolutionary synthesis in the 1930s and 1940s.[11] In the 1930s the study of animal behavior (ethology) emerged with the work of Dutch biologist Nikolaas Tinbergen and Austrian biologists Konrad Lorenz and Karl von Frisch. W.D. Hamilton's (1964) papers on inclusive fitness and Robert Trivers's (1972) theories on reciprocity and parental investment helped to establish evolutionary thinking in psychology and the other social sciences. In 1975, Edward O. Wilson combined evolutionary theory with studies of animal and social behavior, building on the works of Lorenz and Tinbergen, in his book Sociobiology: The New Synthesis. In the 1970s, two major branches developed from ethology. Firstly, the study of animal social behavior (including humans) generated sociobiology, defined by its pre-eminent proponent Edward O. Wilson in 1975 as "the systematic study of the biological basis of all social behavior"[12] and in 1978 as "the extension of population biology and evolutionary theory to social organization."[13] Secondly, there was behavioral ecology which placed less emphasis on social behavior by focusing on the ecological and evolutionary basis of both animal and human behavior. In the 1970s and 1980s university departments began to include the term evolutionary biology in their titles. The modern era of evolutionary psychology was ushered in, in particular, by Donald Symons' 1979 book The Evolution of Human Sexuality and Leda Cosmides and John Tooby's 1992 book The Adapted Mind. From psychology there are the primary streams of developmental, social and cognitive psychology. Establishing some measure of the relative influence of genetics and environment on behavior has been at the core of behavioral genetics and its variants, notably studies at the molecular level that examine the relationship between genes, neurotransmitters and behavior. Dual inheritance theory (DIT), developed in the late 1970s and early 1980s, has a slightly different perspective by trying to explain how human behavior is a product of two different and interacting evolutionary processes: genetic evolution and cultural evolution. DIT is seen by some as a "middle-ground" between views that emphasize human universals versus those that emphasize cultural variation.[14]

Theoretical foundations The theories on which evolutionary psychology is based originated with Charles Darwin's work, including his speculations about the evolutionary origins of social instincts in humans. Modern evolutionary psychology, however, is possible only because of advances in evolutionary theory in the 20th century. Evolutionary psychologists say that natural selection has provided humans with many psychological adaptations, in much the same way that it generated humans' anatomical and physiological adaptations.[15] As with adaptations in general, psychological adaptations are said to be specialized for the environment in which an organism evolved, the environment of evolutionary adaptedness, or EEA.[16] Sexual selection provides organisms with adaptations related to mating. For male mammals, which have a relatively high maximal potential reproduction rate, sexual selection leads to adaptations that help them compete for females. For female mammals, with a relatively low maximal potential reproduction rate, sexual selection leads to choosiness, which helps females select higher quality mates. Charles Darwin described both natural selection and sexual selection, and he relied on group selection to explain the evolution of altruistic (self-sacrificing) behavior. But group selection was considered a weak explanation, because in any group the less altruistic individuals will be more likely to survive, and the group will become less self-sacrificing as a whole. In 1964, William D. Hamilton proposed inclusive fitness theory, emphasizing a "gene's-eye" view of evolution. Hamilton noted that individuals can increase the replication of their genes into the next generation by helping close relatives with whom they share genes survive and reproduce. According to "Hamilton's rule", a self-sacrificing behavior can evolve if it helps close relatives so much that it more than compensates for the individual animal's sacrifice. Inclusive fitness theory resolved the issue of how "altruism" evolved. Other theories also help explain the evolution of altruistic behavior, including evolutionary game theory, tit-for-tat reciprocity, and generalized reciprocity. These theories not only help explain the development of altruistic behavior, but also account for hostility toward cheaters (individuals that take advantage of others' altruism).

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Evolutionary psychology Several mid-level evolutionary theories inform evolutionary psychology. The r/K selection theory proposes that some species prosper by having many offspring, while others follow the strategy of having fewer offspring but investing much more in each one. Humans follow the second strategy. Parental investment theory explains how parents invest more or less in individual offspring based on how successful those offspring are likely to be, and thus how much they might improve the parents' inclusive fitness. According to the Trivers-Willard hypothesis, parents in good conditions tend to invest more in sons (who are best able to take advantage of good conditions), while parents in poor conditions tend to invest more in daughters (who are best able to have successful offspring even in poor conditions). According to life history theory, animals evolve life histories to match their environments, determining details such as age at first reproduction and number of offspring. Dual inheritance theory posits that genes and human culture have interacted, with genes affecting the development of culture, and culture, in turn, affecting human evolution on a genetic level (see also the Baldwin effect).

Evolved psychological mechanisms Evolutionary psychology is based on the hypothesis that, just like hearts, lungs, livers, kidneys, and immune systems, cognition has functional structure that has a genetic basis, and therefore has evolved by natural selection. Like other organs and tissues, this functional structure should be universally shared amongst a species, and should solve important problems of survival and reproduction. Evolutionary psychologists seek to understand psychological mechanisms by understanding the survival and reproductive functions they might have served over the course of evolutionary history.[citation needed] These might include abilities to infer others' emotions, discern kin from non-kin, identify and prefer healthier mates, or cooperate with others. Consistent with the theory of natural selection, evolutionary psychology sees humans as often in conflict with others, including mates and relatives. For instance, when breastfeeding offspring, the mother reaches the point in which it is evolutionary more beneficial to wean the pup earlier than the pup itself does. For the mother, weaning implies that she will be able to conceive the next offspring, but for the pup the future siblings are not as important and therefore it will crave to still be breastfed. Evolutionary psychology also recognizes the role of kin selection and reciprocity in evolving prosocial traits such as altruism. Like chimps and bonobos, humans have subtle and flexible social instincts, allowing them to form extended families, lifelong friendships, and political alliances. In studies testing theoretical predictions, evolutionary psychologists have made modest findings on topics such as infanticide, intelligence, marriage patterns, promiscuity, perception of beauty, bride price and parental investment.

Products of evolution: adaptations, exaptations, byproducts, and random variation Not all traits of organisms are adaptations. As noted in the table below, traits may also be exaptations, byproducts of adaptations (sometimes called "spandrels"), or random variation between individuals.[17] Psychological adaptations are hypothesized to be innate or relatively easy to learn, and to manifest in cultures worldwide. For example, the ability of toddlers to learn a language with virtually no training is likely to be a psychological adaptation. On the other hand, ancestral humans did not read or write, thus today, learning to read and write require extensive training, and presumably represent byproducts of cognitive processing that use psychological adaptations designed for other functions.[18] However, variations in manifest behavior can result from universal mechanisms interacting with different local environments. For example, Caucasians who move from a northern climate to the equator will have darker skin. The mechanisms regulating their pigmentation do not change; rather the input to the those mechanisms change, resulting in different output.

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Adaptation

Exaptation

By-Product

Random Noise

Definition

Organismic trait designed to solve an ancestral problem(s). Shows complexity, special "design", functionality

Adaptation that has been "re-designed" to solve a different adaptive problem.

Byproduct of an adaptive mechanism with no current or ancestral function

Random variations in an adaptation or byproduct

Physiological Example

Bones / Umbilical cord

Small bones of the inner ear

White color of bones / Belly button

Bumps on the skull, convex or concave belly button shape

Psychological Example

Toddlers’ ability to learn to talk with minimal instruction.

Voluntary Attention

Ability to learn to read and write.

Within-sex variations in voice pitch.

One of the tasks of evolutionary psychology is to identify which psychological traits are likely to be adaptations, byproducts or random variation. George C Williams suggested that an "adaptation is a special and onerous concept that should only be used where it is really necessary."[19] As noted by Williams and others, adaptations can be identified by their improbable complexity, species universality, and adaptive functionality.

Obligate and facultative adaptations A question that may be asked about an adaptation is whether it is generally obligate (relatively robust in the face of typical environmental variation) or facultative (sensitive to typical environmental variation).[20] The sweet taste of sugar and the pain of hitting one's knee against concrete are the result of fairly obligate psychological adaptations; typical environmental variability during development does not much affect their operation. By contrast, facultative adaptations are somewhat like "if-then" statements. For example, adult attachment style seems particularly sensitive to early childhood experiences. As adults, the propensity to develop close, trusting bonds with others is dependent on whether early childhood caregivers could be trusted to provide reliable assistance and attention. The adaptation for skin to tan is conditional to exposure to sunlight; this is an example of another facultative adaptation. When a psychological adaptation is facultative, evolutionary psychologists concern themselves with how developmental and environmental inputs influence the expression of the adaptation.

Cultural universals Evolutionary psychologists hold that behaviors or traits that occur universally in all cultures are good candidates for evolutionary adaptations. Cultural universals include behaviors related to language, cognition, social roles, gender roles, and technology.[21] Evolved psychological adaptations (such as the ability to learn a language) interact with cultural inputs to produce specific behaviors (e.g., the specific language learned). Basic gender differences, such as greater eagerness for sex among men and greater coyness among women,[22] are explained as sexually dimorphic psychological adaptations that reflect the different reproductive strategies of males and females.[23] Evolutionary psychologists contrast their approach to what they term the "standard social science model," according to which the mind is a general-purpose cognition device shaped almost entirely by culture.[24][25]

Evolutionary psychology

Environment of evolutionary adaptedness EP argues that to properly understand the functions of the brain, one must understand the properties of the environment in which the brain evolved. That environment is often referred to as the "environment of evolutionary adaptedness" (EEA). The idea of an environment of evolutionary adaptedness was first explored as a part of attachment theory by John Bowlby.[citation needed] This is the environment to which a particular evolved mechanism is adapted. More specifically, the EEA is defined as the set of historically recurring selection pressures that formed a given adaptation, as well as those aspects of the environment that were necessary for the proper development and functioning of the adaptation. Humans, comprising the genus Homo, appeared between 1.5 and 2.5 million years ago, a time that roughly coincides with the start of the Pleistocene 2.6 million years ago. Because the Pleistocene ended a mere 12,000 years ago, most human adaptations either newly evolved during the Pleistocene, or were maintained by stabilizing selection during the Pleistocene. Evolutionary psychology therefore proposes that the majority of human psychological mechanisms are adapted to reproductive problems frequently encountered in Pleistocene environments. In broad terms, these problems include those of growth, development, differentiation, maintenance, mating, parenting, and social relationships. The EEA is significantly different from modern society. The ancestors of modern humans lived in smaller groups, had more cohesive cultures, and had more stable and rich contexts for identity and meaning.[] Researchers look to existing hunter-gatherer societies for clues as to how hunter-gatherers lived in the EEA. Unfortunately, the few surviving hunter-gatherer societies are different from each other, and they have been pushed out of the best land and into harsh environments, so it is not clear how closely they reflect ancestral culture. Evolutionary psychologists sometimes look to chimpanzees, bonobos, and other great apes for insight into human ancestral behavior.[] Christopher Ryan and Cacilda Jetha argue that evolutionary psychologists have overemphasized the similarity of humans and chimps, which are more violent, while underestimating the similarity of humans and bonobos, which are more peaceful.[26]

Mismatches Since an organism's adaptations were suited to its ancestral environment, a new and different environment can create a mismatch. Because humans are mostly adapted to Pleistocene environments, psychological mechanisms sometimes exhibit "mismatches" to the modern environment. One example is the fact that although about 10,000 people are killed with guns in the US annually,[27] whereas spiders and snakes kill only a handful, people nonetheless learn to fear spiders and snakes about as easily as they do a pointed gun, and more easily than an unpointed gun, rabbits or flowers. A potential explanation is that spiders and snakes were a threat to human ancestors throughout the Pleistocene, whereas guns (and rabbits and flowers) were not. There is thus a mismatch between humans' evolved fear-learning psychology and the modern environment. This mismatch also shows up in the phenomena of the supernormal stimulus, a stimulus that elicits a response more strongly than the stimulus for which the response evolved. The term was coined by Niko Tinbergen to refer to non-human animal behavior, but psychologist Deirdre Barrett said that supernormal stimulation governs the behavior of humans as powerfully as that of other animals. She explained junk food as an exaggerated stimulus to cravings for salt, sugar, and fats,[28] and she says that television is an exaggeration of social cues of laughter, smiling faces and attention-grabbing action.[29] Magazine centerfolds and double cheeseburgers pull instincts intended for an EEA where breast development was a sign of health, youth and fertility in a prospective mate, and fat was a rare and vital nutrient.

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Research methods Evolutionary theory is heuristic in that it may generate hypotheses that might not be developed from other theoretical approaches. One of the major goals of adaptationist research is to identify which organismic traits are likely to be adaptations, and which are byproducts or random variations. As noted earlier, adaptations are expected to show evidence of complexity, functionality, and species universality, while byproducts or random variation will not. In addition, adaptations are expected to manifest as proximate mechanisms that interact with the environment in either a generally obligate or facultative fashion (see above). Evolutionary psychologists are also interested in identifying these proximate mechanisms (sometimes termed "mental mechanisms" or "psychological adaptations") and what type of information they take as input, how they process that information, and their outputs. Evolutionary developmental psychology, or "evo-devo," focuses on how adaptations may be activated at certain developmental times (e.g., losing baby teeth, adolescence, etc.) or how events during the development of an individual may alter life history trajectories. Evolutionary psychologists use several strategies to develop and test hypotheses about whether a psychological trait is likely to be an evolved adaptation. Buss (2011)[30] notes that these methods include: Cross-cultural Consistency. Characteristics that have been demonstrated to be cross cultural human universals such as smiling, crying, facial expressions are presumed to be evolved psychological adaptations. Several evolutionary psychologists have collected massive datasets from cultures around the world to assess cross-cultural universality. Function to Form (or "problem to solution"). The fact that males, but not females, risk potential misidentification of genetic offspring (referred to as "paternity insecurity") led evolutionary psychologists to hypothesize that, compared to females, male jealousy would be more focused on sexual, rather than emotional, infidelity. Form to Function (reverse-engineering – or "solution to problem"). Morning sickness, and associated aversions to certain types of food, during pregnancy seemed to have the characteristics of an evolved adaptation (complexity and universality). Margie Profet hypothesized that the function was to avoid the ingestion of toxins during early pregnancy that could damage fetus (but which are otherwise likely to be harmless to healthy non-pregnant women). Corresponding Neurological Modules. Evolutionary psychology and cognitive neuropsychology are mutually compatible – evolutionary psychology helps to identify psychological adaptations and their ultimate, evolutionary functions, while neuropsychology helps to identify the proximate manifestations of these adaptations. Evolutionary psychologists also use various sources of data for testing, including experiments, archaeological records, data from hunter-gatherer societies, observational studies, self-reports and surveys, public records, and human products. Recently, additional methods and tools have been introduced based on fictional scenarios, mathematical models, and multi-agent computer simulations.

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Evolutionary psychology

Major areas of research Foundational areas of research in evolutionary psychology can be divided into broad categories of adaptive problems that arise from the theory of evolution itself: survival, mating, parenting, family and kinship, interactions with non-kin, and cultural evolution.

Survival and individual level psychological adaptations Problems of survival are thus clear targets for the evolution of physical and psychological adaptations.Wikipedia:Please clarify Major problems the ancestors of present day humans faced included food selection and acquisition; territory selection and physical shelter; and avoiding predators and other environmental threats.[31] Consciousness Consciousness is likely an evolved adaptation since it meetsWikipedia:ASF#A_simple_formulation George Williams' criteria of species universality, complexity, and functionality, and it is a trait that apparently increases fitness.[32] In his paper "Evolution of consciousness," John Eccles argues that special anatomical and physical adaptations of the mammalian cerebral cortex gave rise to consciousness. In contrast, others have argued that the recursive circuitry underwriting consciousness is much more primitive, having evolved initially in pre-mammalian species because it improves the capacity for interaction with both social and natural environments by providing an energy-saving "neutral" gear in an otherwise energy-expensive motor output machine.[33] Once in place, this recursive circuitry may well have provided a basis for the subsequent development of many of the functions that consciousness facilitates in higher organisms, as outlined by Bernard J. Baars.[34] Richard Dawkins suggested that humans evolved consciousness in order to make themselves the subjects of thought. Daniel Povinelli suggests that large, tree-climbing apes evolved consciousness to take into account one's own mass when moving safely among tree branches. Consistent with this hypothesis, Gordon Gallup found that chimps and orangutans, but not little monkeys or terrestrial gorillas, demonstrated self-awareness in mirror tests. The concept of consciousness can refer to voluntary action, awareness, or wakefulness. However, even voluntary behavior involves unconscious mechanisms. Many cognitive processes take place in the cognitive unconscious, unavailable to conscious awareness. Some behaviors are conscious when learned but then become unconscious, seemingly automatic. Learning, especially implicitly learning a skill, can take place outside of consciousness. For example, plenty of people know how to turn right when they ride a bike, but very few can accurately explain how they actually do so. Sleep may have evolved to conserve energy when activity would be less fruitful or more dangerous, such as at night, especially in winter.[] Sensation and perception Many experts, such as Jerry Fodor, write that the purpose of perception is knowledge, but evolutionary psychologists hold that its primary purpose is to guide action. For example, they say, depth perception seems to have evolved not to help us know the distances to other objects but rather to help us move around in space. Evolutionary psychologists say that animals from fiddler crabs to humans use eyesight for collision avoidance, suggesting that vision is basically for directing action, not providing knowledge.[] Building and maintaining sense organs is metabolically expensive, so these organs evolve only when they improve an organism's fitness. More than half the brain is devoted to processing sensory information, and the brain itself consumes roughly one-fourth of one's metabolic resources, so the senses must provide exceptional benefits to fitness. Perception accurately mirrors the world; animals get useful, accurate information through their senses.

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Evolutionary psychology Scientists who study perception and sensation have long understood the human senses as adaptations. Depth perception consists of processing over half a dozen visual cues, each of which is based on a regularity of the physical world. Vision evolved to respond to the narrow range of electromagnetic energy that is plentiful and that does not pass through objects. Sound waves go around corners and interact with obstacles, creating a complex pattern that includes useful information about the sources of and distances to objects. Larger animals naturally make lower-pitched sounds as a consequence of their size. The range over which an animal hears, on the other hand, is determined by adaptation. Homing pigeons, for example, can hear very low-pitched sound (infrasound) that carries great distances, even though most smaller animals detect higher-pitched sounds. Taste and smell respond to chemicals in the environment that are thought to have been significant for fitness in the EEA. For example, salt and sugar were apparently both valuable to the human or pre-human inhabitants of the EEA, so present day humans have an intrinsic hunger for salty and sweet tastes. The sense of touch is actually many senses, including pressure, heat, cold, tickle, and pain. Pain, while unpleasant, is adaptive. An important adaptation for senses is range shifting, by which the organism becomes temporarily more or less sensitive to sensation. For example, one's eyes automatically adjust to dim or bright ambient light. Sensory abilities of different organisms often coevolve, as is the case with the hearing of echolocating bats and that of the moths that have evolved to respond to the sounds that the bats make. Evolutionary psychologists claim that perception demonstrates the principle of modularity, with specialized mechanisms handling particular perception tasks. For example, people with damage to a particular part of the brain suffer from the specific defect of not being able to recognize faces (prosopagnosia). EP suggests that this indicates a so-called face-reading module. Learning and facultative adaptations In evolutionary psychology, learning is said to be accomplished through evolved capacities, specifically facultative adaptations.[35] Facultative adaptations express themselves differently depending on input from the environment. Sometimes the input comes during development and helps shape that development. For example, migrating birds learn to orient themselves by the stars during a critical period in their maturation. Evolutionary psychologists claim that humans also learn language along an evolved program, also with critical periods. The input can also come during daily tasks, helping the organism cope with changing environmental conditions. For example, animals evolved Pavlovian conditioning in order to solve problems about causal relationships. Animals accomplish learning tasks most easily when those tasks resemble problems that they faced in their evolutionary past, such as a rat learning where to find food or water. Learning capacities sometimes demonstrate differences between the sexes. In many animal species, for example, males can solve spatial problem faster and more accurately than females, due to the effects of male hormones during development. The same might be true of humans. Emotion and motivation Motivations direct and energize behavior, while emotions provide the affective component to motivation, positive or negative.[36] In the early 1970s, Paul Ekman and colleagues began a line of research that suggests that many emotions are universal. He found evidence that humans share at least five basic emotions: fear, sadness, happiness, anger, and disgust. Social emotions evidently evolved to motivate social behaviors that were adaptive in the EEA. For example, spite seems to work against the individual but it can establish an individual's reputation as someone to be feared. Shame and pride can motivate behaviors that help one maintain one's standing in a community, and self-esteem is one's estimate of one's status.

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Evolutionary psychology Cognition Cognition refers to internal representations of the world and internal information processing. From an EP perspective, cognition is not "general purpose," but uses heuristics, or strategies, that generally increase the likelihood of solving problems that the ancestors of present day humans routinely faced. For example, present day humans are far more likely to solve logic problems that involve detecting cheating (a common problem given humans' social nature) than the same logic problem put in purely abstract terms.[37] Since the ancestors of present day humans did not encounter truly random events, present day humans may be cognitively predisposed to incorrectly identify patterns in random sequences. "Gamblers' Fallacy" is one example of this. Gamblers may falsely believe that they have hit a "lucky streak" even when each outcome is actually random and independent of previous trials. Most people believe that if a fair coin has been flipped 9 times and Heads appears each time, that on the tenth flip, there is a greater than 50% chance of getting Tails. Humans find it far easier to make diagnoses or predictions using frequency data than when the same information is presented as probabilities or percentages, presumably because the ancestors of present day humans lived in relatively small tribes (usually with fewer than 150 people) where frequency information was more readily available. Personality Evolutionary psychology is primarily interested in finding commonalities between people, or basic human psychological nature. From an evolutionary perspective, the fact that people have fundamental differences in personality traits initially presents something of a puzzle.[38] (Note: The field of behavioral genetics is concerned with statistically partitioning differences between people into genetic and environmental sources of variance. However, understanding the concept of heritability can be tricky—heritability refers only to the differences between people, never the degree to which the traits of an individual are due to environmental or genetic factors, since traits are always a complex interweaving of both.) Personality traits are conceptualized by evolutionary psychologists as due to normal variation around an optimum, or due to frequency-dependent selection, or facultative adaptations. Like variability in height, some personality traits may be simply reflect inter-individual variability around a general optimum. Or, personality traits may represent different genetically predisposed "behavioral morphs" – alternate behavioral strategies that depend on the frequency of competing behavioral strategies in the population. For example, if most of the population is generally trusting and gullible, the behavioral morph of being a "cheater" (or, in the extreme case, a sociopath) may be advantageous. Finally, like many other psychological adaptations, personality traits may be facultative—sensitive to typical variations in the social environment, especially during early development. For example, later born children are more likely than first borns to be rebellious, less conscientious and more open to new experiences, which may be advantageous to them given their particular niche in family structure.[39] Language According to Steven Pinker, who builds on the work by Noam Chomsky, the universal human ability to learn to talk between the ages of 1 – 4, basically without training, suggests that language acquisition is a distinctly human psychological adaptation (see, in particular, Pinker's The Language Instinct). Pinker and Bloom (1990) argue that language as a mental faculty shares many likenesses with the complex organs of the body which suggests that, like these organs, language has evolved as an adaptation, since this is the only known mechanism by which such complex organs can develop. Pinker follows Chomsky in arguing that the fact that children can learn any human language with no explicit instruction suggests that language, including most of grammar, is basically innate and that it only needs to be activated by interaction. Chomsky himself does not believe language to have evolved as an adaptation, but suggests that it likely evolved as a byproduct of some other adaptation, a so-called spandrel. But Pinker and Bloom argue that the organic nature of language strongly suggests that it has an adaptational origin.[40]

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Evolutionary psychology Evolutionary psychologists hold that the FOXP2 gene may well be associated with the evolution of human language.[41] In the 1980s, psycholinguist Myrna Gropnik identified a dominant gene that causes language impairment in the KE family of Britain. This gene turned out to be a mutation of the FOXP2 gene. Humans have a unique allele of this gene, which has otherwise been closely conserved through most of mammalian evolutionary history. This unique allele seems to have first appeared between 100 and 200 thousand years ago, and it is now all but universal in humans. However, the once-popular idea that FOXP2 is a 'grammar gene' or that it triggered the emergence of language in Homo sapiens is now widely discredited.[42] Currently several competing theories about the evolutionary origin of language coexist, none of them having achieved a general consensus.[43] Researchers of language acquisition in primates and humans such as Michael Tomasello and Talmy Givón, argue that the innatist framework has understated the role of imitation in learning and that it is not at all necessary to posit the existence of an innate grammar module to explain human language acquisition. Tomasello argues that studies of how children and primates actually acquire communicative skills suggests that humans learn complex behavior through experience, so that instead of a module specifically dedicated to language acquisition, language is acquired by the same cognitive mechanisms that are used to acquire all other kinds of socially transmitted behavior.[44] On the issue of whether language is best seen as having evolved as an adaptation or as a spandrel, evolutionary biologist W. Tecumseh Fitch, following Stephen J. Gould, argues that it is unwarranted to assume that every aspect of language is an adaptation, or that language as a whole is an adaptation. He criticizes some strands of evolutionary psychology for suggesting a pan-adaptionist view of evolution, and dismisses Pinker and Bloom's question of whether "Language has evolved as an adaptation" as being misleading. He argues instead that from a biological viewpoint the evolutionary origins of language is best conceptualized as being the probable result of a convergence of many separate adaptations into a complex system.[45] A similar argument is made by Terrence Deacon who in The Symbolic Species argues that the different features of language have co-evolved with the evolution of the mind and that the ability to use symbolic communication is integrated in all other cognitive processes.[46] If the theory that language could have evolved as a single adaptation is accepted, the question becomes which of its many functions has been the basis of adaptation, several evolutionary hypotheses have been posited: that it evolved for the purpose of social grooming, that it evolved to as a way to show mating potential or that it evolved to form social contracts. Evolutionary psychologists recognize that these theories are all speculative and that much more evidence is required to understand how language might have been selectively adapted.[47]

Mating Given that sexual reproduction is the means by which genes are propagated into future generations, sexual selection plays a large role in the direction of human evolution. Human mating, then, is of interest to evolutionary psychologists who aim to investigate evolved mechanisms to attract and secure mates.[48] Several lines of research have stemmed from this interest, such as studies of mate selection[49][50] mate poaching,[51] mate retention,[52] and mating preferences[53] Much of the research on human mating is based on parental investment theory,[54] which makes important predictions about the different strategies men and women will use in the mating domain (see above under "Middle-level evolutionary theories"). In essence, it predicts that women will be more selective when choosing mates, whereas men will not, especially under short-term mating conditions. While some other scientists such as Tim Brikhead assert that promiscuity can have benefits to women, such as fertility insurance, trading up to better genes, reducing risk of inbreeding, and insurance protection of her offspring.[55] Sarah Blaffer-Hrdy asserts that female chimpanzees, bonobos and langur monkeys, and probably human women have seemingly indiscriminate sexuality, and that being selective is biologically less advantageous than being promiscuous.[56] This has led some researchers to predict sex differences in such domains as sexual jealousy,[57][58] wherein females will react more aversively to emotional infidelity and males will react more aversively to sexual infidelity. This particular pattern is predicted

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Evolutionary psychology because the costs involved in mating for each sex are distinct. Women, on average, should prefer a mate who can offer some kind of resources (e.g., financial, commitment), which means that a woman would also be more at risk for losing those valued traits in a mate who commits an emotional infidelity. Men, on the other hand, are limited by the fact that they can never be certain of the paternity of their children because they do not bear the offspring themselves. This obstacle entails that sexual infidelity would be more aversive than emotional infidelity for a man because investing resources in another man's offspring does not lead to propagation of the man's own genes. Evidence for this pattern of sexual jealousy is mixed, with some research suggesting that men and women are both more averse to emotional infidelity than sexual infidelity. Another interesting line of research is that which examines women's mate preferences across the ovulatory cycle. The theoretical underpinning of this research is that ancestral women would have evolved mechanisms to select mates with certain traits depending on their hormonal status. For example, the theory hypothesizes that, during the ovulatory phase of a woman's cycle (approximately days 10–15 of a woman's cycle), a woman who mated with a male with high genetic quality would have been more likely, on average, to produce and rear a healthy offspring than a woman who mated with a male with low genetic quality. These putative preferences are predicted to be especially apparent for short-term mating domains because a potential male mate would only be offering genes to a potential offspring. This hypothesis allows researchers to examine whether women select mates who have characteristics that indicate high genetic quality during the high fertility phase of their ovulatory cycles. Indeed, studies have shown that women's preferences vary across the ovulatory cycle. In particular, Haselton and Miller (2006) showed that highly fertile women prefer creative but poor men as short-term mates. Creativity may be a proxy for good genes.[59] Research by Gangestad et al. (2004) indicates that highly fertile women prefer men who display social presence and intrasexual competition; these traits may act as cues that would help women predict which men may have, or would be able to acquire, resources.

Parenting Reproduction is always costly for women, and can also be for men. Individuals are limited in the degree to which they can devote time and resources to producing and raising their young, and such expenditure may also be detrimental to their future condition, survival and further reproductive output. Parental investment is any parental expenditure (time, energy etc.) that benefits one offspring at a cost to parents' ability to invest in other components of fitness (Clutton-Brock 1991: 9; Trivers 1972). Components of fitness (Beatty 1992) include the well being of existing offspring, parents' future reproduction, and inclusive fitness through aid to kin (Hamilton, 1964). Parental investment theory is a branch of life history theory. Robert Trivers' theory of parental investment predicts that the sex making the largest investment in lactation, nurturing and protecting offspring will be more discriminating in mating and that the sex that invests less in offspring will compete for access to the higher investing sex (see Bateman's principle). Sex differences in parental effort are important in determining the strength of sexual selection. The benefits of parental investment to the offspring are large and are associated with the effects on condition, growth, survival and ultimately, on reproductive success of the offspring. However, these benefits can come at the cost of parent's ability to reproduce in the future e.g. through the increased risk of injury when defending offspring against predators, the loss of mating opportunities whilst rearing offspring and an increase in the time to the next reproduction. Overall, parents are selected to maximize the difference between the benefits and the costs, and parental care will be likely to evolve when the benefits exceed the costs. The Cinderella effect is an alleged high incidence of stepchildren being physically, emotionally or sexually abused, neglected, murdered, or otherwise mistreated at the hands of their stepparents at significantly higher rates than their genetic counterparts. It takes its name from the fairy tale character Cinderella, who in the story was cruelly mistreated by her stepmother and stepsisters.[60] Daly and Wilson (1996) noted: "Evolutionary thinking led to the discovery of the most important risk factor for child homicide – the presence of a stepparent. Parental efforts and

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Evolutionary psychology investments are valuable resources, and selection favors those parental psyches that allocate effort effectively to promote fitness. The adaptive problems that challenge parental decision making include both the accurate identification of one's offspring and the allocation of one's resources among them with sensitivity to their needs and abilities to convert parental investment into fitness increments…. Stepchildren were seldom or never so valuable to one's expected fitness as one's own offspring would be, and those parental psyches that were easily parasitized by just any appealing youngster must always have incurred a selective disadvantage"(Daly & Wilson, 1996, p64 – 65). However, they note that not all stepparents will "want" to abuse their partner's children, or that genetic parenthood is any insurance against abuse. They see step parental care as primarily "mating effort" towards the genetic parent.[61]

Family and kin Inclusive fitness is the sum of an organism's classical fitness (how many of its own offspring it produces and supports) and the number of equivalents of its own offspring it can add to the population by supporting others.[62] The first component is called classical fitness by Hamilton (1964). From the gene's point of view, evolutionary success ultimately depends on leaving behind the maximum number of copies of itself in the population. Until 1964, it was generally believed that genes only achieved this by causing the individual to leave the maximum number of viable offspring. However, in 1964 W. D. Hamilton proved mathematically that, because close relatives of an organism share some identical genes, a gene can also increase its evolutionary success by promoting the reproduction and survival of these related or otherwise similar individuals. Hamilton claimed that this leads natural selection to favor organisms that would behave in ways that maximize their inclusive fitness. It is also true that natural selection favors behavior that maximizes personal fitness. Hamilton's rule describes mathematically whether or not a gene for altruistic behavior will spread in a population:

where • • •

is the reproductive cost to the altruist, is the reproductive benefit to the recipient of the altruistic behavior, and is the probability, above the population average, of the individuals sharing an altruistic gene – commonly viewed as "degree of relatedness".

The concept serves to explain how natural selection can perpetuate altruism. If there is an '"altruism gene"' (or complex of genes) that influences an organism's behavior to be helpful and protective of relatives and their offspring, this behavior also increases the proportion of the altruism gene in the population, because relatives are likely to share genes with the altruist due to common descent. Altruists may also have some way to recognize altruistic behavior in unrelated individuals and be inclined to support them. As Dawkins points out in The Selfish Gene (Chapter 6) and The Extended Phenotype,[63] this must be distinguished from the green-beard effect. Although it is generally true that humans tend to be more altruistic toward their kin than toward non-kin, the relevant proximate mechanisms that mediate this cooperation have been debated (see kin recognition), with some arguing that kin status is determined primarily via social and cultural factors (such as co-residence, maternal association of sibs, etc.), while others have argued that kin recognition can also mediated by biological factors such as facial resemblance and immunogenetic similarity of the major histocompatibility complex (MHC).[64] For a discussion of the interaction of these social and biological kin recognition factors see Lieberman, Tooby, and Cosmides (2007)[65] (PDF [66]). Whatever the proximate mechanisms of kin recognition there is substantial evidence that humans act generally more altruistically to close genetic kin compared to genetic non-kin.[67][68] )[69]

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Interactions with non-kin / reciprocity Although interactions with non-kin are generally less altruistic compared to those with kin, cooperation can be maintained with non-kin via mutually beneficial reciprocity as was proposed by Robert Trivers. If there are repeated encounters between the same two players in an evolutionary game in which each of them can choose either to "cooperate" or "defect," then a strategy of mutual cooperation may be favored even if it pays each player, in the short term, to defect when the other cooperates. Direct reciprocity can lead to the evolution of cooperation only if the probability, w, of another encounter between the same two individuals exceeds the cost-to-benefit ratio of the altruistic act: w > c/b Reciprocity can also be indirect if information about previous interactions is shared. Reputation allows evolution of cooperation by indirect reciprocity. Natural selection favors strategies that base the decision to help on the reputation of the recipient: studies show that people who are more helpful are more likely to receive help. The calculations of indirect reciprocity are complicated and only a tiny fraction of this universe has been uncovered, but again a simple rule has emerged. Indirect reciprocity can only promote cooperation if the probability, q, of knowing someone’s reputation exceeds the cost-to-benefit ratio of the altruistic act: q > c/b One important problem with this explanation is that individuals may be able to evolve the capacity to obscure their reputation, reducing the probability, q, that it will be known. Trivers argues that friendship and various social emotions evolved in order to manage reciprocity.[70] Liking and disliking, he says, evolved to help present day humans' ancestors form coalitions with others who reciprocated and to exclude those who did not reciprocate. Moral indignation may have evolved to prevent one's altruism from being exploited by cheaters, and gratitude may have motivated present day humans' ancestors to reciprocate appropriately after benefiting from others' altruism. Likewise, present day humans feel guilty when they fail to reciprocate. These social motivations match what evolutionary psychologists expect to see in adaptations that evolved to maximize the benefits and minimize the drawbacks of reciprocity. Evolutionary psychologists say that humans have psychological adaptations that evolved specifically to help us identify nonreciprocators, commonly referred to as "cheaters." In 1993, Robert Frank and his associates found that participants in a prisoner's dilemma scenario were often able to predict whether their partners would "cheat," based on a half hour of unstructured social interaction. In a 1996 experiment, for example, Linda Mealey and her colleagues found that people were better at remembering the faces of people when those faces were associated with stories about those individuals cheating (such as embezzling money from a church).

Evolution and culture Evolutionary psychology incorporates insights derived from other disciplines about how cultural phenomena evolve over time. Theories that have applied evolutionary perspectives to cultural phenomena include memetics, cultural ecology, and dual inheritance theory (gene-culture co-evolution).[71] Memetics is a theory of mental content based on an analogy with evolution, originating from Richard Dawkins' 1976 book The Selfish Gene. It purports to be an approach to evolutionary models of cultural information transfer. A meme, analogous to a gene, is essentially a "unit of culture"—an idea, belief, pattern of behavior, etc. which is "hosted" in one or more individual minds, and which can reproduce itself from mind to mind. Thus what would otherwise be regarded as one individual influencing another to adopt a belief is seen memetically as a meme reproducing itself. As with genetics, particularly under Dawkins's interpretation, a meme's success may be due to its contribution to the effectiveness of its host. Memetics is notable for sidestepping the traditional concern with the truth of ideas and beliefs.

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Evolutionary psychology Susan Blackmore (2002) re-stated the definition of meme as: whatever is copied from one person to another person, whether habits, skills, songs, stories, or any other kind of information. Further she said that memes, like genes, are replicators in the sense as defined by Dawkins.[72] That is, they are information that is copied. Memes are copied by imitation, teaching and other methods. The copies are not perfect: memes are copied with variation; moreover, memes compete for humans' limited memory capacity and for the chance to be copied again. Only some of the variants can survive. The combination of these three elements (copies; variation; competition for survival) forms precisely the condition for Darwinian evolution, and so memes (and hence human cultures) evolve. Large groups of memes that are copied and passed on together are called co-adapted meme complexes, or memeplexes. In her definition, the way that a meme replicates is through imitation. Dual inheritance theory (DIT), also known as gene-culture coevolution, suggests that cultural information and genes co-evolve. Marcus Feldman and Luigi Luca Cavalli-Sforza (1976) published perhaps the first dynamic models of gene-culture coevolution. These models were to form the basis for subsequent work on DIT, heralded by the publication of three seminal books in 1980 and 1981. Charles Lumsden and E.O. Wilson's Genes, Mind and Culture (1981).[73] also outlined a series of mathematical models of how genetic evolution might favor the selection of cultural traits and how cultural traits might, in turn, affect the speed of genetic evolution. Another 1981 book relevant to this topic was Cavalli-Sforza and Feldman's Cultural Transmission and Evolution: A Quantitative Approach.[74] Borrowing heavily from population genetics and epidemiology, this book built a mathematical theory concerning the spread of cultural traits. It describes the evolutionary implications of vertical transmission, passing cultural traits from parents to offspring; oblique transmission, passing cultural traits from any member of an older generation to a younger generation; and horizontal transmission, passing traits between members of the same population. Robert Boyd and Peter Richerson's (1985) Culture and the Evolutionary Process presents models of the evolution of social learning under different environmental conditions, the population effects of social learning, various forces of selection on cultural learning rules, different forms of biased transmission and their population-level effects, and conflicts between cultural and genetic evolution. Along with game theory, Herbert Gintis suggested that Dual inheritance theory has potential for unifying the behavioral sciences, including economics, biology, anthropology, sociology, psychology and political science because it addresses both the genetic and cultural components of human inheritance. Laland and Brown hold a similar view.[citation needed]

In psychology sub-fields Developmental psychology According to Paul Baltes, the benefits granted by evolutionary selection decrease with age. Natural selection has not eliminated many harmful conditions and nonadaptive characteristics that appear among older adults, such as Alzheimer disease. If it were a disease that killed 20 year-olds instead of 70 year-olds this may have been a disease that natural selection could have eliminated ages ago. Thus, unaided by evolutionary pressures against nonadaptive conditions, modern humans suffer the aches, pains, and infirmities of aging and as the benefits of evolutionary selection decrease with age, the need for culture increases.[75]

Social psychology As humans are a highly social species, there are many adaptive problems associated with navigating the social world (e.g., maintaining allies, managing hierarchies, interacting with outgroup members). Researchers in the emerging field of evolutionary social psychology have made many discoveries pertaining to topics traditionally studied by social psychologists, including person perception, social cognition, attitudes, altruism, emotions, group dynamics, leadership, motivation, prejudice, intergroup relations, and cross-cultural differences.[76][77]

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Abnormal psychology Adaptationist hypotheses regarding the etiology of psychological disorders are often based on analogies between physiological and psychological dysfunctions,[78] as noted in the table below. Prominent theorists and evolutionary psychiatrists include Michael T. McGuire and Randolph M. Nesse. They, and others, suggest that mental disorders are due to the interactive effects of both nature and nurture, and often have multiple contributing causes. Possible Causes of Psychological 'Abnormalities' from an Adaptationist Perspective Summary based on information in these textbooks (all titled "Evolutionary Psychology"): Buss (2011), Gaulin & McBurney (2004), [79] Workman & Reader (2008) as well as Cosmides & Tooby (1999) Toward an evolutionary taxonomy of treatable conditions Causal mechanism of failure or malfunction of adaptation

Physiological Example

Hypothesized Psychological Example

Functioning adaptation (adaptive defense)

Fever / Vomiting

Mild depression or anxiety

(functional responses to infection or ingestion of toxins)

(functional responses to mild loss or stress / reduction of social [80] interactions to prevent infection by contagious pathogens)

By-product of an adaptation(s)

Intestinal gas

Sexual fetishes (?)

(byproduct of digestion of fiber)

(possible byproduct of normal sexual arousal adaptations that have 'imprinted' on unusual objects or situations)

Gene for malaria resistance, in homozygous form, causes sickle cell anemia

Adaptation(s) for high levels of creativity may also predispose schizophrenia or bi-polar disorder

Allergies

Autism

(over-reactive immunological responses)

(possible malfunctioning of theory of mind module)

The two sexes / Different blood and immune system types

Personality traits and personality disorders

Mismatch between ancestral & current environments

Modern diet-related Type 2 Diabetes

More frequent modern interaction with strangers (compared to family and close friends) may predispose greater incidence of depression & anxiety

Tails of normal (bell shaped) curve

Very short or tall height

Tails of the distribution of personality traits (e.g., extremely introverted or extroverted)

Adaptations with multiple effects

Malfunctioning adaptation

Frequency-dependent morphs

(adaptations with both positive and negative effects, perhaps dependent on alternate developmental trajectories)

(may represent alternative behavioral strategies dependent on the frequency of the strategy in the population)

Evolutionary psychologists have suggested that schizophrenia and bipolar disorder may reflect a side-effect of genes with fitness benefits, such as increased creativity.[81] (Some individuals with bipolar disorder are especially creative during their manic phases and the close relatives of schizophrenics have been found to be more likely to have creative professions.) A 1994 report by the American Psychiatry Association found that people suffered from schizophrenia at roughly the same rate in Western and non-Western cultures, and in industrialized and pastoral societies, suggesting that schizophrenia is not a disease of civilization nor an arbitrary social invention. Sociopathy may represent an evolutionarily stable strategy, by which a small number of people who cheat on social contracts benefit in a society consisting mostly of non-sociopaths. Mild depression may be an adaptive response to withdraw from, and re-evaluate, situations that have led to disadvantageous outcomes (see Evolutionary approaches to depression). Some of these speculations have yet to be developed into fully testable hypotheses, and a great deal of research is required to confirm their validity.[82] Clinical psychology and psychiatry are relatively isolated from the field of evolutionary psychology and the etiological speculations of evolutionary psychology have yet to pass the scrutiny and demanding research criteria of these larger disciplines. Some psychiatrists raise the concern that evolutionary psychologists seek to explain hidden adaptive advantages without engaging the rigorous empirical testing required to

Evolutionary psychology back up such claims. While there is strong research to suggest a genetic link to bipolar disorder and schizophrenia, there is significant debate within clinical psychology about the relative influence and the mediating role of cultural or environmental factors.[83] For example, epidemiological research suggests that different cultural groups may have divergent rates of diagnosis, symptomatology, and expression of mental illnesses.[84] There has also been increasing acknowledgment of culture-bound disorders,[85] which may be viewed as an argument for an environmental versus genetic psychological adaptation.[86] While certain mental disorders may have psychological traits that can be explained as 'adaptive' on an evolutionary scale, these disorders cause afflicted individuals significant emotional and psychological distress and negatively influence the stability of interpersonal relationships and day-to-day adaptive functioning.[87]

Psychology of religion Adaptationist perspectives on religious belief suggest that, like all behavior, religious behaviors are a product of the human brain. As with all other organ functions, cognition's functional structure has been argued to have a genetic foundation, and is therefore subject to the effects of natural selection and sexual selection. Like other organs and tissues, this functional structure should be universally shared amongst humans and should have solved important problems of survival and reproduction in ancestral environments. However, evolutionary psychologists remain divided on whether religious belief is more likely a consequence of evolved psychological adaptations, or is a byproduct of other cognitive adaptations.

Reception Critics of evolutionary psychology accuse it of promoting genetic determinism, panadaptionism (the idea that all behaviors and anatomical features are adaptations), unfalsifiable hypotheses, distal or ultimate explanations of behavior when proximate explanations are superior, and malevolent political or moral ideas.[88]

Ethical implications Critics have argued that evolutionary psychology might be used to justify existing social hierarchies and reactionary policies. It has also been suggested by critics that evolutionary psychologists' theories and interpretations of empirical data rely heavily on ideological assumptions about race and gender. Other critics contend that some theories – for example, the authors of A Natural History of Rape view rape as a form of mate choice that enhances male fitness – could have potentially far-reaching ethical implications. In response to such criticism, evolutionary psychologists often invoke the naturalistic fallacy – the idea that "what is natural" is necessarily a moral good.[89]Wikipedia:Citing sources However, their use of the naturalistic fallacy has been criticized as fallacious and a means to stifle legitimate ethical discussions.

Standard social science model Evolutionary psychology has been entangled in the larger philosophical and social science controversies related to the nature versus nurture debate. Evolutionary psychologists typically contrast evolutionary psychology with what they call the standard social science model (SSSM). They characterize the SSSM as the "blank slate," social constructionist, or "cultural determinist" perspective that they claim dominated the social sciences throughout the 20th century and assumed that the mind was shaped almost entirely by culture. Critics have argued that evolutionary psychologists created a false dichotomy between their own view and the caricature of the SSSM. Other critics regard the SSSM as a rhetorical device or a straw man and suggest that the scientists whom evolutionary psychologists associate with the SSSM did not believe that the mind was a blank state devoid of any natural predispositions.

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Reductionism and determinism Some critics view evolutionary psychology as a form of genetic reductionism and genetic determinism,[90] a common critique being that evolutionary psychology does not address the complexity of individual development and experience and fails to explain the influence of genes on behavior in individual cases.[91] Evolutionary psychologists respond that EP works within a nature-nurture interactionist framework that acknowledges that many psychological adaptations are facultative (sensitive to environmental variations during individual development). EP is generally not focused on proximate analyses of behavior but rather its focus is on the study of distal/ultimate causality (the evolution of psychological adaptations). The field of behavioral genetics is focused on the study of the proximate influence of genes on behavior.

Testability of hypotheses A frequent critique of the discipline is that the hypotheses of evolutionary psychology are frequently arbitrary and difficult or impossible to adequately test, thus questioning its status as an actual scientific discipline, for example because many current traits probably evolved to serve different functions than they do now. While evolutionary psychology hypotheses are difficult to test, evolutionary psychologists assert that it is not impossible.[92] Part of the critique of the scientific base of evolutionary psychology includes a critique of the concept of the Environments of Evolutionary Adaptation (EEA). Some critics have argued that researchers know so little about the environment in which Homo sapiens evolved that explaining specific traits as an adaption to that environment becomes highly speculative.[93] Evolutionary psychologists respond that they do know many things about this environment, including the facts that only women became pregnant, present day humans' ancestors were hunter-gatherers that generally lived in small tribes, etc.[94]

Modularity of mind Evolutionary psychologists generally presume that, like the body, the mind is made up of many evolved modular adaptations, although there is some disagreement within the discipline regarding the degree of general plasticity, or "generality," of some modules. It has been suggested that modularity evolves because, compared to non-modular networks, it would have conferred an advantage in terms of fitness and because connection costs are lower. In contrast, some academics argue that it is unnecessary to posit the existence of highly domain specific modules, and, suggest that the neural anatomy of the brain supports a model based on more domain general faculties and processes. Moreover, empirical support for the domain-specific theory stems almost entirely from performance on variations of the Wason selection task which is extremely limited in scope as it only tests one subtype of deductive reasoning.

Evolutionary psychology defense Evolutionary psychologists have addressed many of their critics (see, for example, books by Segerstråle (2000), Defenders of the Truth: The Battle for Science in the Sociobiology Debate and Beyond,[95] Barkow (2005), Missing the Revolution: Darwinism for Social Scientists,[96] and Alcock (2001), The Triumph of Sociobiology.[97]). Among their rebuttals are that some criticisms are straw men, are based on an incorrect nature vs. nurture dichotomy, are based on misunderstandings of the discipline, etc.[98][99][100][101][102][103][104] Robert Kurzban suggested that " ...critics of the field, when they err, are not slightly missing the mark. Their confusion is deep and profound. It’s not like they are marksman who can’t quite hit the center of the target; they’re holding the gun backwards."[105]

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Notes [1] Confer et al. 2010; Buss, 2005; Durrant & Ellis, 2003; Pinker, 2002; Tooby & Cosmides, 2005 [2] Duntley and Buss 2008 [3] Carmen, R.A., et al. (2013). Evolution Integrated Across All Islands of the Human Behavioral Archipelago: All Psychology as Evolutionary Psychology. EvoS Journal: The Journal of the Evolutionary Studies Consortium, 5, pp. 108-126. ISSN: 1944-1932 PDF (http:/ / evostudies. org/ wp-content/ uploads/ 2013/ 03/ Carmen_-Vol5Iss1. pdf) [4] "Despite this difficulty, there have been many careful and informative studies of human social behavior from an evolutionary perspective. Infanticide, intelligence, marriage patterns, promiscuity, perception of beauty, bride price, altruism, and the allocation of parental care have all been explored by testing predictions derived from the idea that conscious and unconscious behaviours have evolved to maximize inclusive fitness. The findings have been impressive." "social behaviour, animal." Encyclopædia Britannica. Encyclopædia Britannica Online. Encyclopædia Britannica, 2011. Web. 23 Jan 2011. (http:/ / www. britannica. com/ EBchecked/ topic/ 550897/ animal-social-behaviour). [5] The Oxford Handbook of Evolutionary Psychology, Edited by Robin Dunbar and Louise Barret, Oxford University Press, 2007 [6] The Handbook of Evolutionary Psychology, edited by David M. Buss, John Wiley & Sons, Inc., 2005 [7] Tooby & Cosmides 2005, p. 5 [8] Nesse, R.M. (2000). Tingergen's Four Questions Organized. Read online (http:/ / www-personal. umich. edu/ ~nesse/ Nesse-Tinbergen4Q. PDF). [9] Gaulin and McBurney 2003 p 1-24. [10] Evolutionary Psychology at the University of Texas (http:/ / homepage. psy. utexas. edu/ homepage/ Group/ BussLAB/ about. htm) [11] Sterelny, Kim. 2009. In Ruse, Michael & Travis, Joseph (eds) Wilson, Edward O. (Foreword) Evolution: The First Four Billion Years. Harvard University Press, Cambridge, Ma. ISBN 978-0-674-03175-3. p. 314. [12] Wilson, Edward O. 1975. Sociobiology: The New Synthesis. (http:/ / www. hup. harvard. edu/ catalog/ WILSOR. html) Harvard University Press, Cambridge, Ma. ISBN 0-674-00089-7 p.4. [13] Wilson, Edward O. 1978. On Human Nature. Harvard University Press, Cambridge, Ma. p. x. [14] Laland, Kevin N. and Gillian R. Brown. 2002. Sense & Nonsense: Evolutionary Perspectives on Human Behavior. Oxford University Press, Oxford. pp. 287–319. [15] Gaulin and McBurney 2003 p 25-56. [16] See also "Environment of evolutionary adaptation," a variation of the term used in Economics, e.g., in Rubin, Paul H., 2003, "Folk economics" Southern Economic Journal, 70:1, July 2003, 157–171. [17] Buss et al. 1998 [18] Pinker, Steven. (1994)The Language Instinct [19] George C Williams, Adaptation and Natural Selection. p.4. [20] Buss, D. M. (2011). Evolutionary psychology. [21] Brown, Donald E. (1991) Human Universals. New York: McGraw-Hill. [22] Symons, D. (1979). The evolution of human sexuality. Oxford: Oxford University Press. Chapter 6. [23] Pinker 2002 [24] Barkow et al. 1992 [25] "instinct." Encyclopædia Britannica. Encyclopædia Britannica Online. Encyclopædia Britannica, 2011. Web. 18 Feb 2011. (http:/ / www. britannica. com/ EBchecked/ topic/ 289249/ instinct). [26] Ryan, Christopher and Cacilda Jethá. Sex at Dawn: The Prehistoric Origins of Modern Sexuality. Harper. 2010. [27] CDC pdf (http:/ / www. cdc. gov/ nchs/ data/ nvsr/ nvsr54/ nvsr54_10. pdf) [28] Barrett, Deirdre. Waistland: The R/Evolutionary Science Behind Our Weight and Fitness Crisis (2007). New York: W.W. Norton. p. 31-51. [29] Barrett, Deirdre. Supernormal Stimuli: How Primal Urges Overran Their Evolutionary Purpose. New York: W.W. Norton, 2010 [30] Buss, D.M. (2011). Evolutionary psychology. Chapter 2. Boston: Pearson/A and B. [31] Buss, D.M. (2011). Evolutionary Psychology: The New Science of the Mind [32] Freeman and Herron. Evolutionary Analysis. 2007. Pearson Education, NJ. [33] Peters, Frederic "Consciousness as Recursive, Spatiotemporal Self-Location" (http:/ / precedings. nature. com/ documents/ 2444/ version/ 1) [34] Baars, Bernard J. A Cognitive Theory of Consciousness. 1993. Cambridge University Press. [35] Gaulin and McBurney 2003 Chapter 8. [36] Gaulin and McBurney 2003 p 121-142. [37] Gaulin and McBurney 2003 Chapter 7. [38] Gaulin and McBurney 2003 Chapter 9. [39] Sulloway, F. (1996). Born to rebel. NY: Pantheon. [40] Workman, Lance and Will Reader (2004) Evolutionary psychology: an introduction. Cambridge University Press p. 259 [41] Workman, Lance and Will Reader (2008). Evolutionary psychology: an introduction. 2nd Ed. Cambridge University Press. Chapter 10. [42] Diller, K. C. and R. L. Cann 2009. Evidence against a genetic-based revolution in language 50,000 years ago. In R. Botha and C. Knight (eds), The Cradle of Language. Oxford: Oxford University Press, pp. 135-149. [43] Workman & Reader 2008:277 "There are a number of hypotheses suggesting that language evolved to fulfil a social function such as social grooming (to bind large groups together), the making of social contracts (to enable monogamy and male provisioning) and the use of language

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Evolutionary psychology to impress potential mates. While each of these hypotheses has its merits, each is still highly speculative and requires more evidence from different areas of research (such as linguistics and anthropology)." [44] Workman, Lance and Will Reader (2004) Evolutionary psychology: an introduction. Cambridge University Press p. 267 [45] W. Tecumseh Fitch (2010) The Evolution of Language. Cambridge University Press p.65-66 [46] Deacon, Terrence W. (1997) The Symbolic Species: The Co-evolution of Language and the Brain. W.W. Norton & Co [47] Workman, Lance and Will Reader (2004) Evolutionary psychology: an introduction. Cambridge University Press p. 277 [48] Wilson, G.D. Love and Instinct. London: Temple Smith, 1981. [49] Buss 1994 [50] Buss & Barnes 1986 [51] Schmitt and Buss 2001 [52] Buss 1988. [53] Shackelford, Schmitt, & Buss (2005) Universal dimensions of human mate preferences; Personality and Individual Differences 39 [54] Trivers, R. (1972). Parental investment and sexual selection. In B. Campbell (Ed.), Sexual Selection and the Descent of Man. Chicago: Aldine-Atherton. [55] http:/ / www. guardian. co. uk/ uk/ 2000/ sep/ 03/ anthonybrowne. theobserver [56] www.citrona.com/hrdy/documents/findingmrright.pdf [57] Buss 1989 [58] Buss et al. 1992 [59] Miller, G. F. (2000b) The mating mind: How sexual choice shaped the evolution of human nature. Anchor Books: New York. [60] Daly, Matin, and Margo I. Wilson. (1999) [61] Daly & Wilson 1998 [62] Definition and explanation of inclusive fitness from Personality Research.org (http:/ / www. personalityresearch. org/ evolutionary/ inclusive. html) [63] Dawkins, Richard, "The Extended Phenotype", Oxford University Press 1982 (Chapter 9) [64] Villinger, J., and Waldman, B. (2012). Social discrimination by quantitative assessment of immunogenetic similarity. Published online before print September 5, 2012, doi: 10.1098/rspb.2012.1279 Proc. R. Soc. B [65] Lieberman, D., Tooby, J. & Cosmides, L. The architecture of human kin detection Nature 445, 727-731 (15 February 2007) | doi:10.1038/nature05510 [66] http:/ / www. psych. ucsb. edu/ research/ cep/ papers/ kinnature05510. pdf [67] Buss, D.M. (2011). Evolutionary Psychology. Monterey: Brooks-Cole. [68] Gaulin & McBurney (2004), Evolutionary Psychology, 2nd Ed. NY: Prentice Hall [69] Workman & Reader (2008), Evolutionary Psychology, 2nd Ed. Cambridge: Cambridge University Press [70] Gaulin, Steven J. C. and Donald H. McBurney. Evolutionary Psychology. Prentice Hall. 2003. ISBN 978-0-13-111529-3, Chapter 14, p 323-352. [71] Workman, Lance and Will Reader (2004) Evolutionary psychology: an introduction. Cambridge University Press, Chapter 13. [72] Dawkins, R. (1982) "Replicators and Vehicles" (http:/ / www. stephenjaygould. org/ library/ dawkins_replicators. html) King's College Sociobiology Group, eds., Current Problems in Sociobiology, Cambridge, Cambridge University Press, pp. 45–64. "A replicator may be defined as any entity in the universe of which copies are made." [73] Lumsden C., and E. Wilson. 1981. Genes, Mind and Culture: The Coevolutionary Process. Cambridge, MA: Harvard University Press. [74] Cavalli-Sfornza, L. and M. Feldman. 1981. Cultural Transmission and Evolution: A Quantitative Approach. Princeton, New Jersey: Princeton University Press. [75] Santrock, W. John (2005). A Topical Approach to Life-Span Development (3rd ed.). New York, NY: McGraw-Hill. pp.62. [76] Neuberg, S. L., Kenrick, D. T., & Schaller, M. (2010). Evolutionary social psychology. In S. T. Fiske, D. T. Gilbert, & G. Lindzey (Eds.), Handbook of social psychology (5th Edition, Vol. 2, pp. 761–796). New York: John Wiley & Sons. [77] Schaller, M., Simpson, J. A., & Kenrick, D. T. (Eds.) (2006). Evolution and social psychology. New York: Psychology Press. [78] (adaptationist perspective to both physiological and psychological dysfunctions) [79] Workman & Reader (2008), Evolutionary Psychology, 2nd Ed. Cambridge: Cambridge University Press, [80] Raison, C.L, Miller, A. N. (2012). The evolutionary significance of depression in Pathogen Host Defense (PATHOS-D) Molecular Psychiatry 1-23. PDF (http:/ / www. nature. com/ mp/ journal/ vaop/ ncurrent/ pdf/ mp20122a. pdf). [81] Gaulin and McBurney 2003 p 239-256. [82] O’Connell, H. (2004) Evolutionary theory in psychiatry and psychology. Irish Journal of Psychological Medicine, 21 (1), pp. 37–37. [83] Adams, H. and Sutker, P. Comprehensive Handbook of Psychopathology, 3rd Ed. Springer. 2001. Chapter 3, p 53-84. [84] Adams, H. and Sutker, P. Comprehensive Handbook of Psychopathology, 3rd Ed. Springer. 2001. Chapter 5, p 105-127 [85] American Psychiatric Association. Diagnostic and statistical manual of mental disorders (4th ed., text revision). 2000. Culture Bound Syndromes, p 897-903. [86] Schumaker, J. The Age of Insanity: Modernity and Mental Health. Praeger. 2001. Chapter 3, p 29-49. [87] Adams, H. and Sutker, P. Comprehensive Handbook of Psychopathology, 3rd Ed. Springer. 2001 [88] Kurzban, Robert. Alas poor evolutionary psychology (http:/ / human-nature. com/ nibbs/ 02/ apd. html). The Human Nature Review 2002 Volume 2: 99-109 (14 March ). Retrieved 14 July 2013.

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Evolutionary psychology [89] Pinker, S. (2003). The blank slate. NY: Penguin [90] Plotkin, Henry. 2004 Evolutionary thought in Psychology: A Brief History. Blackwell. p.150. [91] "instinct." Encyclopædia Britannica. Encyclopædia Britannica Online. Encyclopædia Britannica, 2011. Web. 9 Feb 2011. (http:/ / www. britannica. com/ EBchecked/ topic/ 289249/ instinct). [92] "Testing ideas about the evolutionary origins of psychological phenomena is indeed a challenging task, but not an impossible one" Buss et al. 1998; Pinker, 1997b). [93] Plotkin, Henry. 2004 Evolutionary thought in Psychology: A Brief History. Blackwell. p.149. [94] The Handbook of Evolutionary Psychology (2005), David M. Buss, Chapter 1, pp. 5–67, Conceptual Foundations of Evolutionary Psychology, John Tooby and Leda Cosmides [95] Segerstråle, Ullica Christina Olofsdotter (2000). Defenders of the truth: The battle for science in the sociobiology debate and beyond. Oxford [Oxfordshire]: Oxford University Press. ISBN 0-19-850505-1. [96] Jerome H. Barkow, (2005), Missing the Revolution: Darwinism for Social Scientists Oxford, Oxford University Press. [97] Alcock, John (2001). The Triumph of Sociobiology. Oxford: Oxford University Press. ISBN 978-0-19-516335-3 [98] Segerstråle, Ullica Christina Olofsdotter (2000). Defenders of the truth : the battle for science in the sociobiology debate and beyond. Oxford [Oxfordshire]: Oxford University Press. ISBN 0-19-850505-1. [99] Barkow, Jerome (Ed.). (2006) Missing the Revolution: Darwinism for Social Scientists. Oxford: Oxford University Press. ISBN 978-0-19-513002-7 [100] Tooby, J., Cosmides, L. & Barrett, H. C. (2005). Resolving the debate on innate ideas: Learnability constraints and the evolved interpenetration of motivational and conceptual functions. In Carruthers, P., Laurence, S. & Stich, S. (Eds.), The Innate Mind: Structure and Content. NY: Oxford University Press. [101] Controversies surrounding evolutionary psychology by Edward H. Hagen, Institute for Theoretical Biology, Berlin. In D. M. Buss (Ed.), The Handbook of Evolutionary Psychology (pp. 5–67). Hoboken, NJ: Wiley. [102] The Never-Ending Misconceptions About Evolutionary Psychology: Persistent Falsehoods About Evolutionary Psychology (http:/ / www. psychologytoday. com/ blog/ homo-consumericus/ 200906/ the-never-ending-misconceptions-about-evolutionary-psychology) by Gad Saad, Psychology Today blog. [103] Geher, G. (2006). Evolutionary psychology is not evil! … and here’s why … Psihologijske Teme (Psychological Topics); Special Issue on Evolutionary Psychology, 15, 181-202. (http:/ / www2. newpaltz. edu/ ~geherg/ ep_not_evil. pdf) [104] What Anti-Evolutionary Psychologists are Really Worried About (http:/ / www. psychologytoday. com/ blog/ cui-bono/ 201110/ what-anti-evolutionary-psychologists-are-really-worried-about) by John Johnson, Psychology Today blog. [105] Kurzban, R. (2013). This One Goes to Eleven, PZ Myers, and Other Punch Lines. (http:/ / www. epjournal. net/ blog/ 2013/ 07/ this-one-goes-to-eleven-pz-myers-and-other-punch-lines/ ) Evolutionary Psychology.

References • Barkow, Jerome H. (2006). Missing the Revolution: Darwinism for Social Scientists. Oxford University Press, USA. ISBN 0-19-513002-2. • Barkow, J., Cosmides, L. & Tooby, J. 1992. The adapted mind: Evolutionary psychology and the generation of culture. Oxford: Oxford University Press. • Buss, D. M.; Barnes, M. (1986). "Preferences in human mate selection" (http://homepage.psy.utexas.edu/ homepage/Group/BussLAB/pdffiles/prefs_mate_selection_1986_jpsp.pdf) (PDF). Journal of Personality and Social Psychology 50 (3): 559–570. doi: 10.1037/0022-3514.50.3.559 (http://dx.doi.org/10.1037/0022-3514. 50.3.559). • Buss, D. M. (1988). "From vigilance to violence: Tactics of mate retention in American undergraduates" (http:// homepage.psy.utexas.edu/homepage/Group/BussLAB/pdffiles/Vigilance_to_Violence_1988.pdf) (PDF). Ethology and Sociobiology 9 (5): 291–317. doi: 10.1016/0162-3095(88)90010-6 (http://dx.doi.org/10.1016/ 0162-3095(88)90010-6). • Buss, D. M. (1989). "Sex differences in human mate preferences: Evolutionary hypotheses tested in 37 cultures" (http://homepage.psy.utexas.edu/homepage/Group/BussLAB/pdffiles/SexDifferencesinHuman.PDF). Behavioral and Brain Sciences 12: 1–49. doi: 10.1017/S0140525X00023992 (http://dx.doi.org/10.1017/ S0140525X00023992). • Buss, D. M.; Larsen, R. J.; Westen, D.; Semmelroth, J. (1992). "Sex differences in jealousy: Evolution, physiology, and psychology" (http://homepage.psy.utexas.edu/homepage/Group/BussLAB/pdffiles/ SexDifferencesinJealousy.PDF). Psychological Science 3 (4): 251–255. doi: 10.1111/j.1467-9280.1992.tb00038.x (http://dx.doi.org/10.1111/j.1467-9280.1992.tb00038.x).

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Evolutionary psychology • Buss, D. M. (1994). The evolution of desire: Strategies of human mating. New York: Basic Books. • Buss, David M. (2004). Evolutionary psychology: the new science of the mind. Boston: Pearson/A and B. ISBN 0-205-37071-3. • Buss, David M.; Haselton, Martie G.; Shackelford, Todd K.; Bleske, April L.; Wakefield, Jerome C. (1998). "Adaptations, Exaptations, and Spandrels" (http://www.sscnet.ucla.edu/comm/haselton/webdocs/spandrels. html). American Psychologist 53 (5): 533–548. doi: 10.1037/0003-066X.53.5.533 (http://dx.doi.org/10.1037/ 0003-066X.53.5.533). PMID  9612136 (http://www.ncbi.nlm.nih.gov/pubmed/9612136). Retrieved 29 August 2011. • Clarke, Murray (2004). Reconstructing reason and representation. Cambridge, Mass: MIT Press. ISBN 0-262-03322-4. • Confer, Easton, Fleischman, Goetz, Lewis, Perilloux & Buss Evolutionary Psychology (http://homepage.psy. utexas.edu/homepage/Group/BussLAB/pdffiles/evolutionary_psychology_AP_2010.pdf), American Psychologist, 2010. • Duntley, J.D.; Buss, D.M. (2008). "Evolutionary psychology is a metatheory for psychology" (http://homepage. psy.utexas.edu/homepage/Group/BussLAB/pdffiles/duntleybuss2008.pdf) (PDF). Psychological Inquiry 19: 30–34. doi: 10.1080/10478400701774105 (http://dx.doi.org/10.1080/10478400701774105). • Durrant, R., & Ellis, B.J. (2003). Evolutionary Psychology. In M. Gallagher & R.J. Nelson (Eds.), Comprehensive Handbook of Psychology, Volume Three: Biological Psychology (pp. 1–33). New York: Wiley & Sons. • Evan, Dylan (2000). Introducing Evolutionary Psychology. Lanham, MD: Totem Books USA. ISBN 1-84046-043-1. • Fruehwald, Edwin Scott, Law and Human Behavior: A Study in Behavioral Biology, Neuroscience, and The Law (Vandeplas 2011). ISBN 978-1-60042-144-0 • Gaulin, Steven J. C. and Donald H. McBurney. Evolutionary psychology. Prentice Hall. 2003. ISBN 978-0-13-111529-3 • Joyce, Richard (2006). The Evolution of Morality (Life and Mind: Philosophical Issues in Biology and Psychology). Cambridge, Mass: The MIT Press. ISBN 0-262-10112-2. • Miller, Geoffrey P. (2000). The mating mind: how sexual choice shaped the evolution of human nature. Garden City, N.Y: Doubleday. ISBN 0-385-49516-1. • Nesse, R.M. (2000). Tingergen's Four Questions Organized (http://www-personal.umich.edu/~nesse/ Nesse-Tinbergen4Q.PDF). • Nesse, R; Williams, George C. (1996). Why We Get Sick. NY: Vintage. • Pinker, Steven (1997). How the mind works. New York: Norton. ISBN 0-393-04535-8. • Pinker, Steven (2002). The blank slate: the modern denial of human nature. New York, N.Y: Viking. ISBN 0-670-03151-8. • Richards, Janet C. (2000). Human nature after Darwin: a philosophical introduction. New York: Routledge. ISBN 0-415-21243-X. • Ryan, C. & Jethá, C. (2010). Sex at Dawn: The Prehistoric Origins of Modern Sexuality. New York, NY: Harper. ISBN 0-06-170780-5. • Santrock, John W. (2005). The Topical Approach to Life-Span Development(3rd ed.). New York, N.Y: McGraw Hill. ISBN 0-07-322626-2. • Schacter, Daniel L, Daniel Wegner and Daniel Gilbert. 2007. Psychology. Worth Publishers. ISBN 0-7167-5215-8 ISBN 9780716752158. • Schmitt, D. P.; Buss, D. M. (2001). "Human mate poaching: Tactics and temptations for infiltrating existing relationships" (http://homepage.psy.utexas.edu/homepage/Group/BussLAB/pdffiles/ Human_Mate_Poaching_2001.pdf) (PDF). Journal of Personality and Social Psychology 80 (6): 894–917. doi: 10.1037/0022-3514.80.6.894 (http://dx.doi.org/10.1037/0022-3514.80.6.894). PMID  11414373 (http:// www.ncbi.nlm.nih.gov/pubmed/11414373).

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Evolutionary psychology • Tooby, J. & Cosmides, L. (2005). Conceptual foundations of evolutionary psychology. In D. M. Buss (Ed.), The Handbook of Evolutionary Psychology (pp. 5–67). Hoboken, NJ: Wiley. Full text (http://www.psych.ucsb.edu/ research/cep/papers/bussconceptual05.pdf) • Wilson, Edward Raymond (2000). Sociobiology: the new synthesis. Cambridge: Belknap Press of Harvard University Press. ISBN 0-674-00089-7. • Wright, Robert C. M. (1995). The moral animal: evolutionary psychology and everyday life. New York: Vintage Books. ISBN 0-679-76399-6.

Further reading • Buss, D. M. (1995). "Evolutionary psychology: A new paradigm for psychological science" (http://homepage. psy.utexas.edu/homepage/Group/BussLAB/pdffiles/ANewParadigmforPsych.PDF). Psychological Inquiry 6: 1–30. doi: 10.1207/s15327965pli0601_1 (http://dx.doi.org/10.1207/s15327965pli0601_1). • Confer, J.C.; Easton, J.A.; Fleischman, D.S.; Goetz, C. D.; Lewis, D.M.G.; Perilloux, C.; Buss, D. M. (2010). "Evolutionary Psychology: Controversies, Questions, Prospects, and Limitations" (http://homepage.psy.utexas. edu/homepage/Group/BussLAB/pdffiles/evolutionary_psychology_AP_2010.pdf) (PDF). American Psychologist 65 (2): 110–126. doi: 10.1037/a0018413 (http://dx.doi.org/10.1037/a0018413). PMID  20141266 (http://www.ncbi.nlm.nih.gov/pubmed/20141266). • Heylighen F. (2012). " Evolutionary Psychology (http://pcp.vub.ac.be/Papers/EvolutionaryPsychology-QOL. pdf)", in: A. Michalos (ed.): Encyclopedia of Quality of Life Research (Springer, Berlin). • Kennair, L. E. O. (2002). "Evolutionary psychology: An emerging integrative perspective within the science and practice of psychology" (http://www.human-nature.com/nibbs/02/ep.html). Human Nature Review 2: 17–61. • Medicus, G. (2005). "Evolutionary Theory of Human Sciences" (http://homepage.uibk.ac.at/~c720126/ humanethologie/ws/medicus/block1/inhalt.html). pp. 9, 10, 11. Retrieved 2009-09-08.

External links • Evolutionary Psychology (http://www.dmoz.org/Science/Social_Sciences/Psychology/ Evolutionary_Psychology/) at the Open Directory Project • What Is Evolutionary Psychology? by Clinical Evolutionary Psychologist Dale Glaebach (http://www. systemsthinker.com/interests/mind/glabachep/glabachwhatisep.shtml). • Evolutionary Psychology-Approaches in Psychology (http://www.psychegames.com/evolutionary-psychology. htm)

Academic societies • Human Behavior and Evolution Society (http://www.hbes.com); international society dedicated to using evolutionary theory to study human nature • The International Society for Human Ethology (http://evolution.anthro.univie.ac.at/ishe); promotes ethological perspectives on the study of humans worldwide • European Human Behaviour and Evolution Association (http://www.ehbea.com/) an interdisciplinary society that supports the activities of European researchers with an interest in evolutionary accounts of human cognition, behavior and society • The Association for Politics and the Life Sciences (http://www.aplsnet.org/); an international and interdisciplinary association of scholars, scientists, and policymakers concerned with evolutionary, genetic, and ecological knowledge and its bearing on political behavior, public policy and ethics. • Society for Evolutionary Analysis in Law (http://www.sealsite.org/) a scholarly association dedicated to fostering interdisciplinary exploration of issues at the intersection of law, biology, and evolutionary theory

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Evolutionary psychology • The New England Institute for Cognitive Science and Evolutionary Psychology (http://www.une.edu/nei/) aims to foster research and education into the interdisciplinary nexus of cognitive science and evolutionary studies • The NorthEastern Evolutionary Psychology Society (http://www.neepsociety.com/); regional society dedicated to encouraging scholarship and dialogue on the topic of evolutionary psychology • Feminist Evolutionary Psychology Society (http://fepsociety.org/) researchers that investigate the active role that females have had in human evolution

Journals • Evolutionary Psychology (http://www.epjournal.net/) free access online scientific journal • Evolution and Human Behavior (http://www.sciencedirect.com/science/journal/10905138); journal of the Human Behavior and Evolution Society (http://www.hbes.com) • Politics and the Life Sciences (http://www.politicsandthelifesciences.org/) is an interdisciplinary peer-reviewed journal published by the Association for Politics and the Life Sciences (http://www.aplsnet.org/ ) • Human Nature: An Interdisciplinary Biosocial Perspective (http://www.springer.com/social+sciences/ anthropology+and+archaeology/journal/12110) advances the interdisciplinary investigation of the biological, social, and environmental factors that underlie human behavior. It focuses primarily on the functional unity in which these factors are continuously and mutually interactive. These include the evolutionary, biological, and sociological processes as they interact with human social behavior. • Biological Theory: Integrating Development, Evolution and Cognition (http://www.kli.ac.at/publications-a. html) devoted to theoretical advances in the fields of biology and cognition, with an emphasis on the conceptual integration afforded by evolutionary and developmental approaches. • Evolutionary Anthropology (http://www3.interscience.wiley.com/cgi-bin/jtoc?ID=38641) • [[Behavioral and Brain Sciences (http://www.bbsonline.org/)] interdisciplinary articles in psychology, neuroscience, behavioral biology, cognitive science, artificial intelligence, linguistics and philosophy. About 30% of the articles have focused on evolutionary analyses of behavior. • Evolution and Development (http://www3.interscience.wiley.com/journal/118546131/home) Research relevant to interface of evolutionary and developmental biology • The Evolutionary Review – Art, Science, and Culture (http://www.evolutionaryreview.com/ed.htm)

Videos • Brief video clip re what EP is (from the "Evolution" PBS Series) (http://www.youtube.com/ watch?v=pEmX8Rim-hs) • TED talk (http://www.ted.com/index.php/talks/steven_pinker_chalks_it_up_to_the_blank_slate.html) by Steven Pinker about his book The Blank Slate: The Modern Denial of Human Nature • RSA talk (https://www.youtube.com/watch?v=PWHlvFiv70Q) by evolutionary psychologist Robert Kurzban on modularity of mind, based on his book Why Everyone (Else) is a Hypocrite • Richard Dawkins' lecture on natural selection and evolutionary psychology (http://www.youtube.com/ watch?v=BzJUCG7L9I4) • Evolutionary Psychology-Steven Pinker & Frans de Waal (http://www.youtube.com/watch?v=z3X5AuKE9rg) Audio recording • Stone Age Minds: A conversation with evolutionary psychologists Leda Cosmides and John Tooby (http://www. youtube.com/watch?v=nNW_B8EwgH4) • Margaret Mead and Samoa (http://video.google.com/videoplay?docid=4165874976901589227&q=margaret+ mead+and+samoa&total=8&start=0&num=10&so=0&type=search&plindex=0). Review of the nature vs. nurture debate triggered by Mead's book "Coming of Age in Samoa."

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Evolutionary psychology • Secrets of the Tribe (http://vimeo.com/18751423) Documents the conflicts between cultural and evolutionary anthropologists who have studied the Yanomamo tribes. • Video interview (http://video.google.com/videoplay?docid=3554279466299738997) with Steven Pinker by Robert Wright (journalist) discussing evolutionary psychology • Video interview (http://video.google.com/videoplay?docid=-4975549474851602314) with Edward O. Wilson by Robert Wright (journalist), contextualizing evolutionary psychology within science, politics, academics and philosophy

Human behavioral ecology Human behavioral ecology (HBE) or human evolutionary ecology applies the principles of evolutionary theory and optimization to the study of human behavioral and cultural diversity. HBE examines the adaptive design of traits, behaviors, and life histories of humans in an ecological context. One aim of modern human behavioral ecology is to determine how ecological and social factors influence and shape behavioral flexibility within and between human populations. Among other things, HBE attempts to explain variation in human behavior as adaptive solutions to the competing life-history demands of growth, development, reproduction, parental care, and mate acquisition. HBE overlaps with evolutionary psychology, human or cultural ecology, and decision theory. It is most prominent in disciplines such as anthropology and psychology where human evolution is considered relevant for a holistic understanding of human behavior or in economics where self-interest, methodological individualism, and maximization are key elements in modeling behavior Wikipedia:Disputed statement. It has been resisted in fields such as sociology and political science where the findings on human evolution are either ignored or regarded as irrelevant.[citation needed]

Evolutionary theory Human behavioral ecology rests upon a foundation of evolutionary theory. This includes aspects of both general evolutionary theory and established middle-level evolutionary theories, as well. Aspects of general evolutionary theory include: • Natural selection, the process by which individual organisms with favorable traits are more likely to survive and reproduce. • Sexual selection, the theory that competition for mates between individuals of the same sex results in differential mating and reproduction. • Kin selection, the changes in gene frequency across generations that are driven at least in part by interactions between related individuals, and • Inclusive fitness, the sum of an individual's own reproductive success, (natural and sexual selection), plus the effects the individual's actions have on the reproductive success of that individual's kin, (kin selection). Middle-level evolutionary theories used in HBE include: • The theory of parental investment, which predicts that the sex making the largest investment in lactation, nurturing and protecting offspring will be more discriminating in mating and that the sex that invests less in offspring will compete for access to the higher investing sex. • Parent-offspring conflict, which predicts that because the genetic interests of parents and offspring are not identical, offspring will be selected to manipulate their parents in order to ensure higher investment, and that, conversely, parents will be selected to manipulate their offspring. • The theory of reciprocal altruism, a form of altruism in which one organism provides a benefit to another in the expectation of future reciprocation.

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Human behavioral ecology • The Trivers–Willard hypothesis, which proposes that parents should invest more in the sex that gives them the greatest reproductive payoff (grandchildren) with increasing or marginal investment. • r/K selection theory, which, in ecology, relates to the selection of traits in organisms that allow success in particular environments. r-selected species - in unstable or unpredictable environments - produce many offspring, any individual one of which is unlikely to survive to adulthood, while K-selected species - in stable or predictable environments - invest more heavily in fewer offspring, each of which has a better chance of surviving to adulthood. • Evolutionary game theory, the application of population genetics-inspired models of change in gene frequency in populations to game theory. • Evolutionarily stable strategy, which refers to a strategy, which if adopted by a population, cannot be invaded by any competing alternative strategy.

Basic principles of HBE Ecological selectionism Ecological selectionism refers to the assumption that humans are highly flexible in their behaviors. Furthermore, it assumes that various ecological forces select for various behaviors that optimize humans' inclusive fitness in their given ecological context.

The piecemeal approach The piecemeal approach refers to taking a reductionist approach as opposed to a holistic approach in studying human socioecological behavior. Human behavioral ecologists assume that by taking complex social phenomena, (e.g., marriage patterns, foraging behaviors, etc.), and then breaking them down into sets of components involving decisions and constraints that they are in a better position to create models and make predictions involving human behavior. An example would be examining marriage systems by examining the ecological context, mate preferences, the distribution of particular characteristics within the population, and so forth.

Conditional strategies Human behavioral ecologists assume that what might be the most adaptive strategy in one environment might not be the most adaptive strategy in another environment. Conditional strategies, therefore, can be represented in the following statement: • In environmental context X, engage in adaptive strategy A. • In environmental context Y, engage in adaptive strategy B.

The phenotypic gambit The phenotypic gambit refers to the assumption that humans possess a high amount of phenotypic plasticity. Human behavioral ecologists attempt to control for culture, genetic variation, human cognition, and human phylogeny. It is not that human behavioral ecologists think that these concepts are irrelevant. It is simply that the primary focus of HBE is to discover correlations between variations of ecological contexts and variations of human behavior.

Modeling Theoretical models that human behavioral ecologists employ include, but are not limited to: • Optimal foraging theory, which states that organisms focus on consuming the most energy while expending the least amount of energy.

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Human behavioral ecology

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• Life history theory, which postulates that many of the physiological traits and behaviors of individuals may be best understood in relation to the key maturational and reproductive characteristics that define the life course. • Sex allocation theory, which predicts that parents should bias their reproductive investments toward the offspring sex generating the greatest fitness return. • The polygyny threshold model, which suggests that polygyny is driven by female choice of mates who control more resources relative to other potential mates in the population.

Further reading • Borgerhoff Mulder, M. & Schacht, R. (2012). Human Behavioural Ecology [1]. Nature Encyclopedia of Life Sciences. • Hames, R. (2001). Human Behavioral Ecology. [2] International Encyclopedia of the Social & Behavioral Sciences. Elsevier Science Ltd. • Cronk, L. (1991). Human behavioral ecology. Annual Review of Anthropology, 20, 25-53. • Smith, Eric Alden (1999). Three Styles in the Evolutionary Analysis of Human Behavior [3] in Lee Cronk, Napoleon Chagnon and William Irons Adaptation and Human Behavior: An Anthropological Perspective [4], 27-48, New York: Aldine de Gruyter. • Winterhalder, Bruce & Smith, Eric Alden (2000). Analysing Adaptive Strategies: Human Behavioral Ecology at Twenty-Five. [5] Evolutionary Anthropology: Issues, News, and Reviews, Volume 9, Issue 2.

External links • The Human Behavioral Ecology Bibliography [6] - Maintained by Kermyt G. Anderson

References [1] [2] [3] [4]

http:/ / www. academia. edu/ 2166350/ Human_Behavioural_Ecology http:/ / www. unl. edu/ rhames/ ms/ behavioral-ecology-hames. pdf#search='human%20behavioral%20ecology' http:/ / faculty. washington. edu/ easmith/ ThreeStyles. pdf http:/ / shopping. yahoo. com/ p:Adaptation%20and%20Human%20Behavior%3A%20An%20Anthropological%20Perspective:3000093488;_ylc=X3oDMTB1c21tcDhkBF9TAzk2NjMyOTA3B [5] http:/ / www. hbes. com/ HBES/ evolanth. pdf [6] http:/ / faculty-staff. ou. edu/ A/ Kermyt. G. Anderson-1/ HBE/

Prisoner's dilemma

Prisoner's dilemma The prisoner's dilemma (or prisoners' dilemma) is a canonical example of a game analyzed in game theory that shows why two individuals might not cooperate, even if it appears that it is in their best interests to do so. It was originally framed by Merrill Flood and Melvin Dresher working at RAND in 1950. Albert W. Tucker formalized the game with prison sentence rewards and gave it the name "prisoner's dilemma" (Poundstone, 1992), presenting it as follows: Two members of a criminal gang are arrested and imprisoned. Each prisoner is in solitary confinement with no means of speaking to or exchanging messages with the other. The police admit they don't have enough evidence to convict the pair on the principal charge. They plan to sentence both to a year in prison on a lesser charge. Simultaneously, the police offer each prisoner a Faustian bargain. Here's how it goes: • If A and B both confess the crime, each of them serves 2 years in prison • If A confesses but B denies the crime, A will be set free whereas B will serve 3 years in prison (and vice versa) • If A and B both deny the crime, both of them will only serve 1 year in prison Because betraying your partner (by confessing) always rewards more than cooperating with them, all purely rational self-interested prisoners would betray the other, and so the only possible outcome for two purely rational prisoners is for them to betray each other. The interesting part of this result is that pursuing individual reward logically leads both of the prisoners to betray, but they would get a better reward if they both cooperated. In reality, humans display a systematic bias towards cooperative behavior in this and similar games, much more so than predicted by simple models of "rational" self-interested action. A model based on a different kind of rationality, where people forecast how the game would be played if they formed coalitions and then they maximize their forecasts, has been shown to make better predictions of the rate of cooperation in this and similar games given only the payoffs of the game. There is also an extended "iterative" version of the game, where the classic game is played over and over between the same prisoners, and consequently, both prisoners continuously have an opportunity to penalize the other for previous decisions. If the number of times the game will be played is known to the players, then (by backward induction) two purely rational prisoners will betray each other repeatedly, for the same reasons as the classic version. In an infinite or unknown length game there is no fixed optimum strategy, and Prisoner's Dilemma tournaments have been held to compete and test algorithms. The prisoner's dilemma game can be used as a model for many real world situations involving cooperative behaviour. In casual usage, the label "prisoner's dilemma" may be applied to situations not strictly matching the formal criteria of the classic or iterative games: for instance, those in which two entities could gain important benefits from cooperating or suffer from the failure to do so, but find it merely difficult or expensive, not necessarily impossible, to coordinate their activities to achieve cooperation.

Strategy for the classic prisoners' dilemma The normal game is shown below:

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Prisoner's dilemma

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Prisoner B stays silent (cooperates) Prisoner B betrays (defects) Prisoner A stays silent (cooperates) Each serves 1 year

Prisoner A betrays (defects)

Prisoner A: 3 years Prisoner B: goes free

Prisoner A: goes free Prisoner B: 3 years

Each serves 2 years

Here, regardless of what the other decides, each prisoner gets a higher pay-off by betraying the other ("defecting"). The reasoning involves an argument by dilemma: B will either cooperate or defect. If B cooperates, A should defect, since going free is better than serving 1 year. If B defects, A should also defect, since serving 2 years is better than serving 3. So either way, A should defect. Parallel reasoning will show that B should defect. In traditional game theory, some very restrictive assumptions on prisoner behaviour are made. It is assumed that both understand the nature of the game, and that despite being members of the same gang, they have no loyalty to each other and will have no opportunity for retribution or reward outside the game. Most importantly, a very narrow interpretation of "rationality" is applied in defining the decision-making strategies of the prisoners. Given these conditions and the payoffs above, prisoner A will betray prisoner B. The game is symmetric, so Prisoner B should act the same way. Since both "rationally" decide to defect, each receives a lower reward than if both were to stay quiet. Traditional game theory results in both players being worse off than if each chose to lessen the sentence of his accomplice at the cost of spending more time in jail himself.

Generalized form The structure of the traditional Prisoners’ Dilemma can be generalized from its original prisoner setting. Suppose that the two players are represented by the colors, red and blue, and that each player chooses to either "Cooperate" or "Defect". If both players cooperate, they both receive the reward, R, for cooperating. If Blue defects while Red cooperates, then Blue receives the temptation, T payoff while Red receives the "sucker's", S, payoff. Similarly, if Blue cooperates while Red defects, then Blue receives the sucker's payoff S while Red receives the temptation payoff T. If both players defect, they both receive the punishment payoff P. This can be expressed in normal form:

Canonical PD payoff matrix Cooperate Defect Cooperate R, R Defect

T, S

S, T P, P

and to be a prisoner's dilemma game in the strong sense, the following condition must hold for the payoffs: T>R>P>S The payoff relationship R > P implies that mutual cooperation is superior to mutual defection, while the payoff relationships T > R and P > S imply that defection is the dominant strategy for both agents. That is, mutual defection is the only strong Nash equilibrium in the game (i.e., the only outcome from which each player could only do worse by unilaterally changing strategy). The dilemma then is that mutual cooperation yields a better outcome than mutual defection but it is not the rational outcome because the choice to cooperate, at the individual level, is not rational from a self-interested point of view.

Prisoner's dilemma

The iterated prisoners' dilemma If two players play prisoners' dilemma more than once in succession and they remember previous actions of their opponent and change their strategy accordingly, the game is called iterated prisoners' dilemma. In addition to the general form above, the iterative version also requires that 2R > T + S, to prevent alternating cooperation and defection giving a greater reward than mutual cooperation. The iterated prisoners' dilemma game is fundamental to certain theories of human cooperation and trust. On the assumption that the game can model transactions between two people requiring trust, cooperative behaviour in populations may be modeled by a multi-player, iterated, version of the game. It has, consequently, fascinated many scholars over the years. In 1975, Grofman and Pool estimated the count of scholarly articles devoted to it at over 2,000. The iterated prisoners' dilemma has also been referred to as the "Peace-War game". If the game is played exactly N times and both players know this, then it is always game theoretically optimal to defect in all rounds. The only possible Nash equilibrium is to always defect. The proof is inductive: one might as well defect on the last turn, since the opponent will not have a chance to punish the player. Therefore, both will defect on the last turn. Thus, the player might as well defect on the second-to-last turn, since the opponent will defect on the last no matter what is done, and so on. The same applies if the game length is unknown but has a known upper limit. Unlike the standard prisoners' dilemma, in the iterated prisoners' dilemma the defection strategy is counter-intuitive and fails badly to predict the behavior of human players. Within standard economic theory, though, this is the only correct answer. The superrational strategy in the iterated prisoners' dilemma with fixed N is to cooperate against a superrational opponent, and in the limit of large N, experimental results on strategies agree with the superrational version, not the game-theoretic rational one. For cooperation to emerge between game theoretic rational players, the total number of rounds N must be random, or at least unknown to the players. In this case 'always defect' may no longer be a strictly dominant strategy, only a Nash equilibrium. Amongst results shown by Robert Aumann in a 1959 paper, rational players repeatedly interacting for indefinitely long games can sustain the cooperative outcome.

Strategy for the iterated prisoners' dilemma Interest in the iterated prisoners' dilemma (IPD) was kindled by Robert Axelrod in his book The Evolution of Cooperation (1984). In it he reports on a tournament he organized of the N step prisoners' dilemma (with N fixed) in which participants have to choose their mutual strategy again and again, and have memory of their previous encounters. Axelrod invited academic colleagues all over the world to devise computer strategies to compete in an IPD tournament. The programs that were entered varied widely in algorithmic complexity, initial hostility, capacity for forgiveness, and so forth. Axelrod discovered that when these encounters were repeated over a long period of time with many players, each with different strategies, greedy strategies tended to do very poorly in the long run while more altruistic strategies did better, as judged purely by self-interest. He used this to show a possible mechanism for the evolution of altruistic behaviour from mechanisms that are initially purely selfish, by natural selection. The winning deterministic strategy was tit for tat, which Anatol Rapoport developed and entered into the tournament. It was the simplest of any program entered, containing only four lines of BASIC, and won the contest. The strategy is simply to cooperate on the first iteration of the game; after that, the player does what his or her opponent did on the previous move. Depending on the situation, a slightly better strategy can be "tit for tat with forgiveness." When the opponent defects, on the next move, the player sometimes cooperates anyway, with a small probability (around 1–5%). This allows for occasional recovery from getting trapped in a cycle of defections. The exact probability depends on the line-up of opponents. By analysing the top-scoring strategies, Axelrod stated several conditions necessary for a strategy to be successful.

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Prisoner's dilemma Nice The most important condition is that the strategy must be "nice", that is, it will not defect before its opponent does (this is sometimes referred to as an "optimistic" algorithm). Almost all of the top-scoring strategies were nice; therefore, a purely selfish strategy will not "cheat" on its opponent, for purely self-interested reasons first. Retaliating However, Axelrod contended, the successful strategy must not be a blind optimist. It must sometimes retaliate. An example of a non-retaliating strategy is Always Cooperate. This is a very bad choice, as "nasty" strategies will ruthlessly exploit such players. Forgiving Successful strategies must also be forgiving. Though players will retaliate, they will once again fall back to cooperating if the opponent does not continue to defect. This stops long runs of revenge and counter-revenge, maximizing points. Non-envious The last quality is being non-envious, that is not striving to score more than the opponent (note that a "nice" strategy can never score more than the opponent). The optimal (points-maximizing) strategy for the one-time PD game is simply defection; as explained above, this is true whatever the composition of opponents may be. However, in the iterated-PD game the optimal strategy depends upon the strategies of likely opponents, and how they will react to defections and cooperations. For example, consider a population where everyone defects every time, except for a single individual following the tit for tat strategy. That individual is at a slight disadvantage because of the loss on the first turn. In such a population, the optimal strategy for that individual is to defect every time. In a population with a certain percentage of always-defectors and the rest being tit for tat players, the optimal strategy for an individual depends on the percentage, and on the length of the game. In the strategy called Pavlov, win-stay, lose-switch, If the last round outcome was P,P, a Pavlov player switches strategy the next turn, which means P,P would be considered as a failure to cooperate.[citation needed] For a certain range of parametersWikipedia:Citing sources, Pavlov beats all other strategies by giving preferential treatment to co-players which resemble Pavlov. Deriving the optimal strategy is generally done in two ways: 1. Bayesian Nash Equilibrium: If the statistical distribution of opposing strategies can be determined (e.g. 50% tit for tat, 50% always cooperate) an optimal counter-strategy can be derived analytically.[1] 2. Monte Carlo simulations of populations have been made, where individuals with low scores die off, and those with high scores reproduce (a genetic algorithm for finding an optimal strategy). The mix of algorithms in the final population generally depends on the mix in the initial population. The introduction of mutation (random variation during reproduction) lessens the dependency on the initial population; empirical experiments with such systems tend to produce tit for tat players (see for instance Chess 1988), but there is no analytic proof that this will always occur. Although tit for tat is considered to be the most robust basic strategy, a team from Southampton University in England (led by Professor Nicholas Jennings [2] and consisting of Rajdeep Dash, Sarvapali Ramchurn, Alex Rogers, Perukrishnen Vytelingum) introduced a new strategy at the 20th-anniversary iterated prisoners' dilemma competition, which proved to be more successful than tit for tat. This strategy relied on cooperation between programs to achieve the highest number of points for a single program. The university submitted 60 programs to the competition, which were designed to recognize each other through a series of five to ten moves at the start.[3] Once this recognition was made, one program would always cooperate and the other would always defect, assuring the maximum number of points for the defector. If the program realized that it was playing a non-Southampton player, it

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Prisoner's dilemma would continuously defect in an attempt to minimize the score of the competing program. As a result,[4] this strategy ended up taking the top three positions in the competition, as well as a number of positions towards the bottom. This strategy takes advantage of the fact that multiple entries were allowed in this particular competition and that the performance of a team was measured by that of the highest-scoring player (meaning that the use of self-sacrificing players was a form of minmaxing). In a competition where one has control of only a single player, tit for tat is certainly a better strategy. Because of this new rule, this competition also has little theoretical significance when analysing single agent strategies as compared to Axelrod's seminal tournament. However, it provided the framework for analysing how to achieve cooperative strategies in multi-agent frameworks, especially in the presence of noise. In fact, long before this new-rules tournament was played, Richard Dawkins in his book The Selfish Gene pointed out the possibility of such strategies winning if multiple entries were allowed, but he remarked that most probably Axelrod would not have allowed them if they had been submitted. It also relies on circumventing rules about the prisoners' dilemma in that there is no communication allowed between the two players. When the Southampton programs engage in an opening "ten move dance" to recognize one another, this only reinforces just how valuable communication can be in shifting the balance of the game.

Continuous iterated prisoners' dilemma Most work on the iterated prisoners' dilemma has focused on the discrete case, in which players either cooperate or defect, because this model is relatively simple to analyze. However, some researchers have looked at models of the continuous iterated prisoners' dilemma, in which players are able to make a variable contribution to the other player. Le and Boyd found that in such situations, cooperation is much harder to evolve than in the discrete iterated prisoners' dilemma. The basic intuition for this result is straightforward: in a continuous prisoners' dilemma, if a population starts off in a non-cooperative equilibrium, players who are only marginally more cooperative than non-cooperators get little benefit from assorting with one another. By contrast, in a discrete prisoners' dilemma, tit for tat cooperators get a big payoff boost from assorting with one another in a non-cooperative equilibrium, relative to non-cooperators. Since nature arguably offers more opportunities for variable cooperation rather than a strict dichotomy of cooperation or defection, the continuous prisoners' dilemma may help explain why real-life examples of tit for tat-like cooperation are extremely rare in nature (ex. Hammerstein[5]) even though tit for tat seems robust in theoretical models.

Real-life examples These particular examples, involving prisoners and bag switching and so forth, may seem contrived, but there are in fact many examples in human interaction as well as interactions in nature that have the same payoff matrix. The prisoner's dilemma is therefore of interest to the social sciences such as economics, politics, and sociology, as well as to the biological sciences such as ethology and evolutionary biology. Many natural processes have been abstracted into models in which living beings are engaged in endless games of prisoner's dilemma. This wide applicability of the PD gives the game its substantial importance.

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Prisoner's dilemma

In environmental studies In environmental studies, the PD is evident in crises such as global climate change. It is argued all countries will benefit from a stable climate, but any single country is often hesitant to curb CO2 emissions. The immediate benefit to an individual country to maintain current behavior is perceived to be greater than the purported eventual benefit to all countries if behavior was changed, therefore explaining the current impasse concerning climate change. An important difference between climate change politics and the prisoner's dilemma is uncertainty. The pace at which pollution will change climate is not known precisely. The dilemma faced by government is therefore different from the prisoner's dilemma in that the payoffs of cooperation are largely unknown. This difference suggests states will cooperate much less than in a real iterated prisoner's dilemma, so that the probability of avoiding a climate catastrophe is much smaller than that suggested by a game-theoretical analysis of the situation using a real iterated prisoner's dilemma. Osang and Nandy provide a theoretical explanation with proofs for a regulation-driven win-win situation along the lines of Michael Porter's hypothesis, in which government regulation of competing firms is substantial.[6]

In psychology In addiction research/behavioral economics, George Ainslie points out that addiction can be cast as an intertemporal PD problem between the present and future selves of the addict. In this case, defecting means relapsing, and it is easy to see that not defecting both today and in the future is by far the best outcome, and that defecting both today and in the future is the worst outcome. The case where one abstains today but relapses in the future is clearly a bad outcome – in some sense the discipline and self-sacrifice involved in abstaining today have been "wasted" because the future relapse means that the addict is right back where he started and will have to start over (which is quite demoralizing, and makes starting over more difficult). The final case, where one engages in the addictive behavior today while abstaining "tomorrow" will be familiar to anyone who has struggled with an addiction. The problem here is that (as in other PDs) there is an obvious benefit to defecting "today", but tomorrow one will face the same PD, and the same obvious benefit will be present then, ultimately leading to an endless string of defections. John Gottman in his research described in "the science of trust" defines good relationships as those where partners know not to enter the (D,D) cell or at least not to get dynamically stuck there in a loop.

In economics Advertising is sometimes cited as a real life example of the prisoner’s dilemma. When cigarette advertising was legal in the United States, competing cigarette manufacturers had to decide how much money to spend on advertising. The effectiveness of Firm A’s advertising was partially determined by the advertising conducted by Firm B. Likewise, the profit derived from advertising for Firm B is affected by the advertising conducted by Firm A. If both Firm A and Firm B chose to advertise during a given period the advertising cancels out, receipts remain constant, and expenses increase due to the cost of advertising. Both firms would benefit from a reduction in advertising. However, should Firm B choose not to advertise, Firm A could benefit greatly by advertising. Nevertheless, the optimal amount of advertising by one firm depends on how much advertising the other undertakes. As the best strategy is dependent on what the other firm chooses there is no dominant strategy, which makes it slightly different than a prisoner's dilemma. The outcome is similar, though, in that both firms would be better off were they to advertise less than in the equilibrium. Sometimes cooperative behaviors do emerge in business situations. For instance, cigarette manufacturers endorsed the creation of laws banning cigarette advertising, understanding that this would reduce costs and increase profits across the industry.[7] This analysis is likely to be pertinent in many other business situations involving advertising.[citation needed] Without enforceable agreements, members of a cartel are also involved in a (multi-player) prisoners' dilemma. 'Cooperating' typically means keeping prices at a pre-agreed minimum level. 'Defecting' means selling under this minimum level, instantly taking business (and profits) from other cartel members. Anti-trust authorities want

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Prisoner's dilemma potential cartel members to mutually defect, ensuring the lowest possible prices for consumers.

In sport Doping in sport has been cited as an example of a prisoner's dilemma. If two competing athletes have the option to use an illegal and dangerous drug to boost their performance, then they must also consider the likely behaviour of their competitor. If neither athlete takes the drug, then neither gains an advantage. If only one does, then that athlete gains a significant advantage over their competitor (reduced only by the legal or medical dangers of having taken the drug). If both athletes take the drug, however, the benefits cancel out and only the drawbacks remain, putting them both in a worse position than if neither had used doping.

Multiplayer dilemmas Many real-life dilemmas involve multiple players.[8] Although metaphorical, Hardin's tragedy of the commons may be viewed as an example of a multi-player generalization of the PD: Each villager makes a choice for personal gain or restraint. The collective reward for unanimous (or even frequent) defection is very low payoffs (representing the destruction of the "commons"). The commons are not always exploited: William Poundstone, in a book about the prisoner's dilemma (see References below), describes a situation in New Zealand where newspaper boxes are left unlocked. It is possible for people to take a paper without paying (defecting) but very few do, feeling that if they do not pay then neither will others, destroying the system. Subsequent research by Elinor Ostrom, winner of the 2009 Sveriges Riksbank Prize in Economic Sciences in Memory of Alfred Nobel, hypothesized that the tragedy of the commons is oversimplified, with the negative outcome influenced by outside influences. Without complicating pressures, groups communicate and manage the commons among themselves for their mutual benefit, enforcing social norms to preserve the resource and achieve the maximum good for the group, an example of effecting the best case outcome for PD.

The Cold War The Cold War and similar arms races can be modelled as a Prisoner's Dilemma situation. During the Cold War the opposing alliances of NATO and the Warsaw Pact both had the choice to arm or disarm. From each side's point of view: Disarming whilst your opponent continues to arm would have led to military inferiority and possible annihilation. If both sides chose to arm, neither could afford to attack each other, but at the high cost of maintaining and developing a nuclear arsenal. If both sides chose to disarm, war would be avoided and there would be no costs. If your opponent disarmed while you continue to arm, then you achieve superiority. Although the 'best' overall outcome is for both sides to disarm, the rational course for both sides is to arm. This is indeed what happened. Both sides poured enormous resources into military research and armament for the next thirty years until Soviet President Mikhail Gorbachev and US President Ronald Reagan negotiated arms reductions and reform in the Soviet Union caused ideological differences to abate.

Related games Closed-bag exchange Hofstadter[9] once suggested that people often find problems such as the PD problem easier to understand when it is illustrated in the form of a simple game, or trade-off. One of several examples he used was "closed bag exchange": Two people meet and exchange closed bags, with the understanding that one of them contains money, and the other contains a purchase. Either player can choose to honor the deal by putting into his or her bag what he or she agreed, or he or she can defect by handing over an empty bag.

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In this game, defection is always the best course, implying that rational agents will never play. However, in this case both players cooperating and both players defecting actually give the same result, assuming there are no gains from trade, so chances of mutual cooperation, even in repeated games, are few.

Friend or Foe? Friend or Foe? is a game show that aired from 2002 to 2005 on the Game Show Network in the USA. It is an example of the prisoner's dilemma game tested on real people, but in an artificial setting. On the game show, three pairs of people compete. When a pair is eliminated, they play a game similar to the prisoner's dilemma to determine how the winnings are split. If they both cooperate (Friend), they share the winnings 50–50. If one cooperates and the other defects (Foe), the defector gets all the winnings and the cooperator gets nothing. If both defect, both leave with nothing. Notice that the payoff matrix is slightly different from the standard one given above, as the payouts for the "both defect" and the "cooperate while the opponent defects" cases are identical. This makes the "both defect" case a weak equilibrium, compared with being a strict equilibrium in the standard prisoner's dilemma. If you know your opponent is going to vote Foe, then your choice does not affect your winnings. In a certain sense, Friend or Foe has a payoff model between prisoner's dilemma and the game of Chicken. The payoff matrix is Cooperate Defect Cooperate 1, 1 Defect

2, 0

0, 2 0, 0

This payoff matrix has also been used on the British television programmes Trust Me, Shafted, The Bank Job and Golden Balls, and on the American shows Bachelor Pad and Take It All. Game data from the Golden Balls series has been analyzed by a team of economists, who found that cooperation was "surprisingly high" for amounts of money that would seem consequential in the real world, but were comparatively low in the context of the game.

Iterated Snowdrift Researchers from the University of Lausanne and the University of Edinburgh have suggested that the "Iterated Snowdrift Game" may more closely reflect real-world social situations. Although this model is actually a Chicken game, it will be described here. In this model, the risk of being exploited through defection is lower, and individuals always gain from taking the cooperative choice. The Snowdrift game imagines two drivers who are stuck on opposite sides of a snowdrift, each of whom is given the option of shovelling snow to clear a path, or remaining in their car. A player's highest payoff comes from leaving the opponent to clear all the snow by themselves, but the opponent is still nominally rewarded for their work. This may better reflect real world scenarios, the researchers giving the example of two scientists collaborating on a report, both of whom would benefit if the other worked harder. "But when your collaborator doesn’t do any work, it’s probably better for you to do all the work yourself. You’ll still end up with a completed project."

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Example Snowdrift Payouts (A, B) B cooperates B defects A cooperates

200, 200

100, 300

A defects

300, 100

0, 0

Example PD Payouts (A, B) B cooperates B defects A cooperates

200, 200

-100, 300

A defects

300, -100

0, 0

References [1] For example see the 2003 study "Bayesian Nash equilibrium; a statistical test of the hypothesis" (http:/ / econ. hevra. haifa. ac. il/ ~mbengad/ seminars/ whole1. pdf) for discussion of the concept and whether it can apply in real economic or strategic situations (from Tel Aviv University). [2] http:/ / www. ecs. soton. ac. uk/ ~nrj [3] http:/ / www. southampton. ac. uk/ mediacentre/ news/ 2004/ oct/ 04_151. shtml [4] The 2004 Prisoners' Dilemma Tournament Results (http:/ / www. prisoners'-dilemma. com/ results/ cec04/ ipd_cec04_full_run. html) show University of Southampton's strategies in the first three places, despite having fewer wins and many more losses than the GRIM strategy. (Note that in a PD tournament, the aim of the game is not to "win" matches – that can easily be achieved by frequent defection). It should also be pointed out that even without implicit collusion between software strategies (exploited by the Southampton team) tit for tat is not always the absolute winner of any given tournament; it would be more precise to say that its long run results over a series of tournaments outperform its rivals. (In any one event a given strategy can be slightly better adjusted to the competition than tit for tat, but tit for tat is more robust). The same applies for the tit for tat with forgiveness variant, and other optimal strategies: on any given day they might not 'win' against a specific mix of counter-strategies. An alternative way of putting it is using the Darwinian ESS simulation. In such a simulation, tit for tat will almost always come to dominate, though nasty strategies will drift in and out of the population because a tit for tat population is penetrable by non-retaliating nice strategies, which in turn are easy prey for the nasty strategies. Richard Dawkins showed that here, no static mix of strategies form a stable equilibrium and the system will always oscillate between bounds. [5] Hammerstein, P. (2003). Why is reciprocity so rare in social animals? A protestant appeal. In: P. Hammerstein, Editor, Genetic and Cultural Evolution of Cooperation, MIT Press. pp. 83–94. [6] Osang and Nandy 2003 (http:/ / faculty. smu. edu/ tosang/ pdf/ regln0803. pdf) [7] This argument for the development of cooperation through trust is given in The Wisdom of Crowds , where it is argued that long-distance capitalism was able to form around a nucleus of Quakers, who always dealt honourably with their business partners. (Rather than defecting and reneging on promises – a phenomenon that had discouraged earlier long-term unenforceable overseas contracts). It is argued that dealings with reliable merchants allowed the meme for cooperation to spread to other traders, who spread it further until a high degree of cooperation became a profitable strategy in general commerce [8] Gokhale CS, Traulsen A. Evolutionary games in the multiverse. Proceedings of the National Academy of Sciences. 2010 Mar 23;107(12):5500–4. [9] – see Ch.29 The Prisoner's Dilemma Computer Tournaments and the Evolution of Cooperation.

Prisoner's dilemma

Further reading • Robert Aumann, "Acceptable points in general cooperative n-person games", in R. D. Luce and A. W. Tucker (eds.), Contributions to the Theory 23 of Games IV, Annals of Mathematics Study 40, 287–324, Princeton University Press, Princeton NJ. • Axelrod, R. (1984). The Evolution of Cooperation. ISBN 0-465-02121-2 • Bicchieri, Cristina (1993). Rationality and Coordination. Cambridge University Press. • David M. Chess (1988). Simulating the evolution of behavior: the iterated prisoners' dilemma problem. Complex Systems, 2:663–670. • Dresher, M. (1961). The Mathematics of Games of Strategy: Theory and Applications Prentice-Hall, Englewood Cliffs, NJ. • Greif, A. (2006). Institutions and the Path to the Modern Economy: Lessons from Medieval Trade. Cambridge University Press, Cambridge, UK. • Rapoport, Anatol and Albert M. Chammah (1965). Prisoner's Dilemma. University of Michigan Press.

External links • Prisoner's Dilemma (Stanford Encyclopedia of Philosophy) (http://plato.stanford.edu/entries/ prisoner-dilemma/) • Iterated prisoner's dilemma game (http://www.iterated-prisoners-dilemma.net/) • The Bowerbird's Dilemma (http://www.msri.org/ext/larryg/pages/15.htm) The Prisoner's Dilemma in ornithology – mathematical cartoon by Larry Gonick. • Dixit, Avinash; Nalebuff, Barry (2008). "Prisoner's Dilemma" (http://www.econlib.org/library/Enc/ PrisonersDilemma.html). In David R. Henderson (ed.). Concise Encyclopedia of Economics (2nd ed.). Indianapolis: Library of Economics and Liberty. ISBN 978-0865976658. OCLC  237794267 (http://www. worldcat.org/oclc/237794267).

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Kin selection

Kin selection Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Kin altruism is altruistic behaviour whose evolution is driven by kin selection. Kin selection is an instance of inclusive fitness, which combines the number of offspring produced with the number an individual can produce by supporting others, such as siblings. Charles Darwin discussed the concept of kin selection in his 1859 book, The Origin of Species, where he reflected The co-operative behaviour of social insects like the honey bee on the puzzle of sterile social insects, such as honey bees, can be explained by kin selection. which leave reproduction to their sisters, arguing that a selection benefit to related organisms (the same "stock") would allow the evolution of a trait that confers the benefit but destroys an individual at the same time. R.A. Fisher in 1930 and J.B.S. Haldane in 1932 set out the mathematics of kin selection, with Haldane famously joking that he would willingly die for two brothers or eight cousins. In 1964, W.D. Hamilton popularized the concept and the major advance in the mathematical treatment of the phenomenon by George R. Price which has become known as "Hamilton's rule". In the same year John Maynard Smith used the actual term kin selection for the first time. According to Hamilton's rule, kin selection causes genes to increase in frequency when the genetic relatedness of a recipient to an actor multiplied by the benefit to the recipient is greater than the reproductive cost to the actor. The rule is difficult to test but was verified experimentally in 2010 by observing adoption of orphans by surrogate mothers in a wild red squirrel population. Hamilton proposed two mechanisms for kin selection: kin recognition, where individuals are able to identify their relatives, and viscous populations, where dispersal is rare enough for populations to be closely related. The viscous population mechanism makes kin selection and social cooperation possible in the absence of kin recognition. Nurture kinship, the treatment of individuals as kin when they live together, is sufficient for kin selection, given reasonable assumptions about dispersal rates. Kin selection is not the same thing as group selection, where natural selection acts on the group as a whole. In humans, altruism is more likely and on a larger scale with kin than with unrelated individuals; for example, humans give presents according to how closely related they are to the recipient. In other species, vervet monkeys use allomothering, where related females such as older sisters or grandmothers often care for young, according to their relatedness. The social shrimp Synalpheus regalis protects juveniles within highly related colonies.

Historical overview Charles Darwin was the first to discuss the concept of kin selection. In The Origin of Species, he wrote clearly about the conundrum represented by altruistic sterile social insects that This difficulty, though appearing insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be applied to the family, as well as to the individual, and may thus gain the desired end. Breeders of cattle wish the flesh and fat to be well marbled together. An animal thus characterized has been slaughtered, but the breeder has gone with confidence to the same stock and has succeeded. —Darwin In this passage "the family" and "stock" stand for a kin group. These passages and others by Darwin about "kin selection" are highlighted in D.J. Futuyma's textbook of reference Evolutionary Biology and inE. O. Wilson's

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Kin selection Sociobiology. The earliest mathematically formal treatments of kin selection were by R.A. Fisher in 1930 and J.B.S. Haldane in 1932 and 1955. J.B.S. Haldane fully grasped the basic quantities and considerations in kin selection, famously writing "I would lay down my life for two brothers or eight cousins".[1] Haldane's remark alluded to the fact that if an individual loses its life to save two siblings, four nephews, or eight cousins, it is a "fair deal" in evolutionary terms, as siblings are on average 50% identical by descent, nephews 25%, and cousins 12.5% (in a diploid population that is randomly mating and previously outbred). But Haldane also joked that he would truly die only to save more than a single identical twin of his or more than two full siblings. In 1955 he clarified: Let us suppose that you carry a rare gene that affects your behavior so that you jump into a flooded river and save a child, but you have one chance in ten of being drowned, while I do not possess the gene, and stand on the bank and watch the child drown. If the child's your own child or your brother or sister, there is an even chance that this child will also have this gene, so five genes will be saved in children for one lost in an adult. If you save a grandchild or a nephew, the advantage is only two and a half to one. If you only save a first cousin, the effect is very slight. If you try to save your first cousin once removed the population is more likely to lose this valuable gene than to gain it. … It is clear that genes making for conduct of this kind would only have a chance of spreading in rather small populations when most of the children were fairly near relatives of the man who risked his life. W. D. Hamilton, in 1963 and especially in 1964, popularized the concept and the more thorough mathematical treatment given to it by George Price. John Maynard Smith may have coined the actual term "kin selection" in 1964: These processes I will call kin selection and group selection respectively. Kin selection has been discussed by Haldane and by Hamilton. … By kin selection I mean the evolution of characteristics which favour the survival of close relatives of the affected individual, by processes which do not require any discontinuities in the population breeding structure. Kin selection causes changes in gene frequency across generations, driven by interactions between related individuals. This dynamic forms the conceptual basis of the theory of social evolution. Some cases of evolution by natural selection can only be understood by considering how biological relatives influence each other's fitness. Under natural selection, a gene encoding a trait that enhances the fitness of each individual carrying it should increase in frequency within the population; and conversely, a gene that lowers the individual fitness of its carriers should be eliminated. However, a hypothetical gene that prompts behaviour which enhances the fitness of relatives but lowers that of the individual displaying the behavior, may nonetheless increase in frequency, because relatives often carry the same gene. According to this principle, the enhanced fitness of relatives can at times more than compensate for the fitness loss incurred by the individuals displaying the behaviour, making kin selection possible. This is a special case of a more general model, "inclusive fitness". This analysis has been challenged,[2] Wilson writing that "the foundations of the general theory of inclusive fitness based on the theory of kin selection have crumbled" and that he now relies instead on the theory of eusociality and "gene-culture co-evolution" for the underlying mechanics of sociobiology. "Kin selection" should not be confused with "group selection" according to which a genetic trait can become prevalent within a group because it benefits the group as a whole, regardless of any benefit to individual organisms. All known forms of group selection conform to the principle that an individual behaviour can be evolutionarily successful only if the genes responsible for this behaviour conform to Hamilton's Rule, and hence, on balance and in the aggregate, benefit from the behaviour.

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Hamilton's rule Formally, genes should increase in frequency when

where r = the genetic relatedness of the recipient to the actor, often defined as the probability that a gene picked randomly from each at the same locus is identical by descent. B = the additional reproductive benefit gained by the recipient of the altruistic act, C = the reproductive cost to the individual performing the act. This inequality is known as Hamilton's rule after W. D. Hamilton who in 1964 published the first formal quantitative treatment of kin selection. The relatedness parameter (r) in Hamilton's rule was introduced in 1922 by Sewall Wright as a coefficient of relationship that gives the probability that at a random locus, the alleles there will be identical by descent. Subsequent authors, including Hamilton, sometimes reformulate this with a regression, which, unlike probabilities, can be negative. A regression analysis producing statistically significant negative relationships indicates that two individuals are less genetically alike than two random ones (Hamilton 1970, Nature & Grafen 1985 Oxford Surveys in Evolutionary Biology). This has been invoked to explain the evolution of spiteful behaviour consisting of acts that result in harm, or loss of fitness, to both the actor and the recipient. There has been little empirical support for Hamilton's rule in wild animals, as it is hard to quantify the costs and benefits of different behaviours. The first study to test Hamilton's rule successfully was in 2010, involving a wild population of red squirrels in Yukon, Canada. The researchers found that surrogate mothers would adopt related orphaned squirrel pups but not unrelated orphans. The researchers calculated the cost of adoption by measuring a decrease in the survival probability of the entire litter after increasing the litter by one pup, while benefit was measured as the increased chance of survival of the orphan. The degree of relatedness of the orphan and surrogate mother for adoption to occur depended on the number of pups the surrogate mother already had in her nest, as this affected the cost of adoption. The study showed that females always adopted orphans when rB > C, but never adopted when rB < C, providing strong support for Hamilton's rule.

Mechanisms Altruism occurs where the instigating individual suffers a fitness loss while the receiving individual experiences a fitness gain. The sacrifice of one individual to help another is an example. Hamilton (1964) outlined two ways in which kin selection altruism could be favoured: The selective advantage which makes behaviour conditional in the right sense on the discrimination of factors which correlate with the relationship of the individual concerned is therefore obvious. It may be, for instance, that in respect of a certain social action performed towards neighbours indiscriminately, an individual is only just breaking even in terms of inclusive fitness. If he could learn to recognise those of his neighbours who really were close relatives and could devote his beneficial actions to them alone an advantage to inclusive fitness would at once appear. Thus a mutation causing such discriminatory behaviour itself benefits inclusive fitness and would be selected. In fact, the individual may not need to perform any discrimination so sophisticated as we suggest here; a difference in the generosity of his behaviour according to whether the situations evoking it were encountered near to, or far from, his own home might occasion an advantage of a similar kind." (1996 [1964], 51) Kin recognition: First, if individuals have the capacity to recognize kin and to discriminate (positively) on the basis of kinship, then the average relatedness of the recipients of altruism could be high enough for kin selection. Because of the facultative nature of this mechanism, kin recognition and discrimination are expected to be unimportant except

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Kin selection among 'higher' forms of life (although there is some evidence for it among protozoa). Note also that kin recognition may be selected for inbreeding avoidance, and little evidence indicates that 'innate' kin recognition plays a role in mediating altruism. A thought experiment on the kin recognition/discrimination distinction is the hypothetical 'green beard', where a gene for social behaviour is imagined also to cause a distinctive phenotype that can be recognised by other carriers of the gene. Due to conflicting genetic similarity in the rest of the genome, there would be selection pressure for green-beard altruistic sacrifices to be suppressed, making common ancestry the most likely form of inclusive fitness. Viscous populations: Secondly, even indiscriminate altruism may be favoured in "viscous" populations with low rates or short ranges of dispersal. Here, social partners are typically genealogically close kin, and so altruism can flourish even in the absence of kin recognition and kin discrimination faculties — spatial proximity and circumstantial cues serving as a rudimentary form of discrimination. This suggests a rather general explanation for altruism. Directional selection always favours those with higher rates of fecundity within a certain population. Social individuals can often enhance the survival of their own kin by participating in and following the rules of their own group. Hamilton later modified his thinking to suggest that an innate ability to recognise actual genetic relatedness was unlikely to be the dominant mediating mechanism for kin altruism: But once again, we do not expect anything describable as an innate kin recognition adaptation, used for social behaviour other than mating, for the reasons already given in the hypothetical case of the trees.(Hamilton 1987, 425)[3] Hamilton's later clarifications often go unnoticed, and because of the long standing assumption that kin selection requires innate powers of kin recognition, some theorists have tried to clarify the position in recent work: In his original papers on inclusive fitness theory, Hamilton pointed out a sufficiently high relatedness to favour altruistic behaviours could accrue in two ways — kin discrimination or limited dispersal ( Hamilton, 1964, 1971,1972, 1975). There is a huge theoretical literature on the possible role of limited dispersal reviewed by Platt & Bever (2009) and West et al. (2002a), as well as experimental evolution tests of these models (Diggle et al., 2007; Griffin et al., 2004; Kümmerli et al., 2009 ). However, despite this, it is still sometimes claimed that kin selection requires kin discrimination (Oates & Wilson, 2001; Silk, 2002 ). Furthermore, a large number of authors appear to have implicitly or explicitly assumed that kin discrimination is the only mechanism by which altruistic behaviours can be directed towards relatives... [T]here is a huge industry of papers reinventing limited dispersal as an explanation for cooperation. The mistakes in these areas seem to stem from the incorrect assumption that kin selection or indirect fitness benefits require kin discrimination (misconception 5), despite the fact that Hamilton pointed out the potential role of limited dispersal in his earliest papers on inclusive fitness theory (Hamilton, 1964; Hamilton, 1971; Hamilton, 1972; Hamilton, 1975). (West et al. 2010, p.243 and supplement)[4] The assumption that kin recognition must be innate, and that cue-based mediation of social cooperation based on limited dispersal and shared developmental context are not sufficient, has obscured significant progress made in applying kin selection and inclusive fitness theory to a wide variety of species, including humans,[5] on the basis of cue-based mediation of social bonding and social behaviours.

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Kin selection and human social patterns Evolutionary psychologists, following Darwinian anthropologists' interpretation[6] of kin selection theory initially attempted to explain human altruistic behaviour through kin selection by stating that "behaviors that help a genetic relative are favored by natural selection." However, most Evolutionary psychologists recognize that this common shorthand formulation is inaccurate; [M]any misunderstandings persist. In many cases, they result from conflating "coefficient of relatedness" and Families are important in human behaviour, but kin selection may be based on closeness and other "proportion of shared genes," which is a short step from cues. the intuitively appealing—but incorrect—interpretation that "animals tend to be altruistic toward those with whom they share a lot of genes." These misunderstandings don’t just crop up occasionally; they are repeated in many writings, including undergraduate psychology textbooks—most of them in the field of social psychology, within sections describing evolutionary approaches to altruism. (Park 2007, p860)[7] As with the earlier sociobiological forays into the cross-cultural data, typical approaches are not able to find explanatory fit with the findings of ethnographers insofar that human kinship patterns are not necessarily built upon blood-ties. However, as Hamilton's later refinements of his theory make clear, it does not simply predict that genetically related individuals will inevitably recognise and engage in positive social behaviours with genetic relatives: rather, indirect context-based mechanisms may have evolved, which in historical environments have met the inclusive fitness criterion (see above section). Consideration of the demographics of the typical evolutionary environment of any species is crucial to understanding the evolution of social behaviours. As Hamilton himself puts it, "Altruistic or selfish acts are only possible when a suitable social object is available. In this sense behaviours are conditional from the start." (Hamilton 1987, 420). Under this perspective, and noting the necessity of a reliable context of interaction being available, the data on how altruism is mediated in social mammals is readily made sense of. In social mammals, primates and humans, altruistic acts that meet the kin selection criterion are typically mediated by circumstantial cues such as shared developmental environment, familiarity and social bonding.[8] That is, it is the context that mediates the development of the bonding process and the expression of the altruistic behaviours, not genetic relatedness per se. This interpretation thus is compatible with the cross-cultural ethnographic data and has been called nurture kinship.

Examples Eusociality (true sociality) is used to describe social systems with three characteristics: one is an overlap in generations between parents and their offspring, two is cooperative brood care, and the third characteristic is specialized castes of nonreproductive individuals. The social insects provide good examples of organisms with what appear to be kin selected traits. The workers of some species are sterile, a trait Experiment about kin selection that would not occur if individual selection was the only process at work. The relatedness coefficient r is abnormally high between the worker sisters in a colony of Hymenoptera due to haplodiploidy. Hamilton's rule is presumed to be satisfied because the benefits in fitness for the workers are believed to exceed the costs in terms of lost reproductive opportunity, though this has never been demonstrated empirically. There are competing hypotheses, as well, which may also explain the evolution of social behavior in such organisms.

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Kin selection Another example is preference among individuals in sun-tailed monkey communities. A study showed that maternal kin (kin related to by mothers) favoured each other, but that with relatives more distant than half-siblings, this bias dropped significantly. Alarm calls in ground squirrels are another example. While they may alert others of the same species to danger, they draw attention to the caller and expose it to increased risk of predation. Paul Sherman studied the alarm calls of ground squirrels, observing that the calls occurred most frequently when the caller had relatives nearby. In a similar study, John Hoogland followed individual males through different stages of life. Male prairie dogs modified their rate of calling when closer to kin. These behaviours show that self-sacrifice is directed towards close relatives, and that there is an indirect fitness gain. Alan Krakauer of University of California, Berkeley has studied kin selection in the courtship behavior of wild turkeys. Like a teenager helping her older sister prepare for a party, a subordinate turkey may help his dominant brother put on an impressive team display that is only of direct benefit to the dominant member.[9] Recent studies provide evidence that even certain plants can recognize and respond to kinship ties. Using sea rocket for her experiments, Susan Dudley at McMaster University in Canada compared the growth patterns of unrelated plants sharing a pot to plants from the same clone. She found that unrelated plants competed for soil nutrients by aggressive root growth. This did not occur with sibling plants. In the wood mouse (Apodemus sylvaticus), aggregates of spermatozoa form mobile trains, some of the spermatozoa undergo premature acrosome reactions that correlate to improved mobility of the mobile trains towards the female egg for fertilization. This association is thought to proceed as a result of a "green beard effect" in which the spermatozoa perform a kin-selective altruistic act after identifying genetic similarity with the surrounding spermatozoa.

Human examples Whether or not Hamilton's rule always applies, studies have demonstrated that relatedness is often important for human altruism in that humans are inclined to behave more altruistically toward kin than toward unrelated individuals.[10] Many people choose to live near relatives, exchange sizable gifts with relatives, and favour relatives in wills in proportion to their relatedness.

Experimental studies, interviews, and surveys A study interviewed several hundred women in Los Angeles to study patterns of helping between kin versus non-kin. While non-kin friends were willing to help one another, their assistance was far more likely to be reciprocal. The largest amounts of non-reciprocal help, however, were reportedly provided by kin. Additionally, more closely related kin were considered more likely sources of assistance than distant kin. Similarly, several surveys of American college students found that individuals were more likely to incur the cost of assisting kin when a high probability that relatedness and benefit would be greater than cost existed. Participants’ feelings of helpfulness were stronger toward family members than non-kin. Additionally, participants were found to be most willing to help those individuals most closely related to them. Interpersonal relationships between kin in general were more supportive and less Machiavellian than those between non-kin. In one experiment, the longer participants (from both the UK and the South African Zulus) held a painful skiing position, the more money or food was presented to a given relative. Participants repeated the experiment for individuals of different relatedness (parents and siblings at r = .5, grandparents, nieces, and nephews at r = .25, etc.). The results showed that participants held the position for longer intervals the greater the degree of relatedness between themselves and those receiving the reward.

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Observational studies A study of food-sharing practices on the West Caroline islets of Ifaluk determined that food-sharing was more common among people from the same islet, possibly because the degree of relatedness between inhabitants of the same islet would be higher than relatedness between inhabitants of different islets. When food was shared between islets, the distance the sharer was required to travel correlated with the relatedness of the recipient—a greater distance meant that the recipient needed to be a closer relative. The relatedness of the individual and the potential inclusive fitness benefit needed to outweigh the energy cost of transporting the food over distance. Humans may use the inheritance of material goods and wealth to maximize their inclusive fitness. By providing close kin with inherited wealth, an individual may improve his or her kin’s reproductive opportunities and thus increase his or her own inclusive fitness even after death. A study of a thousand wills found that the beneficiaries who received the most inheritance were generally those most closely related to the will’s writer. Distant kin received proportionally less inheritance, with the least amount of inheritance going to non-kin. A study of childcare practices among Canadian women found that respondents with children provide childcare reciprocally with non-kin. The cost of caring for non-kin was balanced by the benefit a woman received—having her own offspring cared for in return. However, respondents without children were significantly more likely to offer childcare to kin. For individuals without their own offspring, the inclusive fitness benefits of providing care to closely related children might outweigh the time and energy costs of childcare. Family investment in offspring among black South African households also appears consistent with an inclusive fitness model. A higher degree of relatedness between children and their caregivers frequently correlated with a higher degree of investment in the children, with more food, health care, and clothing being provided. Relatedness between the child and the rest of the household also positively associated with the regularity of a child’s visits to local medical practitioners and with the highest grade the child had completed in school. Additionally, relatedness negatively associated with a child’s being behind in school for his or her age. Observation of the Dolgan hunter-gatherers of northern Russia suggested that, while reciprocal food-sharing occurs between both kin and non-kin, there are larger and more frequent asymmetrical transfers of food to kin. Kin are also more likely to be welcomed to non-reciprocal meals, while non-kin are discouraged from attending. Finally, even when reciprocal food-sharing occurs between families, these families are often very closely related, and the primary beneficiaries are the offspring. Other research indicates that violence in families is more likely to occur when step-parents are present and that "genetic relationship is associated with a softening of conflict, and people's evident valuations of themselves and of others are systematically related to the parties' reproductive values". Numerous other studies suggest how inclusive fitness may work amongst different peoples, such as the Ye’kwana of southern Venezuela, the Gypsies of Hungary, and the doomed Donner Party of the United States.[11][12]

Non-human examples Vervet monkeys display kin selection between siblings, mothers and offspring, and grandparent-grandchild. These monkeys utilize allomothering, where the allomother is typically an older female sibling or a grandmother. Other studies have shown that individuals will act aggressively toward other individuals that were aggressive toward their relatives.[13][14] Synalpheus regalis is a eusocial shrimp that protects juveniles in the colony. By defending the young, the large defender shrimp can increase its inclusive fitness. Allozyme data revealed that relatedness within colonies is high, averaging 0.50, indicating that colonies in this species represent close kin groups.

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Criticism The theory of kin selection was criticized in two studies, one published in 1998 and another in 2002 in PNAS. Alonso and Schuck-Paim argue that the behaviours which kin selection attempts to explain are not altruistic (in pure Darwinian terms) because: (1) they may directly favour the performer as an individual aiming to maximize its progeny (so the behaviours can be explained as ordinary individual selection); (2) these behaviours benefit the group (so they can be explained as group selection); or (3) they are by-products of a developmental system of many "individuals" performing different tasks (like a colony of bees, or the cells of multicellular organisms, which are the focus of selection). They also argue that the genes involved in sex ratio conflicts could be treated as "parasites" of (already established) social colonies, not as their "promoters", and, therefore the sex ratio in colonies would be irrelevant to the transition to eusociality. Those papers were mostly ignored until they were re-edited by Martin Nowak, Corina Tarnita, and E. O. Wilson. These latter authors argue that Inclusive fitness theory is not a simplification over the standard approach. It is an alternative accounting method, but one that works only in a very limited domain. Whenever inclusive fitness does work, the results are identical to those of the standard approach. Inclusive fitness theory is an unnecessary detour, which does not provide additional insight or information. —Nowak, Tarnita, and Wilson They, like Alonso (1998) and Alonso and Schuck-Paim (2002) earlier, argue for a multi-level selection model instead. This aroused a strong response, including a rebuttal published in Nature from over a hundred researchers.[15]

References [1] (see also: Haldane's Wikiquote entry) [2] Martin Nowak, Corina Tarnita & EO Wilson "The evolution of eusociality" Nature 466 1057–1062(26 August 2010) [3] Hamilton, W.D. (1987) Discriminating nepotism: expectable, common and overlooked. In Kin recognition in animals, edited by D. J. C. Fletcher and C. D. Michener. New York: Wiley. [4] West et al. 2011. Sixteen common misconceptions about the evolution of cooperation in humans. Evolution and Social Behaviour 32 (2011) 231-262 [5] Holland, Maximilian. (2004) Social Bonding and Nurture Kinship: Compatibility between Cultural and Biological Approaches. London School of Economics, PhD Thesis (http:/ / papers. ssrn. com/ sol3/ papers. cfm?abstract_id=1791365) [6] Daly, M. and Wilson, M.I. (1999) An evolutionary psychological perspective on homicide. In Homicide Studies: A Sourcebook of Social Research, edited by D. Smith and M. Zahn. Thousand Oaks: Sage Publications [7] Park, J.H. (2007) Persistent Misunderstandings of Inclusive Fitness and Kin Selection: Their Ubiquitous Appearance in Social Psychology Textbooks. Evolutionary Psychology 5(4): 860-873. [8] Sherman et al (1997) Recognition Systems. In Behavioural Ecology, edited by J. R. Krebs and N. B. Davies. Oxford: Blackwell Scientific. [9] In the mating game, male wild turkeys benefit even when they do not get the girl (http:/ / www. berkeley. edu/ news/ media/ releases/ 2005/ 03/ 02_turkeys. shtml), Robert Sanders [10] Cartwright, J. (2000). Evolution and human behavior: Darwinian perspectives on human nature. Massachusetts: MIT Press. [11] Hughes, A.L. (1988). Evolution and human kinship. Oxford: Oxford University Press. [12] Dunbar, R. (2008). Kinship in biological perspective. In N.J. Allen, H. Callan, R. Dunbar, W. James (Eds.), Early human kinship: From sex to social reproduction (131-150). New Jersey: Blackwell Publishing Ltd. [13] Lee, P.C. “Sibships: Cooperation and Competition Among Immature Vervet Monkeys.” Primates. Vol 28(1): 47-59. 1987 [14] CFairbanks, Lynn A. “Reciprocal benefits of allomothering for female vervet monkeys.” Animal Behaviour. Vol 40: 553-562. 1990 [15] Abbot et al (2011) Inclusive fitness theory and eusociality. Nature 471: E1-E4 ()

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Further reading • Hamilton, W.D. (1964). "The Genetical Evolution of Social Behaviour. I". Journal of Theoretical Biology 7 (1): 1–16. doi: 10.1016/0022-5193(64)90038-4 (http://dx.doi.org/10.1016/0022-5193(64)90038-4). PMID  5875341 (http://www.ncbi.nlm.nih.gov/pubmed/5875341). • Hamilton, W.D. (1964). "The Genetical Evolution of Social Behaviour. II". Journal of Theoretical Biology 7 (1): 17–52. doi: 10.1016/0022-5193(64)90039-6 (http://dx.doi.org/10.1016/0022-5193(64)90039-6). PMID  5875340 (http://www.ncbi.nlm.nih.gov/pubmed/5875340). • Lucas, J.R.; Creel, S.R.; Waser, P.M. (1996). "How to Measure Inclusive Fitness, Revisited". Animal Behaviour 51 (1): 225–228. doi: 10.1006/anbe.1996.0019 (http://dx.doi.org/10.1006/anbe.1996.0019). • Madsen, E.A.; Tunney, R.J.; Fieldman, G.; Plotkin, H.C.; Dunbar, R.I.M.; Richardson, J.M.; McFarland, D. (2007). "Kinship and Altruism: a Cross-Cultural Experimental Study". British Journal of Psychology 98 (2): 339–359. doi: 10.1348/000712606X129213 (http://dx.doi.org/10.1348/000712606X129213). PMID  17456276 (http://www.ncbi.nlm.nih.gov/pubmed/17456276). • Queller, D.C. & Strassman, J.E. (2002) Quick Guide: Kin Selection (http://www.ruf.rice.edu/~evolve/pdf/ 20002001/CurBio2002_12_R832.pdf). Current Biology,12,R832. • West, S.A., Gardner, A. & Griffin, A.S. (2006) Quick Guide: Altruism (http://westgroup.biology.ed.ac.uk/ pdf/West_etal_06_altruism.pdf). Current Biology,16,R482-R483.

Social evolution Social evolution is a subdiscipline of evolutionary biology that is concerned with social behaviors that have fitness consequences for individuals other than the actor. Social behaviors can be categorized according to the fitness consequences they entail for the actor and recipient. • • • •

Mutually beneficial – a behavior that increases the direct fitness of both the actor and the recipient Selfish – a behavior that increases the direct fitness of the actor, but the recipient suffers a loss Altruistic – a behavior that increases the direct fitness of the recipient, but the actor suffers a loss Spiteful – a behavior that decreases the direct fitness of both the actor and the recipient

This classification was proposed by W. D. Hamilton.[citation needed] He proposes that natural selection favors mutually beneficial or selfish behaviors. Hamilton's insight was to show how kin selection could explain altruism and spite. Social evolution is also often regarded (especially, in the field of social anthropology) as evolution of social systems and structures.[1] In 2010, famed Harvard biologist E. O. Wilson, a founder of modern sociobiology, proposed a new theory of social evolution. He argued that the traditional approach of focusing on eusociality had limitations, which he illustrated primarily with examples from the insect world. A parallel theory of progressive social evolution has been advanced by followers of Herbert Spencer (1820–1903).[2] This theory rejects the conventional religious concept of human sin, which was based on the idea that, after the fall from grace, the human condition was eternally corrupt.[3]

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Notes [1] see, e.g., Evolution and culture. Ed. by Marshall David Sahlins and Elman Service. Ann Arbor, MI: Univ. of Michigan Press, 1960; Andrey Korotayev, Nikolay Kradin, Victor de Munck, and Valeri Lynsha. Alternatives of Social Evolution: An Introduction (http:/ / www. scribd. com/ doc/ 16861617/ Korotayev-Kradin-Lynsha-Alternatives-of-Social-Evolution-An-Introduction-Alternatives-of-Social-Evolution-Ed-by-NKradin-AKorotayev-Vde-M). Alternatives of Social Evolution. Vladivostok: Far Eastern Branch of the Russian Academy of Sciences, 2009. P.12-59. [2] http:/ / www. britannica. com/ EBchecked/ topic/ 551217/ social-evolution [3] http:/ / www. britannica. com/ EBchecked/ topic/ 222234/ Christian-fundamentalism/ 2530/ Origins?anchor=ref883412

References • Carver, Thomas Nixon (1935). The Essential Factors of Social Evolution. Chapter links, pp. ix (http://books. google.com/books?id=6EoW9jPkORAC&pg=PP13&dq=bl&source=gbs_selected_pages& cad=0_1#PPP13,M1)- xi. (http://books.google.com/books?id=6EoW9jPkORAC&pg=PP13&dq=bl& source=gbs_selected_pages&cad=0_1#PPP15,M1) • Frank, S.A. (1998). Foundations of social evolution. Princeton University Press, Princeton NJ. (http://stevefrank. org/foundations/foundations.pdf) • Hamilton, W.D. (1964). The genetical evolution of social behavior I and II. — Journal of Theoretical Biology 7: 1-16 and 17-52. • Korotayev, Andrey (2004). World Religions and Social Evolution of the Old World Oikumene Civilizations: A Cross-cultural Perspective (First Edition ed.). Lewiston, New York: Edwin Mellen Press. ISBN 0-7734-6310-0.

External links

• Alternatives of Social Evolution: An Introduction (http://www.scribd.com/doc/16861617/ Korotayev-Kradin-Lynsha-Alternatives-of-Social-Evolution-An-Introduction-Alternatives-of-Social-Evolution-Ed-by-NKradin-A • Carneiro R. Stellar Evolution and Social Evolution: A Study in Parallel Processes (http://www.socionauki.ru/ authors/carneiro_r/). Social Evolution & History 2005. Vol. 4(1), pp. 136-159

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Social neuroscience is an interdisciplinary field devoted to understanding how biological systems implement social processes and behavior, and to using biological concepts and methods to inform and refine theories of social processes and behavior. Humans are fundamentally a social species, rather than individualists. As such, Homo sapiens create emergent organizations beyond the individual—structures that range from dyads, families, and groups to cities, civilizations, and cultures. These emergent structures evolved hand in hand with neural and hormonal mechanisms to support them because the consequent social behaviors helped these organisms survive, reproduce, and care for offspring sufficiently long that they too survived to reproduce. The term "social neuroscience" can be traced to a publication entitled "Social Neuroscience Bulletin" that was published quarterly between 1988 and 1994. The term was subsequently popularized in an article by John Cacioppo and Gary Berntson, published in the American Psychologist in 1992.[1] Cacioppo and Berntson are considered as the legitimate fathers of social neuroscience. Still a young field, social neuroscience is closely related to affective neuroscience and cognitive neuroscience, focusing on how the brain mediates social interactions.[2]

Overview Traditional neuroscience has for many years considered the nervous system as an isolated entity and largely ignored influences of the social environments in which humans and many animal species live. In fact, we now recognize the considerable impact of social structures on the operations of the brain and body. These social factors operate on the individual through a continuous interplay of neural, neuroendocrine, metabolic and immune factors on brain and body, in which the brain is the central regulatory organ and also a malleable target of these factors.[3] Social neuroscience investigates the biological mechanisms that underlie social processes and behavior, widely considered one of the major problem areas for the neurosciences in the 21st century, and applies concepts and methods of biology to develop theories of social processes and behavior in the social and behavioral sciences. Social neuroscience capitalizes on biological concepts and methods to inform and refine theories of social behavior, and it

Social neuroscience uses social and behavioral constructs and data to advance theories of neural organization and function.[4][5] Throughout most of the 20th century, social and biological explanations were widely viewed as incompatible. But advances in recent years have led to the development of a new approach synthesized from the social and biological sciences. The new field of social neuroscience emphasizes the complementary relationship between the different levels of organization, spanning the social and biological domains (e.g., molecular, cellular, system, person, relational, collective, societal) and the use of multi-level analyses to foster understanding of the mechanisms underlying the human mind and behavior.

Methods A number of methods are used in social neuroscience to investigate the confluence of neural and social processes. These methods draw from behavioral techniques developed in social psychology, cognitive psychology, and neuropsychology), and are associated with a variety of neurobiological techniques including functional magnetic resonance imaging (fMRI), magnetoencephalography (MEG), positron emission tomography (PET), facial electromyography (EMG), transcranial magnetic stimulation (TMS), electroencephalography (EEG), event-related potentials (ERPs), electrocardiograms, electromyograms, endocrinology, immunology, galvanic skin response (GSR), single-cell recording, and studies of focal brain lesion patients.[6][7][8] [9] Animal models are also important to investigate the putative role of specific brain structures, circuits, or processes (e.g., the reward system and drug addiction). In addition, quantitative meta-analyses are important to move beyond idiosyncrasies of individual studies, and neurodevelopmental investigations can contribute to our understanding of brain-behavior associations.[10][11] Primarily psychological methods include performance-based measures that record response time and/or accuracy,[12] such as the Implicit Association Test;[13] observational measures such as preferential looking in infant studies; and, self-report measures, such as questionnaire and interviews.[14] Neurobiological methods can be grouped together into ones that measure more external bodily responses, electrophysiological methods, hemodynamic measures, and lesion methods. Bodily response methods include GSR (also known as skin conductance response (SCR)), facial EMG, and the eyeblink startle response. Electrophysiological methods include single-cell recordings, EEG, and ERPs. Hemodynamic measures, which, instead of directly measuring neural activity, measure changes in blood flow, include PET and fMRI. Lesion methods traditionally study brains that have been damaged via natural causes, such as strokes, traumatic injuries, tumors, neurosurgery, infection, or neurodegenerative disorders. In its ability to create a type of 'virtual lesion' that is temporary, TMS may also be included in this category.[15]

Society for social neuroscience A dinner to discuss the challenges and opportunities in the interdisciplinary field of social neuroscience at the Society for Neuroscience meeting (Chicago, November 2009) resulted in a series of meetings led by John Cacioppo and Jean Decety with social neuroscientists, psychologists, neuroscientists, psychiatrists and neurologists in Argentina, Australia, Chile, China, Colombia, Hong Kong, Israel, Japan, the Netherlands, New Zealand, Singapore, South Korea, Taiwan, the United Kingdom, and the United States. Social neuroscience was defined broadly as the interdisciplinary study of the neural, hormonal, cellular, and genetic mechanisms underlying the emergent structures that define social species. Thus, among the participants in these meetings were scientists who used a wide variety of methods in studies of animals as well as humans, and patients as well as normal participants. The consensus also emerged that a Society for Social Neuroscience should be established to give scientists from diverse disciplines and perspectives the opportunity to meet, communicate with, and benefit from the work of each other. The international, interdisciplinary Society for social neuroscience (http:/ / S4SN. org) was launched at the conclusion of these consultations in Auckland, New Zealand on 20 January 2010, and the inaugural meeting for the Society was held on November 12, 2010, the day prior to the 2010 Society for Neuroscience meeting (San Diego, CA).

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Social Neuroscience Journals • Social Neuroscience - Social Neuroscience [16]. • Social Cognitive and Affective Neuroscience - home page [17]. • The Journal of Cognitive Neuroscience publishes articles on Social Neuroscience, including a special issue [18] in 2004.

Social Neuroscience Sections • • • • •

JPSP published a special section [19] in 2003. Neuropsychologia published a special issue [20] in 2003. NeuroImage published a Special Section on Social Cognitive Neuroscience [21] in the December 2005 issue. Psychophysiology has published several articles dealing with Social Neuroscience. Biological Psychology regularly publishes social neuroscience work.

Key readings • Brune, M., Ribbert, H., & Schiefenhovel, W. (2003). The social brain: evolution and pathology. Hoboken, NJ: Wiley & Sons Ltd. • Cacioppo, J.T. (2002). Social neuroscience: Understanding the pieces fosters understanding the whole and vice versa. American Psychologist, 57, 819-831. • Cacioppo, J. T., & Berntson, G. G. (1992). Social psychological contributions to the decade of the brain: Doctrine of multilevel analysis. American Psychologist, 47, 1019-1028. • Cacioppo, J.T., Berntson, G.G., Sheridan, J.F., & McClintock, M.K. (2000). Multilevel integrative analyses of human behavior: social neuroscience and the complementing nature of social and biological approaches. Psychological Bulletin, 126, 829-843. • Cacioppo, John T.; Gary G. Berntson (2004). Social Neuroscience: Key Readings, [22]. Psychology Press [23]. ISBN 978-1-84169-099-5.. • Cacioppo, John T.; Penny S. Visser, Cynthia L. Pickett (eds.) (2005). Social Neuroscience: People Thinking about Thinking People. MIT Press. ISBN 0-262-03335-6.. • Cozolino, L. (2006). The Neuroscience of Human Relationships: Attachment And the Developing Social Brain. W. W. Norton & Company. • de Haan, M., & Gunnar, M.R. (2009). Handbook of Developmental Social Neuroscience. The Guilford Press. • Decety, J., & Cacioppo, J.T. (2011). Handbook of Social Neuroscience. New York: Oxford University Press. • Decety, J., & Ickes, W. (2009). The Social Neuroscience of Empathy. Cambridge: MIT press. • Emery, N.J. (2007). Cognitive Neuroscience of Social Behavior. Taylor & Francis. • Harmon-Jones, E.; P. Winkielman (2007). Social Neuroscience: Integrating Biological and Psychological Explanations of Social Behavior. Guilford Press. ISBN 978-1-59385-404-1.. • van Lange, P.A.M. (2006). Bridging social psychology: benefits of transdisciplinary Approaches. Mahwah, NJ: Lawrence Erlbaum Associates. • Wolpert, D. & Frith, C. (2004). The Neuroscience of Social Interactions: Decoding, Influencing, and Imitating the Actions of Others. Oxford: Oxford University Press.

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References [1] Cacioppo, J. T., & Berntson, G. G. (1992). Social psychological contributions to the decade of the brain: Doctrine of multilevel analysis. American Psychologist, 47, 1019-1028. [2] Cacioppo, J. T., Berntson, G. G., & Decety, J. (2010). Social neuroscience and its relation to social psychology. Social Cognition, 28, 675-684 [3] Cacioppo, J. T., Berntson, G. G., & Decety, J. (2011). A history of social neuroscience. In A. W. Kruglanski and W. Stroebe (Eds.), Handbook of the History of Social Psychology. New York: Psychology Press. [4] Cacioppo, J.T. et al. (2007). Social neuroscience: progress and implications for mental health. Perspectives on Psychological Science, 2, 99-123. [5] Cacioppo, J. T., & Decety, J. (2011). Social neuroscience: challenges and opportunities in the study of complex behavior. Annals of the New York Academy of Sciences, Epub ahead of print [6] Adolphs, R. (2003). Investigating the cognitive neuroscience of social behavior. Neuropsychologia, 41, 119-126. [7] Cacioppo, J.T., & Berntson, G.G. (2009), Handbook of Neuroscience for the Behavioral Science. New York: John Wiley and Sons. [8] Harmon-Jones, E., & Beer, J.S. (2009). Methods in Social Neuroscience. New York: The Guilford Press [9] Ward, J. (2012). The Student's Guide to Social Neuroscience. New York: Psychology Press [10] de Haan, M., & Gunnar, M.R. (2009). Handbook of Developmental Social Neuroscience. The Guilford Press. [11] Decety, J., & Cacioppo, T.T. (2010). Frontiers in human neuroscience, the golden triangle, and beyond. Perspectives on Psychological Science, 5(6), 767-771. [12] Ward, J. (2012). The Student's Guide to Social Neuroscience. New York: Psychology Press [13] Greenwald, A. G., McGhee, D. E., & Schwartz, J. L. K. (1998). Measuring individual differences in implicit cognition: The Implicit Association Test. Journal of Personality and Social Psychology, 74, 1464-1480. doi:10.1037//00223514.74.6.1464 [14] [15] [16] [17] [18] [19] [20] [21] [22] [23]

Ward, J. (2012). The Student's Guide to Social Neuroscience. New York: Psychology Press Ward, J. (2012). The Student's Guide to Social Neuroscience. New York: Psychology Press http:/ / www. psypress. com/ social-neuroscience-1747-0919/ http:/ / scan. oxfordjournals. org http:/ / jocn. mitpress. org/ content/ vol16/ issue10/ http:/ / content. apa. org/ journals/ psp/ 85/ 4 http:/ / www. sciencedirect. com/ science/ issue/ 4860-2003-999589997-366869/ http:/ / www. sciencedirect. com/ science/ journal/ 10538119/ 28/ 4 http:/ / www. cognitiveneurosciencearena. com/ social-neuroscience-9781841690995 http:/ / www. psypress. com

External links • Society for Social Neuroscience (http://s4sn.org/drupal/). • New Society for Social Neuroscience to help guide emerging field (http://news.uchicago.edu/news. php?asset_id=2189) from the University of Chicago News Office. • University of Chicago Center for Cognitive and Social Neuroscience (http://ccsn.uchicago.edu/). • "What is social neuroscience?" (http://www.informaworld.com/smpp/section?content=a743874678& fulltext=713240928) Introduction from the first issue (March 2006) of the journal Social Neuroscience (http:// www.psypress.com/social-neuroscience-1747-0919/) defining social neuroscience, listing the tools of social neuroscience, and addressing the impact of social neuroscience.

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Sociobiological theories of rape

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Sociobiological theories of rape Criminology and penology

Sociobiological theories of rape explores how evolutionary adaptation influences the psychology of rapists. Such theories are highly controversial, as traditional theories typically do not consider rape to be a behavioral adaptation. Some object to such theories on ethical, religious, political, or scientific grounds. Others argue that a correct knowledge of the causes of rape is necessary to develop effective preventive measures.

A Natural History of Rape The idea that rape evolved under some circumstances as a genetically advantageous behavioral adaptation was popularized by biologist Randy Thornhill and anthropologist Craig T. Palmer in their 2000 book A Natural History of Rape: Biological Bases of Sexual Coercion.

Animal coercive sex It has been noted that behavior resembling rape in humans is observed in the animal kingdom, including ducks and geese, bottlenose dolphins,[1] and chimpanzees.[2] Indeed in orangutans, close human relatives, copulations of this nature may account for up to half of all observed matings.[3] Such behaviours, referred to as ‘forced copulations’, involve an animal being approached and sexually penetrated whilst it struggles or attempts to escape. These observations of forced sex among animals are not controversial. What is controversial is the interpretation of these observations and the extension of theories based on them to humans. “Thornhill introduces this theory by describing the sexual behavior of scorpion flies. In which the male may gain sex from the female either by presenting a gift of food during courtship or without a nuptial offering, in which case force is necessary to restrain her.” [4]

Human rape It is hypothesized that rape is homologous to similar behavior in other animals. “Human rape appears not as an aberration but as an alternative gene-promotion strategy that is most likely to be adopted by the 'losers' in the competitive, harem-building struggle. If the means of access to legitimate, consenting sex is not available, then a male may be faced with the choice between force or genetic extinction.” Thornhill and Palmer write that "In short, a man can have many children, with little inconvenience to himself; a woman can have only a few, and with great effort." Females therefore tend to be more choosy with partners. Rape is seen as one potential strategy for males for achieving reproductive success. They point to several other factors indicating that rape may be a reproductive strategy. It is during the potentially childbearing years that women most often are rape victims. Rapists usually do not use more force than necessary to subdue their victims which is argued to be the case since physically injuring the victims would reduce the chance of reproduction. Furthermore, "In many cultures rape is treated as a crime against the victim's husband." Anthropologist Edward H. Hagen states in his Evolutionary Psychology FAQ [5] from 2002 that he believes there is no clear evidence for the hypothesis that rape is adaptive. He believes the adaptivity of rape is possible, but claims there is not enough evidence to be certain one way or the other. However, he encourages such evidence to be

Sociobiological theories of rape obtained: "Whether human males possess psychological adaptations for rape will only be answered by careful studies seeking evidence for such cognitive specializations. To not seek such evidence is like failing to search a suspect for a concealed weapon." He also describes some conditions in the ancestral environment during which the reproductive gains from rape may have outweighed the costs: • "High status males may be have been able to coerce matings with little fear of reprisal." • "Low status women (e.g., orphans) may have been particularly vulnerable to being raped because males need not have feared reprisals from the woman's family." • "During war, raping enemy women may have had few negative repercussions." • "Men who were low status, who were likely to remain low status, and who had few opportunities to invest in kin may have realized reproductive benefits that outweighed the considerable costs (e.g., reprisal by the woman's family)." McKibbin et al. (2008) argue that there may be several different types of rapists or rape strategies. One is rape by disadvantaged men who cannot get sex otherwise. Another is "specialized rapists" who are more sexually aroused from rape than from consensual sex. A third type is opportunistic rapists who switches between forced and consensual sex depending on circumstances. A fourth type is psychopathic rapists. A fifth type is partner rape due to sperm competition when the male suspects or knows that the female has had sex with another male. There are varying degrees of empirical support for the existence of each of these types. More generally they mention research finding that at least one-third of males "admit they would rape under specific conditions" and that other surveys find that many menWikipedia:Manual of Style (dates and numbers) state having coercive sexual fantasies. They, as have others, "propose that rape is a conditional strategy that may potentially be deployed by any man."

Women’s defenses Women may have developed several defenses against and strategies to avoid rape. One is a partner preference for men that are effective bodyguards against other men such as physically and socially dominant men (although there may also be other evolutionary reasons for such a preference). Another is great psychological pain which according to some research is greatest during the childbearing years. Other researchWikipedia:Avoid weasel words have argued that the emotional pain may cause the women to focus on the social circumstances that enabled the rape with the aim to prevent future rapes. Other researchWikipedia:Avoid weasel words have found that during the fertile phase of the menstrual cycle women perform fewer behaviors that may increase the risk of an assault. Studies have also found that sensitivity for potential coercive behaviors in males as well as handgrip strength (but only in a simulated coercive situation) increase during the fertile phase of the menstrual cycle. On the other hand, a 2003 study found that the frequency of pregnancy from rape is significantly higher than that of pregnancy in non-coercive intercourse, and advanced the hypothesis that male rapists disproportionately target women exhibiting biological indications of fertility.

Naturalistic fallacy Thornhill and Palmer write that "Rape is viewed as a natural, biological phenomenon that is a product of the human evolutionary heritage". They further state that by categorizing a behavior as "natural" and "biological" we do not in any way mean to imply that the behavior is justified or even inevitable. "Biological" means "of or pertaining to life," so the word applies to every human feature and behavior. But to infer from that, as many critics assert that Thornhill and Palmer do, that what is biological is somehow right or good, would be to fall into the so-called naturalistic fallacy. They make a comparison to "natural disasters as epidemics, floods and tornadoes". This shows that what can be found in nature is not always good and that measures should be and are taken against natural phenomena. They further argue that a good knowledge of the causes of rape, including evolutionary ones, are necessary in order to develop effective preventive measures.[]

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Sociobiological theories of rape Evolutionary psychologists McKibbin et al. argue that the claim that evolutionary theories are justifying rape, is a fallacy in the same way that it would be a fallacy, to accuse the scientists doing research on the causes of cancer, that they are justifying cancer. Instead, they say that understanding the causes of rape may help create preventive measures. Wilson et al. (2003) argue that evolutionary psychologists like Thornhill and Palmer use the naturalistic fallacy inappropriately to forestall legitimate discussion about the ethical implications of their theory. According to Thornhill and Palmer, a naturalistic fallacy is to infer ethical conclusions (e.g., rape is good) from (true or false) statements of fact (e.g., rape is natural). Wilson et al. point out that combining a factual statement with an ethical statement to derive an ethical conclusion is standard ethical reasoning, not a naturalistic fallacy, because the moral judgment is not deduced exclusively from the factual statement. They further argue that if one combines Thornhill and Palmer's factual premise that rape increases the fitness of a woman's offspring with the ethical premise that it is right to increase fitness of offspring, the resulting deductively valid conclusion is that rape has also positive effects and that its ethical status is ambiguous. Wilson et al. state that Thornhill and Palmer dismiss all ethical objections with the phrase 'naturalistic fallacy' although "it is Thornhill and Palmer who are thinking fallaciously by using the naturalistic fallacy in this way."

Preventing rape Thornhill and Palmer (2000) suggest a number of possible strategies for preventing rape. One example is explaining to males that they may have predispositions to misread the female invitation of sex. Generally, viewing rape as being due to a desire for domination, and not related to sexual desire, is harmful. One example being the claim that the way women dress will not affect the risk of rape. They argue that the much greater societal freedom of dating without supervision, and removal of many barriers between males and females, have created an environment that has also removed many earlier societal controls against rape. It is suggested that "men and women interact only in public places during the early stages of their relationships".

Victim counseling Counseling of rape victims may also be improved by evolutionary considerations, according to Thornhill and Palmer. They argue that the view that rape is due to a domination desire, cannot explain to the victim why the rapist seemed to be sexually motivated. Evolutionary considerations can also help explain the emotional pain felt, as well as the form it takes. They may also help the rape victim understand why the rape victim's partner may see the rape as a form of infidelity. They also argued that the victim's partner may be helped by such understanding, and be more able to change his reaction.

Criticism The 2003 book Evolution, Gender, and Rape, written in response to A Natural History of Rape, compiles the views of twenty-eight scholars in opposition to sociobiological theories of rape. One contributor, Michael Kimmel, criticizes Thornhill and Palmer's argument that female rape victims tend to be sexually attractive young women, rather than children or older women, contrary to what would be expected if rapists selected victims based on inability to resist. Kimmel argues that younger women are the least likely to be married and the most likely to be out on dates with men, and therefore are the most likely to be raped because of opportunity arising from social exposure and marital status. Palmer and Thornhill responded to these critics in an article in the journal Evolutionary Psychology. Smith et al. (2001) have criticized the scientific accuracy of Thornhill and Palmer's rape adaptation claim. In a study on the Aché in Paraguay, they found that the disadvantages of rape outweighed the reproductive advantages by a 10 to one margin. The study suggests that this would make it very unlikely that rape would be an adaptive behavior for increased reproduction.[6]

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Sociobiological theories of rape While defending the evolutionary psychology theory of rape against its more vehement critics, Vandermassen (2010) provides a critique of some aspects of the view. She characterises the view of Thornhill and Palmer as "extreme" (p. 736), as they fail to allow for the influence of any non-sexual motivations in the crime of rape. Vandermassen also notes two problems with the data cited by Thornhill and Palmer regarding the psychological trauma caused by the violence associated with rape: firstly, this data is inaccurately and confusingly presented in the book, often obscuring the fact that it does not support Thornhill and Palmer's "counterintuitive hypothesis" (p. 744) that more physical violence during rape is associated with less psychological pain. Secondly, more recent research has failed to support this hypothesis. A more moderate position, integrating the evolutionary psychology and feminist theories on rape, is presented by Vandermassen, based in part on the work of feminist evolutionary researcher Barbara Smuts. Hamilton (2008) has criticized Thornhill and Palmer's definition of rape as the coerced vaginal penetration of women of reproductive age. He has suggested that the exclusion of male rape, rape on women outside the reproductive age range, murderous rape, and non-vaginal forms of rape virtually guaranteed the confirmation of their hypothesis that rape is an evolved reproductive strategy and not a crime of violence. Hamilton has argued that evolutionary psychology fails to explain rape because, by evolutionary psychology's own criteria, an adaptation to rape children or men, or non-vaginal rape, would have been eliminated in the course of evolution because it did not confer reproductive advantage on our ancestors.

References [1] Connor, Richard and Vollmer, Nicole (ed. Buss, David). 2005. Sexual Coercion in Dolphin Consortships: A comparison with Chimpanzees, pp 218. [2] Akiko Matsumoto-Oda, Miya Hamai, Hitosige Hayaki, Kazuhiko Hosaka, Kevin D. Hunt, Eiiti Kasuya, Kenji Kawanaka, John C. Mitani, Hiroyuki Takasaki, and Yukio Takahata. 2007. Estrus Cycle Asynchrony in Wild Female Chimpanzees, Pan troglodytes schweinfurthii. [3] Wrangham, R., & Peterson, D. 1996. Demonic males. New York: Houghton Mifflin. [4] Wilson, Glenn. The Science of Sex: Glenn Wilson on Rape. The Great Sex Divide, pp. 128–131. http:/ / www. heretical. com/ wilson/ rape. html [5] http:/ / www. anth. ucsb. edu/ projects/ human/ evpsychfaq. html [6] Why Do We Rape, Kill and Sleep Around? (http:/ / www. thedailybeast. com/ newsweek/ 2009/ 06/ 19/ why-do-we-rape-kill-and-sleep-around. html), Sharon Begley, The Daily Beast

Further reading Forced sex in animals • Abele, L. and Gilchrist, S. (1977), "Homosexual rape and sexual selection in Acanthocephalan worms", Science 197: 81–83. • Barash, D. (1977), "Sociobiology of Rape in Mallards (Anas platyrhynchos): Responses of the Mated Male", Science 197: 788–789. Theories regarding rape in humans • McKibbin, W.F., Shackelford, T.K., Goetz, A.T., & Starratt, V.G. (2008). Why do men rape? An evolutionary psychological perspective. Review of General Psychology, 12, 86–97. Full text (http://www.toddkshackelford. com/downloads/McKibbin-et-al-RGP-2008.pdf) • Thornhill, R. and Palmer, C. (2000), A Natural History of Rape: Biological Bases of Sexual Coercion. Cambridge: MIT Press. ISBN 0-262-20125-9 • Thornhill, R. and Thornhill, N. (1983), "Human Rape: An Evolutionary Analysis", Ethology and Sociobiology 4:137–173. Responses to these theories • Fausto-Sterling, A. "Putting Woman in Her (Evolutionary) Place," in Myths of Gender. Basic Books, (1992). ISBN 0-465-04792-0

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Sociobiological theories of rape • Travis, C. B. (ed.) (2003) Evolution, Gender, and Rape. A Bradford Book, The MIT Press, Cambridge, MA. 454 pp., ISBN 0-262-70090-5. • Palmer, C. T. and Thornhill, R. (2003). A posse of good citizens brings outlaw evolutionists to justice. A response to Evolution, Gender, and Rape. Edited by Cheryl Brown Travis. (2003). Cambridge, MA: MIT Press. Evolutionary Psychology, 1: 10–27. http://www.epjournal.net/filestore/ep011027.pdf Other evidence • Chavanne, T. J. & Gallup, G. G., Jr. (1998) Variation in risk taking behavior among female college students as a function of the menstrual cycle. Evolution and Human Behavior, 19, 27–32. • Bandura, Albert. Social Learning Theory. http://tip.psychology.org/bandura.html CliffsNotes.com. Gender Stereotypes. 9 Dec 2010 http://www.cliffsnotes.com/study_guide/topicArticleId-26957,articleId-26896.html. • Malamuth, Neil M. & Check, Joseph. Repeated Exposure to Violent and Nonviolent Pornography: Likelihood of Raping Ratings and Laboratory Aggression Against Women. http://www.apa.org/divisions/div46/articles/ malamuth.pdf • Siegel, Larry J. Criminonlogy. Thomson & Wadswoth.Tenth Edition. 2009. (http://books.google.com/ books?id=9nH0kSUMwGIC&pg=PA294&lpg=PA294&dq=men+with+psychological+abnormalities+who+ rape&source=bl&ots=6gEcLBaRm9&sig=gC6G_tj_bb0wAcgxedGepRorH9Q&hl=en& ei=zTECTafGLIO8lQfskuGBCA&sa=X&oi=book_result&ct=result&resnum=1& ved=0CBMQ6AEwAA#v=onepage&q&f=false) • Thornhill, Randy & Palmer, Craig T. Why Men Rape. New York Academy of Sciences.JANUARY-FEBRUARY 2000. http://iranscope.ghandchi.com/Anthology/Women/rape.htm Webster, Murray & Rashotte, Lisa. Fixed Roles and Situated Actions. Sep2009, Vol. 61 Issue 5/6, p325-337, 13p, 1 Chart • Wilson, Glenn. The Science of Sex: Glenn Wilson on Rape. The Great Sex Divide, pp. 128–131. http://www. heretical.com/wilson/rape.html

External links • The Evolutionary Psychology FAQ entry on rape (http://www.anth.ucsb.edu/projects/human/epfaq/rape. html) • Insult to injury (http://www.newscientist.com/article.ns?id=dn907) 20 June 2001 New Scientist Print Edition by Matt Walker • Link between rape and pregnancy (http://news.bbc.co.uk/1/hi/health/1398894.stm) 20 June 2001 BBC News.

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Sociophysiology

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Sociophysiology Sociophysiology is the "interplay between society and physical functioning" (Freund 1988: 856) involving "collaboration of two neighboring sciences: physiology and sociology" (Mauss 1936: 373).[1] In other words, sociophysiology is physiological sociology, a special science that studies the physiological side of human (and other animals') interrelations (Zeliony 1912: 405–406).[2]

Interdisciplinary field of research In addition to having been termed an "interdisciplinary area for research, an area which demonstrates the concomitant relationship between physiology and social behavior" (Di Mascio et al. 1955: 4), sociophysiology may also be described as "social ethology" and "social energetics" (Waxweiler 1906: 62). That is, the "physiology of reactive phenomena caused by the mutual excitations of individuals of the same species" (Waxweiler 1906: 62).[3] The interdisciplinary nature of sociophysiology largely entails a "synthesis of psychophysiology and social interaction" (Adler 2002: 884) such that a "socio-psycho-biological study" (Mauss 1936: 386) of "biologico-sociological phenomena" (Mauss 1936: 385) may ensue.[4] Such "socio-psycho-biological study" has uncovered a "sharing of physiology between people involved in a meaningful interaction" (Adler 2002: 884), as well as "mutually responsive physiologic engagement having normative function in maintaining social cohesion and well-being in higher social animals" (Adler 2002: 885). This "mutually responsive physiologic engagement" brings into play the "close links uniting social phenomena to the biological phenomena from which they immediately derive" (Solvay 1906: 26).[5]

Interpersonal physiology Furthermore, sociophysiology explores the "intimate relationship and mutual regulation between social and physiological systems that is especially vital in human groups" (Barchas 1986: 210). In other words, sociophysiology studies the "physio- and psycho-energetic phenomena at the basis of social groupings" (Solvay 1906: 25).[6] Along these lines, Zeliony (1912) noted that The changes of society are the result of the activities of the nervous system. Excitations vary with the same animal and with the same class of animals. The problem of the socio-physiologist is to find out what are the excitors and what the inhibitors. Physiology gives the laws of the nervous system.... Thus the duty of the socio-physiologist is to give a description of the nervous processes of groups which have resulted in changes in the [physical and social] environment.

“

”

—Georgii Pavlovich Zeliony, summarized in Ellwood, C. A. (1916). Objectivism in sociology. American Journal of Sociology, vol. 22, no. 3, p. 297-298.

In addition, sociophysiology "describes structure-function relationships for body structures and interactive functions relevant to psychiatric illness" (Gardner 1997: 351), and also "assumes that psychiatric disorders are pathological variants of the motivation, emotions, and conflict involved in normal communicational processes" (Gardner and Price 1999: 247–248). Psychiatry, thus, involves the diagnosis and treatment of what Lilienfeld (1879: 280) termed "physiological social pathology", and may be classed as a subfield of sociophysiology, called "pathological sociophysiology" by Zeliony (1912: 405).[7] As summarized by Ellwood (1916), Zeliony thought that, in the future, A socio-physiological pathology will become necessary. Its field of observation will be the deviations from the norm which are observed either as a result of the pathological differences in the organism or as a result of other conditions, as in the insane or those addicted to the use of alcohol.

“

”

—Georgii Pavlovich Zeliony, Ellwood, C. A. (1916). Objectivism in sociology. American Journal of Sociology, vol. 22, no. 3, p. 298.

Ellwood (1916: 298) also noted that Zeliony's future sociophysiology, being a natural biological science, must be Darwinian.

Sociophysiology In short, sociophysiology is "reciprocal, interpersonal physiology" (Adler 2002: 885). Such interpersonal physiology may have implications in the realm of human politics. For example, the findings of a recent study "suggest that political attitudes vary with physiological traits linked to divergent manners of experiencing and processing environmental threats" (Oxley et al. 2008: 1669).

Notes and references Notes [1] "Collaboration de deux sciences voisines: physiologie, sociologie" (Mauss 1936: 373). Furthermore, Freund (1988) discusses the work of F. J. Buytendijk (1974), which "is one example of an effort to redefine mechanistic physiology in such a way as to include subjectivity and social existence as interrelated aspects of our physical being" (Freund 1988: 847), such that there exists a "mutually 'conditioning' relation between sociopsychological and biological factors. Social environments 'construct' bodies, which in turn have an impact on social behavior, and this behavior in turn further modifies bodies" (Freund 1988: 849). [2] "Die physische Seite der Wechselbeziehungen der Menschen kann nichtsdestoweniger Gegenstand einer besonderen Naturwissenschaft sein, welche ich als "physiologische Soziologie" oder besser "Soziophysiologie" zu bezeichnen vorschlage" (Zeliony 1912: 405-406). Zeliony’s 1912 paper — the earliest known which uses the term "sociophysiology" and its derivatives, e.g., Soziophysiologie, soziophysiologische, etc. — was originally delivered as a lecture to the Saint Petersburg Philosophical Society on March 19, 1909 (Zeliony 1912: 405). Although Zeliony (1912: 406) attributes the term "sociophysiology" to Russian philosopher A. I. Wedensky, Paul von Lilienfeld, in 1879, published a book titled Social Physiology, which uses throughout the unified concept, conjoined with a dash, "social-physiology" [social-physiologie] in addition to the modified substantive, separated by a space, "social physiology" [sociale Physiologie]. G. P. Zeliony, a worker in Pavlov's lab, took as the starting point for his "future sociophysiology," the study of the "reflexive interrelations" [reflektorischen Wechselbeziehungen] of humans and other animals (Zeliony 1912: 405; 412–413). [3] "Éthologie sociale; énergétique sociale — physiologie des phénomènes réactionnels dus aux excitations mutuelles des individus de même espèce" (Waxweiler 1906: 62). [4] "Étude socio-psycho-biologique" (Mauss 1936: 386); "phénomènes biologico-sociologiques" (Mauss 1936: 385). [5] "Liens étroits qui unissent les phénomènes sociologiques aux phénomènes biologiques dont ils dérivent immédiatement" (Solvay 1906: 26). [6] "Phénomènes physic- et psycho-énergétiques à la base des groupements sociaux" (Rolvaag 1906: 25). Such phenomena are now known to involve neurotransmitters, hormones, pheromones, the immune system, etc. [7] "Die physiologische Socialpathologie" (Lilienfeld 1879: 280); "Pathologische Soziophysiologie" (Zeliony 1912: 405).

References • Adler, H. M. (2002). The sociophysiology of caring in the doctor–patient relationship. Journal of General Internal Medicine, vol. 17, no. 11, pp. 883–890. • Barchas, P. R. (1986). A sociophysiological orientation to small groups. In E. J. Lawler, ed., Advances in Group Processes, vol. 3, pp. 209–246. Greenwich, CT: JAI Press. • Di Mascio, A., Boyd, R. W., Greenblatt, M., and H. C. Solomon. (1955). The psychiatric interview (a sociophysiologic study). Diseases of the Nervous System, vol. 16, no. 1, pp. 4–9. • Ellwood, C. A. (1916). Objectivism in sociology. American Journal of Sociology, vol. 22, no. 3, pp. 289–305. • Freund, P. E. S. (1988). Bringing society into the body: Understanding socialized human nature. Theory and Society, vol. 17, no. 6, pp. 839–864. • Gardner Jr., R. J. (1997). Sociophysiology as the basic science of psychiatry. Theoretical Medicine, vol. 18, no. 4, pp. 335–356. • Gardner Jr., R. J., and J. S. Price. (1999). Sociophysiology and depression. In T. E. Joiner and J. C. Coyne, eds., The Interactional Nature of Depression: Advances in Interpersonal Approaches. Washington, DC: American Psychological Association. • Lilienfeld, P. (1879). Die sociale Physiologie. Volume 4 of Gedanken über die Socialwissenschaft der Zukunft. Mitau: E. Behre’s Verlag. • Mauss, M. (1936). Les techniques du corps. Journal de Psychologie, vol. 32, nos. 3–4, 15 mars – 15 avril 1936. Reprinted in M. Mauss, Sociologie et anthropologie, Paris: PUF, 1950, pp. xxx-xxx. doi: 10.1522/cla.mam.tec (http://dx.doi.org/10.1522/cla.mam.tec)

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Sociophysiology • Oxley, D. R. et al. (2008). Political attitudes vary with physiological traits. Science 19 September 2008: Vol. 321. no. 5896, pp. 1667–1670. (http://www.sciencemag.org/cgi/content/abstract/321/5896/1667) doi: 10.1126/science.1157627 (http://dx.doi.org/10.1126/science.1157627) • Solvay, E. (1906). Note sur des formules d’introduction à l’énergétique physio- et psycho-sociologique. Fascicule 1 des Notes et Mémoires de l’Institut de Sociologie, Instituts Solvay, Parc Léopold, Bruxelles. Bruxelles et Leipzig: Misch et Thron. • Waxweiler, E. (1906). Esquisse d’une sociologie. Fascicule 2 des Notes et Mémoires de l’Institut de Sociologie, Instituts Solvay, Parc Léopold, Bruxelles. Bruxelles et Leipzig: Misch et Thron. • Zeliony, G. P. (1912). Über die zukünftige Soziophysiologie. Archiv für Rassen- und Gesellschafts-Biologie, vol. 9, no. 4, pp. 405–429.

Further reading • Barchas, P. R., ed. (1984). Social Hierarchies: Essays Toward a Sociophysiological Perspective. Westport, CT: Greenwood. • Barchas, P. R. and S. P. Mendoza, eds. (1984). Social Cohesion: Essays Toward a Sociophysiological Perspective. Westport, CT: Greenwood. • Buytendijk, F. J. (1974). Prolegomena to an Anthropological Physiology. Pittsburgh: Duquesne University Press. • Gardner Jr., R. J. and D. R. Wilson. (2004). Sociophysiology and evolutionary aspects of psychiatry. In J. Panksepp, ed., Textbook of Biological Psychiatry. Wiley. (http://books.google.com/ books?id=yeLKvgbrcxgC&dq=Textbook+of+Biological+Psychiatry&pg=PP1&ots=8nCzkG-VNd& sig=Eq02sTs7QQ-qEIp-ZIOdl91qvpA&prev=http://www.google.com/search?client=firefox-a&rls=org. mozilla%3Aen-US%3Aofficial&channel=s&hl=en&q=Textbook+of+Biological+Psychiatry&btnG=Google+ Search&sa=X&oi=print&ct=title#PPP1,M1) • Mysterud, I. (2004). One name for the evolutionary baby? A preliminary guide for everyone confused by the chaos of names. Social Science Information, vol. 43, no. 1, pp. 95–114. • Perec, G. (1976) Lire: esquisse socio-physiologique. Esprit, no. 453, pp. 9–20. Reprinted in G. Perec, Penser/Classer, Paris: Hachette, 1985. • Reinheimer, H. (1920). Symbiosis: A Socio-physiological Study of Evolution. London: Headley Brothers. (Reinheimer seems to use the term "socio-physiology" as a synonym for "eugenics," which usage differs vastly from that of other writers, both early and late.) (http://www.archive.org/details/symbiosissocioph00reinrich) • Waid, W. M., ed. (1984). Sociophysiology. New York: Springer Verlag.

External links • Biozentrum Universität Würzburg: Research: Behavioral Physiology and Sociobiology. (http://www. biozentrum.uni-wuerzburg.de/research0.html) • Universität Bayreuth, Studies on behavioural ecology and sociophysiology in European rabbits. (http://www. uni-bayreuth.de/forschungsberichte/05/2/1/07/00/engl.html) • Hominisation-Anton Fürlinger, Social Brain and Sociophysiology. (http://www.hominisation.at/story.htm) • The Institute of Sociophysiology takes a pataphysical approach to the subject. (http://www.isocphys.org)

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E. O. Wilson

110

E. O. Wilson E. O. Wilson

February, 2003 Born

Edward Osborne Wilson June 10, 1929 Birmingham, Alabama, United States

Nationality

American

Fields

Biologist

Institutions

Harvard University

Alma mater

University of Alabama Harvard University

Thesis

A Monographic Revision of the Ant Genus Lasius

[1]

(1955)

Doctoral students Daniel Simberloff Donald J. Farish Known for

Popularizing sociobiology Epic of Evolution Character displacement Island biogeography

Notable awards

Pulitzer Prize (1979) Crafoord Prize (1990) Pulitzer Prize (1991) Kistler Prize (2000) Nierenberg Prize (2001)

Edward Osborne "E. O." Wilson (born June 10, 1929) is an American biologist, researcher (sociobiology, biodiversity), theorist (consilience, biophilia), naturalist (conservationist) and author. His biological specialty is myrmecology, the study of ants, on which he is considered to be the world's leading authority.[2] Wilson is known for his scientific career, his role as "the father of sociobiology", his environmental advocacy, and his secular-humanist and deist ideas pertaining to religious and ethical matters. Wilson was the Joseph Pellegrino University Research Professor in Entomology for the Department of Organismic and Evolutionary Biology at Harvard University and a Fellow of the Committee for Skeptical Inquiry. He is a Humanist Laureate of the International Academy of Humanism.[3][4] He is a two-time winner of the Pulitzer Prize for General Non-Fiction and a New York Times bestseller for The Social Conquest of Earth[5] and Letters to a Young Scientist.[6]

E. O. Wilson

Early life Wilson was born in Birmingham, Alabama. According to his autobiography Naturalist, he grew up mostly around Washington, D.C. and in the countryside around Mobile, Alabama.[7] From an early age, he was interested in natural history. His parents, Edward and Inez Wilson, divorced when he was seven. The young naturalist grew up in several cities and towns, moving around with his father and his stepmother. In the same year that his parents divorced, Wilson blinded himself in one eye in a fishing accident. He suffered for hours, but he continued fishing. He did not complain because he was anxious to stay outdoors. He never went in for medical treatment. Several months later, his right pupil clouded over with a cataract. He was admitted to Pensacola Hospital to have the lens removed. Wilson writes, in his autobiography, that “[t]he surgery was a terrifying [19th] century ordeal.” Today, he suffers from the phobia of being enclosed in a “closed space with [his] arms immobilized and [his] face covered with an obstruction.” Wilson was left with full sight in his left eye, with a vision of 20/10. He lost his stereoscopy, but he could see fine print and the hairs on the body of small insects. His reduced ability to observe mammals and birds led him to concentrate on insects. At nine, Wilson undertook his first expeditions at the Rock Creek Park in Washington, DC. He began to collect insects and he gained a passion for butterflies. He would capture them using nets made with brooms, coat hangers, and cheesecloth bags. Going on these expeditions lead to Wilson’s fascination with ants. He describes in his autobiography how one day he pulled the bark of a rotting tree away and discovered citronella ants underneath. The worker ants he found were “short, fat, brilliant yellow, and emitted a strong lemony odor. Wilson said the event left a “vivid and lasting impression on [him].” He also earned the Eagle Scout award and served as Nature Director of his Boy Scout summer camp. At the age of 18, intent on becoming an entomologist, he began by collecting flies, but the shortage of insect pins caused by World War II caused him to switch to ants, which could be stored in vials. With the encouragement of Marion R. Smith, a myrmecologist from the National Museum of Natural History in Washington, Wilson began a survey of all the ants of Alabama. This study led him to report the first colony of fire ants in the US, near the port of Mobile.[8] Concerned that he might not be able to afford to go to a university, Wilson attempted to enlist in the United States Army. His plan was to earn U.S. government financial support for his education, but he failed his Army medical examination due to his impaired eyesight. Wilson was able to afford to enroll in the University of Alabama after all. There, he earned his B.S. and M.S. degrees In Biology. He later earned his Ph.D. degree in Biology from Harvard University.

Retirement In 1996 he officially retired from teaching at Harvard and continues to hold the positions of Professor Emeritus and Honorary Curator in Entomology. He and his wife Irene now reside in Lexington, Massachusetts. His daughter, Catherine, and her husband Jonathan, reside in nearby Stow.

Theories and beliefs Epic of evolution "The evolutionary epic," Wilson wrote in his book On Human Nature, "is probably the best myth we will ever have." Wilson's intended usage of the word "myth" does not denote falsehood - rather, a grand narrative that provides people with placement in time—a meaningful placement that celebrates extraordinary moments of shared heritage. Wilson was not the first to use the term, but his fame prompted its usage as the morphed phrase epic of evolution. Wilson explained the need for the epic of evolution:[9] Human beings must have an epic, a sublime account of how the world was created and how humanity became part of it... Religious epics satisfy another primal need. They confirm we are part of something greater than ourselves... The way to achieve our epic that unites human spirituality, instead of cleave it,

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E. O. Wilson it is to compose it from the best empirical knowledge that science and history can provide. The worth of the epic, he said, is that "[t]he true evolutionary epic retold as poetry, is as intrinsically ennobling as any religious epic."[10] Cosmologist Brian Swimme concludes in a 1997 interview:[11] I think that what E. O. Wilson is trying to suggest is that to be fully human, a person has to see that life has a heroic dimension... I think for the scientist, and for other people, it's a question of, "Is the universe valuable? Is it sacred? Is it holy? Or is the human agenda all that matters?" I just don't think we're that stupid to continue in a way that continues to destroy. I'm hopeful that the Epic of Evolution will be yet another strategy in our culture that will lead our consciousness out of a very tight, human-centered materialism. Naturalistic and liberal religious writers have picked up on Wilson's term and have used it in a number of texts. These authors however have at times used other terms to refer to the idea: Universe Story (Brian Swimme, John F. Haught), Great Story (Connie Barlow, Michael Dowd), Everybody's Story (Loyal Rue), New Story (Thomas Berry, Al Gore, Brian Swimme) and Cosmic Evolution (Eric Chaisson).[12]

Sociobiology Michael McGoodwin paraphrasing and quoting Wilson (pp. 16 and 222) on sociobiology:[13] Sociobiology is defined as the scientific or systematic study of the biological basis of all forms of social behavior, in all kinds of organisms including man, and incorporating knowledge from ethology, ecology, and genetics, in order to derive general principles concerning the biological properties of entire societies. "If humankind evolved by Darwinian natural selection, genetic chance and environmental necessity, not God, made the species." "The brain [and the mind] exists because it promotes the survival and multiplication of the genes that direct its assembly." The two apparent dilemmas we face therefore are: (1) We lack any goal external to our biological nature (for even religions evolve to enhance the persistence and influence of their practitioners). Will the transcendental goals of societies dissolve, and will our post-ideological societies regress steadily toward self-indulgence? (2) Morality evolved as instinct. "Which of the censors and motivators should be obeyed and which ones might better be curtailed or sublimated?" Although much human diversity in behavior is culturally influenced, some has been shown to be genetic - rapid acquisition of language, human unpredictability, hypertrophy (extreme growth of pre-existing social structures), altruism and religions. "Religious practices that consistently enhance survival and procreation of the practitioners will propagate the physiological controls that favor the acquisition of the practices during single lifetimes." Unthinking submission to the communal will promotes the fitness of the members of the tribe. Even submission to secular religions and cults involve willing subordination of the individual to the group. Religious practices confer biological advantages. Wilson used sociobiology and evolutionary principles to explain the behavior of the social insects and then to understand the social behavior of other animals, including humans, thus established sociobiology as a new scientific field. He argued that all animal behavior, including that of humans, is the product of heredity, environmental stimuli, and past experiences, and that free will is an illusion. He has referred to the biological basis of behaviour as the "genetic leash."[14] The sociobiological view is that all animal social behavior is governed by epigenetic rules worked out by the laws of evolution. This theory and research proved to be seminal, controversial, and influential.[15] The controversy of sociobiological research is in how it applies to humans. The theory established a scientific argument for rejecting the common doctrine of tabula rasa, which holds that human beings are born without any innate mental content and that culture functions to increase human knowledge and aid in survival and success. In the final chapter of the book Sociobiology and in the full text of his Pulitzer Prize-winning On Human Nature, Wilson

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E. O. Wilson argues that the human mind is shaped as much by genetic inheritance as it is by culture (if not more). There are limits on just how much influence social and environmental factors can have in altering human behavior.

Ants and social insects Wilson, along with Bert Hölldobler, has done a systematic study of ants and ant behavior, culminating in their encyclopedic work, The Ants (1990). Because much self-sacrificing behavior on the part of individual ants can be explained on the basis of their genetic interests in the survival of the sisters, with whom they share 75% of their genes (though the actual case is some species' queens mate with multiple males and therefore some workers in a colony would only be 25% related), Wilson was led to argue for a sociobiological explanation for all social behavior on the model of the behavior of the social insects. In his more recent work, he has sought to defend his views against the criticism of younger scientists such as Deborah Gordon, whose results challenge the idea that ant behavior is as rigidly predictable as Wilson's explanations make it. Edward O. Wilson, referring to ants, once said that "Karl Marx was right, socialism works, it is just that he had the wrong species", meaning that while ants and other eusocial species appear to live in communist-like societies, they only do so because they are forced to do so from their basic biology, as they lack reproductive independence: worker ants, being sterile, need their ant-queen to survive as a colony and a species and individual ants cannot reproduce without a queen, thus being forced to live in centralised societies. Humans, however, do possess reproductive independence so they can give birth to offspring without the need of a "queen", and in fact humans enjoy their maximum level of Darwinian fitness only when they look after themselves and their offspring, while finding innovative ways to use the societies they live in for their own benefit.

Consilience In his 1998 book Consilience: The Unity of Knowledge, Wilson discusses methods that have been used to unite the sciences, and might be able to unite the sciences with the humanities. Wilson prefers and uses the term "consilience" to describe the synthesis of knowledge from different specialized fields of human endeavor. He defines human nature as a collection of epigenetic rules, the genetic patterns of mental development. He argues that culture and rituals are products, not parts, of human nature. He says art is not part of human nature, but our appreciation of art is. He argues that concepts such as art appreciation, fear of snakes, or the incest taboo (Westermarck effect) can be studied by scientific methods of the natural sciences. Previously, these phenomena were only part of psychological, sociological, or anthropological studies. Wilson proposes that they can be part of interdisciplinary research.

The unit and target of selection Wilson has argued that the "unit of selection is a gene, the basic element of heredity. The target of selection is normally the individual who carries an ensemble of genes of certain kinds." With regard to the use of kin selection in explaining the behavior of eusocial insects, Wilson said to Discover magazine, the "new view that I'm proposing is that it was group selection all along, an idea first roughly formulated by Darwin."[16]

Spiritual and political beliefs Views on religion As paraphrased by Michael McGoodwin The predisposition to religious belief is an ineradicable part of human behavior. Mankind has produced 100,000 religions. It is an illusion to think that scientific humanism and learning will dispel religious belief. Men would rather believe than know... A kind of Darwinistic survival of the fittest has occurred with religions... The ecological principle called Gause's law holds that competition is maximal between species with identical needs... Even submission to secular religions such as Communism and guru cults

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E. O. Wilson involve willing subordination of the individual to the group. Religious practices confer biological advantage. The mechanisms of religion include (1) objectification (the reduction of reality to images and definitions that are easily understood and cannot be refuted), (2) commitment through faith (a kind of tribalism enacted through self-surrender), (3) and myth (the narratives that explain the tribe's favored position on the earth, often incorporating supernatural forces struggling for control, apocalypse, and millennium). The three great religion categories of today are Marxism, traditional religion, and scientific materialism... Though theology is not likely to survive as an independent intellectual discipline, religion will endure for a long time to come and will not be replaced by scientific materialism.

Scientific humanism Wilson coined the phrase scientific humanism as "the only worldview compatible with science's growing knowledge of the real world and the laws of nature".[17] Wilson argues that it is best suited to improve the human condition. In 2003 he was one of the signers of the Humanist Manifesto.

God and religion On the question of God, Wilson has described his position as provisional deism.[18] He has explained his faith as a trajectory away from traditional beliefs: "I drifted away from the church, not definitively agnostic or atheistic, just Baptist & Christian no more." Wilson argues that the belief in God and rituals of religion are products of evolution.[19] He argues that they should not be rejected or dismissed, but further investigated by science to better understand their significance to human nature. In his book The Creation, Wilson suggests that scientists ought to "offer the hand of friendship" to religious leaders and build an alliance with them, stating that "Science and religion are two of the most potent forces on Earth and they should come together to save the creation."[20] Wilson makes a similar suggestion, and appeal to the religious community, on the lecture circuit. An article on his September 17, 2009 lecture at Midland College, Texas, reports, "he said the appeal received a 'massive reply' and a covenant has been written. 'I think that partnership will work to a substantial degree as time goes on,' Wilson said."[21] Wilson appears in the documentary Behold The Earth, which inquires into America's "divorce from nature" and the relationship between science and religion.

Ecology When discussing the reinvigoration of his original fields of study since the 1960s, Wilson has said that if he could start his life over he would work in microbial ecology. He studied the mass extinctions of the 20th century and their relationship to modern society, arguing strongly for an ecological approach: Now when you cut a forest, an ancient forest in particular, you are not just removing a lot of big trees and a few birds fluttering around in the canopy. You are drastically imperiling a vast array of species within a few square miles of you. The number of these species may go to tens of thousands. ... Many of them are still unknown to science, and science has not yet discovered the key role undoubtedly played in the maintenance of that ecosystem, as in the case of fungi, microorganisms, and many of the insects.[22] His understanding of the scale of the extinction crisis has led him to advocate a number of strategies for forest protection, including the Forests Now Declaration, which calls for new markets-based mechanisms to protect tropical forests.

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Criticism of human sociobiology Wilson experienced significant criticism for his sociobiological views from several different communities. The scientific response included several of Wilson's colleagues at Harvard, such as Richard Lewontin and Stephen Jay Gould, who were strongly opposed to his ideas regarding sociobiology. Marshall Sahlins's work The Use and Abuse of Biology was a direct criticism of Wilson's theories. Politically, Wilson's sociobiological ideas have offended some Marxists who favored the idea that human behavior was culturally based. Sociobiology re-ignited the nature-versus-nurture debate, and Wilson's scientific perspective on human nature led to public debate. He was accused of "racism, misogyny, and eugenics." In one incident, his lecture was attacked by the International Committee Against Racism, a front group of the Progressive Labor Party, where one member poured a pitcher of water on Wilson's head and chanted "Wilson, you're all wet" at an AAAS conference in November 1978. Wilson later spoke of the incident as a source of pride: "I believe...I was the only scientist in modern times to be physically attacked for an idea." “I believe Gould was a charlatan,” Wilson told The Atlantic. “I believe that he was ... seeking reputation and credibility as a scientist and writer, and he did it consistently by distorting what other scientists were saying and devising arguments based upon that distortion.” Religious objections included those of Paul E. Rothrock, who said: "... sociobiology has the potential of becoming a religion of scientific materialism." [23]

Awards and honors Wilson's scientific and conservation honors include: • Member, National Academy of Sciences, 1969 • U.S. National Medal of Science, 1976 • Pulitzer Prize for On Human Nature, 1979 • Tyler Prize for Environmental Achievement, 1984 • ECI Prize, International Ecology Institute, terrestrial ecology, 1987 • Academy of Achievement Golden Plate Award, 1988 • Crafoord Prize, 1990, a prize awarded by the Royal Swedish Academy of Sciences in certain sciences not covered by the Nobel Prize, and therefore considered the highest award given in the field of ecology

Wilson at a "fireside chat" during which he received the Addison Emery Verrill Medal in 2007

• Pulitzer Prize for The Ants (with Bert Hölldobler), 1991 • International Prize for Biology, 1993 • Carl Sagan Award for Public Understanding of Science, 1994 • Time Magazine's 25 Most Influential People in America, 1995 • Benjamin Franklin Medal for Distinguished Achievement in the Sciences of the American Philosophical Society, 1998. • American Humanist Association's 1999 Humanist of the Year • Lewis Thomas Prize for Writing about Science, 2000 • Nierenberg Prize, 2001 • Distinguished Eagle Scout Award 2004 • Dauphin Island Sea Lab christened its newest research vessel the R/V E.O. Wilson in 2005.

Dr. E.O. Wilson addresses the audience at the dedication of the E.O. Wilson Biophilia Center at Nokuse Plantation in Walton County, Florida.

• The Linnean Tercentenary Silver Medal, 2006 • Addison Emery Verrill Medal from the Peabody Museum of Natural History, 2007

E. O. Wilson • TED Prize 2007 [24] given yearly to honor a maximum of three individuals who have shown that they can, in some way, positively impact life on this planet. • XIX Premi Internacional Catalunya 2007 [25] • Member of the World Knowledge Dialogue [26] Honorary Board, and Scientist in Residence for the 2008 symposium organized in Crans-Montana (Switzerland). • Distinguished Lecturer, University of Iowa, 2008–2009 • E.O. Wilson Biophilia Center [27] on Nokuse Plantation in Walton County, Florida 2009 Video [28] • Explorers Club Medal, 2009 • BBVA Frontiers of Knowledge Award in the Ecology and Conservation Biology Category [29], 2010 • Thomas Jefferson Medal in Architecture, 2010 • 2010 Heartland Prize for fiction for his first novel Anthill: A Novel[30] • EarthSky Science Communicator of the Year,2010

Main works • The Theory of Island Biogeography, 1967, Princeton University Press (2001 reprint), ISBN 0-691-08836-5, with Robert H. MacArthur • The Insect Societies [31], 1971, Harvard University Press, ISBN 0-674-45490-1 • Sociobiology: The New Synthesis 1975, Harvard University Press, (Twenty-fifth Anniversary Edition, 2000 ISBN 0-674-00089-7) • On Human Nature, 1979, Harvard University Press, ISBN 0-674-01638-6 • Genes, Mind and Culture: The coevolutionary process, 1981, Harvard University Press, ISBN 0-674-34475-8 • Promethean fire: reflections on the origin of mind, 1983, Harvard University Press, ISBN 0-674-71445-8 • Biophilia [32], 1984, Harvard University Press, ISBN 0-674-07441-6 • Success and Dominance in Ecosystems: The Case of the Social Insects, 1990, Inter-Research, ISSN 0932-2205 • The Ants, 1990, Harvard University Press, ISBN 0-674-04075-9, Winner of the Pulitzer Prize, with Bert Hölldobler • The Diversity of Life [33], 1992, Harvard University Press, ISBN 0-674-21298-3, The Diversity of Life: Special Edition [34], ISBN 0-674-21299-1 • The Biophilia Hypothesis, 1993, Shearwater Books, ISBN 1-55963-148-1, with Stephen R. Kellert • Journey to the Ants: A Story of Scientific Exploration, 1994, Harvard University Press, ISBN 0-674-48525-4, with Bert Hölldobler • Naturalist, 1994, Shearwater Books, ISBN 1-55963-288-7 • In Search of Nature, 1996, Shearwater Books, ISBN 1-55963-215-1, with Laura Simonds Southworth • Consilience: The Unity of Knowledge, 1998, Knopf, ISBN 0-679-45077-7 • The Future of Life, 2002, Knopf, ISBN 0-679-45078-5 • Pheidole in the New World: A Dominant, Hyperdiverse Ant Genus [35], 2003, Harvard University Press, ISBN 0-674-00293-8 • From So Simple a Beginning: Darwin's Four Great Books. 2005, W. W. Norton. • The Creation: An Appeal to Save Life on Earth, September 2006, W. W. Norton & Company, Inc. ISBN 978-0-393-06217-5 • Nature Revealed: Selected Writings 1949-2006, ISBN 0-8018-8329-6 • The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies, 2009, W.W. Norton & Company, Inc. ISBN 978-0-393-06704-0, with Bert Hölldobler • Anthill: A Novel, April 2010, W. W. Norton & Company, Inc. ISBN 978-0-393-07119-1 • Kingdom of Ants: Jose Celestino Mutis and the Dawn of Natural History in the New World, 2010, Johns Hopkins University Press, Baltimore, with José María Gómez Durán

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E. O. Wilson • The Leafcutter Ants: Civilization by Instinct, 2011, W.W. Norton & Company, Inc. ISBN 978-0-393-33868-3, with Bert Hölldobler • The Social Conquest of Earth, 2012, Liveright Publishing Corporation, New York, ISBN 978-0-87140-413-8 • Letters to a Young Scientist, 2013, WW Norton & Company, ISBN 0871407000

Edited works • From So Simple a Beginning: Darwin's Four Great Books, edited with introductions by Edward O. Wilson (2010 W.W. Norton)

Footnotes [1] http:/ / search. proquest. com/ docview/ 301948222/ [2] http:/ / www. guardian. co. uk/ science/ 2012/ jun/ 24/ battle-of-the-professors [3] "E.O. Wilson Profile" (http:/ / web. archive. org/ web/ 20041014233232/ http:/ / www. metrokc. gov/ dnrp/ swd/ naturalconnections/ edward_wilson_bio. htm) - Comprehensive list of Degrees, Awards and Positions [4] Archive.org; E. O. Wilson biography (http:/ / web. archive. org/ web/ 20101208080544/ http:/ / alabamaliterarymap. org/ author. cfm?AuthorID=16) [5] http:/ / www. nytimes. com/ best-sellers-books/ 2012-05-06/ hardcover-nonfiction/ list. html [6] http:/ / www. nytimes. com/ best-sellers-books/ 2013-05-05/ hardcover-nonfiction/ list. html [7] Edward O. Wilson – Naturalist, Island Press; (April 24, 2006), ISBN 1-59726-088-6 [8] first-hand account, Smithsonian Institution talk, April 22, 2010 [9] Edward O. Wilson, Foreword of Everybody's Story: Wising Up to the Epic of Evolution By Loyal D. Rue, SUNY Press, 1999, page ix and x,ISBN 0-7914-4392-2, [10] Edward O. Wilson, Consilience 1998 (http:/ / www. thegreatstory. org/ ClassicQuotes. pdf) [11] Brian Swimme interview (http:/ / www. earthlight. org/ interview26. html) [12] Kaufman, Gordon. The Epic of Evolution as a Framework for Human Orientation, 1997 [13] Edward O. Wilson: On Human Nature - Summary by Michael McGoodwin, prepared 1991 (http:/ / mcgoodwin. net/ pages/ otherbooks/ eow_humannature. html) [14] E. O. Wilson, Consilience: The Unity of Knowledge, New York, Knopf, 1998, pp. 127-128. [15] Wolfe, Tom (1996). Sorry, But Your Soul Just Died. Vol. 158, Issue 13, pp.210ff. Forbes [16] Richard Conniff "Discover Interview: E.O. Wilson" (http:/ / discovermagazine. com/ 2006/ jun/ e-o-wilson) June 25, 2006. [17] in Harvard Magazine December 2005 p 33. [18] The Creation [19] Human Nature [20] Naturalist E.O. Wilson is optimistic (http:/ / www. news. harvard. edu/ gazette/ 2006/ 06. 15/ 03-biodiversity. html) Harvard Gazette June 15, 2006 [21] Scientist says there is hope to save planet (http:/ / www. mywesttexas. com/ articles/ 2009/ 09/ 18/ news/ top_stories/ doc4ab31071978e4063758641. txt) mywesttexas.com September 18, 2009 [22] Wilson, E. O. (1998 April 28, 1998). Slide show (http:/ / www. saveamericasforests. org/ wilson/ second. htm), page 2. Delivered at Washington, DC. Website of Save America's Forests. Accessed 2008-11-13. [23] Mythology of Scientific Materialism. (http:/ / www. asa3. org/ asa/ PSCF/ 1987/ PSCF6-87Rothrock. html) Paul E. Rothrock and Mary Ellen Rothrock [24] http:/ / www. ted. com/ tedprize/ winners2007. cfm [25] http:/ / www20. gencat. cat/ portal/ site/ msi-dgac/ menuitem. cfb4d0ec869dcac3934fec60b0c0e1a0/ ?vgnextoid=441e7109261a9210VgnVCM1000008d0c1e0aRCRD& vgnextchannel=441e7109261a9210VgnVCM1000008d0c1e0aRCRD& vgnextfmt=default [26] http:/ / www. wkdialogue. ch [27] http:/ / www. eowilsoncenter. org [28] http:/ / www. vimeo. com/ 8678440 [29] http:/ / www. fbbva. es/ TLFU/ tlfu/ ing/ microsites/ premios/ fronteras/ index. jsp [30] E. O. WILSON AND REBECCA SKLOOT: 2010 CHICAGO TRIBUNE HEARTLAND PRIZES (http:/ / www. chicagohumanities. org/ Genres/ Literature/ 2010-Chicago-Tribune-Heartland-Prize-Winners. aspx#) Recorded on November 13, 2010. [31] http:/ / www. hup. harvard. edu/ catalog. php?isbn=9780674454903 [32] http:/ / www. hup. harvard. edu/ catalog. php?isbn=9780674074422 [33] http:/ / www. hup. harvard. edu/ catalog. php?isbn=9780674058170 [34] http:/ / www. hup. harvard. edu/ catalog/ WILDIS. html

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E. O. Wilson [35] http:/ / www. hup. harvard. edu/ catalog. php?isbn=9780674002937

External links • E.O. Wilson Foundation (http://eowilsonfoundation.org/) • Review of The Social Conquest of Earth by Richard Dawkins (http://www.prospectmagazine.co.uk/magazine/ edward-wilson-social-conquest-earth-evolutionary-errors-origin-species/#.UlLItRBELIg)

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W. D. Hamilton

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W. D. Hamilton W. D. Hamilton W. D. Hamilton, 1996 Born

William Donald Hamilton 1 August 1936 Cairo, Egypt

Died

7 March 2000 (aged 63) Fitzrovia, London, United Kingdom

Nationality

British

Fields

Evolutionary biology

Alma mater

University College London London School of Economics St. John's College, Cambridge

Academic advisors John Hajnal Cedric Smith Doctoral students

Laurence Hurst Olivia Judson

Known for

Kin selection, Hamilton's rule

Influences

Ronald Fisher

Influenced

Richard Dawkins

William Donald "Bill" Hamilton, FRS (1 August 1936 – 7 March 2000) was an English evolutionary biologist, widely recognised as one of the greatest evolutionary theorists of the 20th century.[1][2] Hamilton became famous through his theoretical work expounding a rigorous genetic basis for the existence of kin selection and altruism, an insight that was a key part of the development of a gene-centric view of evolution. He is considered one of the forerunners of sociobiology, as popularized by E. O. Wilson. Hamilton also published important work on sex ratios and the evolution of sex. From 1984 to his death in 2000, he was a Royal Society Research Professor at Oxford University.

Early life Hamilton was born in 1936 in Cairo, Egypt, the second of seven children. His father A. M. Hamilton was a New Zealand-born engineer. His mother B. M. Hamilton was a medical doctor, also from New Zealand. The Hamilton family settled in Kent. During the Second World War, the young Hamilton was evacuated to Edinburgh. He had an interest in natural history from an early age and would spend his spare time collecting butterflies and other insects. In 1946 he discovered E.B. Ford's New Naturalist book Butterflies, which introduced him to the principles of evolution by natural selection, genetics and population genetics. He was educated at Tonbridge School, where he was in Smythe House. As a 12-year old he was seriously injured while playing with explosives his father had. These were left over from his making hand grenades for the Home Guard during World War II; the accident might have killed him if his mother had not been a doctor. The boy had to have a thoracotomy in King's College Hospital to save his life, but fingers on his right hand had to be amputated and he was left with scarring on his body. He needed six months to recover. Hamilton stayed on an extra term at Tonbridge to complete the Cambridge entrance examinations, and then travelled in France. He completed two years of national service. As an undergraduate at St. John's College, he was uninspired

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by the "many biologists [who] hardly seemed to believe in evolution". He was intrigued by Ronald Fisher's book The Genetical Theory of Natural Selection; but Fisher lacked standing at Cambridge as he was viewed as only a statistician. Hamilton was excited by Fisher's chapters on eugenics. In earlier chapters, Fisher provided a mathematical basis for the genetics of evolution. Working through the stodgy prose, Hamilton later blamed Fisher's book for his getting only a 2:1 degree.

Hamilton's rule Hamilton enrolled in an MSc course in demography at the London School of Economics (LSE), under Norman Carrier, who helped secure various grants for his studies. Later, when his work became more mathematical and genetical, he had his supervision transferred to John Hajnal of the LSE and Cedric Smith of University College London (UCL). Both Fisher and J. B. S. Haldane had seen a problem in how organisms could increase the fitness of their own genes by aiding their close relatives, but not recognised its significance or properly formulated it. Hamilton worked through several examples, and eventually realised that the number that kept falling out of his calculations was Sewall Wright's coefficient of relationship. This became Hamilton's rule: in each behaviour-evoking situation, the individual assesses his neighbour's fitness against his own according to the coefficients of relationship appropriate to the situation. Algebraically, the rule posits that a costly action should be performed if: Where C is the cost in fitness to the actor, r the genetic relatedness between the actor and the recipient, and B is the fitness benefit to the recipient. Fitness costs and benefits are measured in fecundity. His two 1964 papers entitled The Genetical Evolution of Social Behavior are now widely referenced. The proof and discussion of its consequences, however, involved detailed mathematics, and two reviewers passed over the paper. The third, John Maynard Smith, did not completely understand it either, but recognised its significance. Having his work passed over later led to friction between Hamilton and Maynard Smith, as Hamilton thought Smith had held his work back to claim credit for the idea (during the review period Maynard Smith published a paper that referred briefly to similar ideas). The Hamilton paper was printed in the Journal of Theoretical Biology and, when first published, was largely ignored. Recognition of its significance gradually increased to the point that it is now routinely cited in biology books. Much of the discussion relates to the evolution of eusociality in insects of the order Hymenoptera (ants, bees and wasps) based on their unusual haplodiploid sex-determination system. This system means that females are more closely related to their sisters than to their own (potential) offspring. Thus, Hamilton reasoned, a "costly action" would be better spent in helping to raise their sisters, rather than reproducing themselves.

Spiteful behaviour In his 1970 paper Selfish and Spiteful Behaviour in an Evolutionary Model Hamilton considers the question of whether harm inflicted upon an organism must inevitably be a byproduct of adaptations for survival. What of possible cases where an organism is deliberately harming others without apparent benefit to the self? Such behaviour Hamilton calls spiteful. It can be explained as the increase in the chance of an organism's genetic alleles to be passed to the next generations by harming those that are less closely related than relationship by chance. Spite, however, is unlikely ever to be elaborated into any complex forms of adaptation. Targets of aggression are likely to act in revenge, and the majority of pairs of individuals (assuming a panmictic species) exhibit a roughly average level of genetic relatedness, making the selection of targets of spite problematic.

W. D. Hamilton

Extraordinary sex ratios Between 1964 and 1977 Hamilton was a lecturer at Imperial College London. Whilst there he published a paper in Science on "extraordinary sex ratios". Fisher (1930) had proposed a model as to why "ordinary" sex ratios were nearly always 1:1 (but see Edwards 1998), and likewise extraordinary sex ratios, particularly in wasps, needed explanations. Hamilton had been introduced to the idea and formulated its solution in 1960 when he had been assigned to help Fisher's pupil A.W.F. Edwards test the Fisherian sex ratio hypothesis. Hamilton combined his extensive knowledge of natural history with deep insight into the problem, opening up a whole new area of research. The paper was also notable for introducing the concept of the "unbeatable strategy", which John Maynard Smith and George R. Price were to develop into the evolutionarily stable strategy (ESS), a concept in game theory not limited to evolutionary biology. Price had originally come to Hamilton after deriving the Price equation, and thus rederiving Hamilton's rule. Maynard Smith later peer reviewed one of Price's papers, and drew inspiration from it. The paper was not published but Maynard Smith offered to make Price a co-author of his ESS paper, which helped to improve relations between the men. Price committed suicide in 1975, and Hamilton and Maynard Smith were among the few present at the funeral. Hamilton was regarded as a poor lecturer. This shortcoming would not affect the popularity of his work, however, as it was popularised by Richard Dawkins in Dawkins' 1976 book The Selfish Gene. In 1966 he married Christine Friess and they were to have three daughters, Helen, Ruth and Rowena. 26 years later they amicably separated. Hamilton was a visiting professor at Harvard University and later spent nine months with the Royal Society's and the Royal Geographical Society's Xavantina-Cachimbo Expedition as a visiting professor at the University of São Paulo. From 1978 Hamilton was Professor of Evolutionary Biology at the University of Michigan. Simultaneously, he was elected a Foreign Honorary Member of American Academy of Arts and Sciences. His arrival sparked protests and sit-ins from students who did not like his association with sociobiology. There he worked with the political scientist Robert Axelrod on the prisoner's dilemma, and was a member of the BACH group with original members Arthur Burks, Robert Axelrod, Michael Cohen and John Holland.

Chasing the Red Queen Hamilton was an early proponent of the Red Queen theory of the evolution of sex,[3] first proposed by Leigh Van Valen. This was named for a character in Lewis Carroll's Through the Looking-Glass, who is continuously running but never actually travels any distance: "Well, in our country," said Alice, still panting a little, "you'd generally get to somewhere else—if you ran very fast for a long time, as we've been doing." "A slow sort of country!" said the Queen. "Now, here, you see, it takes all the running you can do, to keep in the same place. If you want to get somewhere else, you must run at least twice as fast as that!" (Carroll, pp. 46) [4]

This theory hypothesizes that sex evolved because new and unfamiliar combinations of genes could be presented to parasites, preventing the parasite from preying on that organism: species with sex were able to continuously "run away" from their parasites. Likewise, parasites were able to evolve mechanisms to get around the organism's new set of genes, thus perpetuating an endless race.

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Return to Britain In 1980 he was elected a Fellow of the Royal Society, and in 1984 he was invited by Richard Southwood to be the Royal Society Research Professor in the Department of Zoology at Oxford, and a Fellow of New College, where he remained until his death. From 1994 Hamilton found companionship with Maria Luisa Bozzi, an Italian science journalist and author. His collected papers, entitled Narrow Roads of Gene Land, began to be published in 1996. The first volume was entitled Evolution of Social Behaviour.

Social evolution The field of social evolution, of which Hamilton's rule has central importance, is broadly defined as being the study of the evolution of social behaviours, i.e. those that impact on the fitness of individuals other than the actor. Social behaviours can be categorized according to the fitness consequences they entail for the actor and recipient. A behaviour that increases the direct fitness of the actor is mutually beneficial if the recipient also benefits, and selfish if the recipient suffers a loss. A behaviour that reduces the fitness of the actor is altruistic if the recipient benefits, and spiteful if the recipient suffers a loss. This classification was first proposed by Hamilton in 1964.[citation needed] Through his collaboration with Hugh N. Comins and Bob May on evolutionarily stable dispersal strategies, Hamilton acquired an Erdős number of 5.[5] Hamilton also proposed the coevolution theory of autumn leaf color as an example of evolutionary signalling theory.

Expedition to the Congo During the 1990s Hamilton became increasingly interested in the controversial argument that the origin of HIV lay in oral polio vaccines trials conducted by Hilary Koprowski in Africa during the 1950s. Letters by Hamilton on the topic to the major peer-reviewed journals were rejected. To look for indirect evidence of the OPV hypothesis by assessing natural levels of simian immunodeficiency virus, in primates, in early 2000 he and two others ventured on a field trip to the war-torn Democratic Republic of the Congo.

Death He returned to London from Africa on 29 Jan 2000. He was admitted to University College Hospital London on 30 Jan 2000. He was transferred to Middlesex Hospital London on 5 Feb 2000 and died there on 7 Mar 2000. An inquest was held on 10 May 2000 at Westminster Coroner's Court to inquire into rumours about the cause of his death. The coroner concluded that his death was due to "Multi-organ failure due to upper gastrointestinal haemorrhage due to a duodenal diverticulum and arterial bleed through a mucosal ulcer". ECU's investigation established that he contracted malaria during his final African expedition. However, the pathologist had suggested the possibility that the ulceration and consequent haemorrhage had resulted from a pill (which might have been taken because of malarial symptoms) lodging in the diverticulum; but, even if this suggestion were correct, the link between malaria and the observed causes of death would be entirely indirect. A secular memorial service (he was an atheist) was held at the Chapel of New College, Oxford on Saturday 1 July 2000, organised by Richard Dawkins. He was buried near Wytham Woods. He, however, had written an essay on My intended burial and why in which he wrote:[6] I will leave a sum in my last will for my body to be carried to Brazil and to these forests. It will be laid out in a manner secure against the possums and the vultures just as we make our chickens secure; and this great Coprophanaeus beetle will bury me. They will enter, will bury, will live on my flesh; and in the shape of their children and mine, I will escape death. No worm for me nor sordid fly, I will buzz in the dusk like a huge bumble bee. I will be many, buzz even as a swarm of motorbikes, be borne, body by flying body out into the Brazilian wilderness beneath the stars, lofted under those beautiful and un-fused elytra which we will all hold over our backs. So finally I too will shine like a violet ground beetle under a stone.

“

”

W. D. Hamilton

The second volume of his collected papers, Evolution of Sex, was published in 2002, and the third and final volume, Last Words, in 2005.

Awards • • • • • • • • • •

1978 Foreign Honorary Member of American Academy of Arts and Sciences 1980 Fellow of the Royal Society of London 1982 Newcomb Cleveland Prize of the American Association for the Advancement of Science 1988 Darwin Medal of the Royal Society of London 1989 Scientific Medal of the Linnean Society 1991 Frink Medal of Zoological Society of London 1992/3 Wander Prize of the University of Bern 1993 Crafoord Prize of the Royal Swedish Academy of Sciences 1993 Kyoto Prize of the Inamori Foundation 1995 Fyssen Prize of the Fyssen Foundation

Biographies • Alan Grafen has written a biographical memoir for the Royal Society. • A biographical book has also been published : Segerstråle, U. 2013. Nature's oracle: the life and work of W. D. Hamilton. Oxford University Press [7].

Works Collected papers Hamilton started to publish his collected papers starting in 1996, along the lines of Fisher's collected papers, with short essays giving each paper context. He died after the preparation of the second volume, so the essays for the third volume come from his coauthors. • Hamilton W.D. (1996) Narrow Roads of Gene Land vol. 1: Evolution of Social Behaviour Oxford University Press,Oxford. ISBN 0-7167-4530-5 • Hamilton W.D. (2002) Narrow Roads of Gene Land vol. 2: Evolution of Sex Oxford University Press,Oxford. ISBN 0-19-850336-9 • Hamilton W.D. (2005) Narrow roads of Gene Land, vol. 3: Last Words (with essays by coauthors, ed. M. Ridley). Oxford University Press, Oxford. ISBN 0-19-856690-5

Significant papers • Hamilton, W. (1964). "The genetical evolution of social behaviour. I". Journal of Theoretical Biology 7 (1): 1–16. doi:10.1016/0022-5193(64)90038-4 [8]. PMID 5875341 [9]. • Hamilton, W. (1964). "The genetical evolution of social behaviour. II". Journal of Theoretical Biology 7 (1): 17–52. doi:10.1016/0022-5193(64)90039-6 [10]. PMID 5875340 [11]. • Hamilton, W. (1966). "The moulding of senescence by natural selection". Journal of Theoretical Biology 12 (1): 12–45. doi:10.1016/0022-5193(66)90184-6 [12]. PMID 6015424 [13]. • Hamilton, W. (1967). "Extraordinary sex ratios. A sex-ratio theory for sex linkage and inbreeding has new implications in cytogenetics and entomology". Science 156 (774): 477–488. doi:10.1126/science.156.3774.477 [14] . PMID 6021675 [15]. • Hamilton, W. (1971). "Geometry for the selfish herd". Journal of Theoretical Biology 31 (2): 295–311. doi:10.1016/0022-5193(71)90189-5 [16]. PMID 5104951 [17].

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W. D. Hamilton • Hamilton W.D. (1975). Innate social aptitudes of man: an approach from evolutionary genetics. [18] in R. Fox (ed.), Biosocial Anthropology, Malaby Press, London, 133-53. • Axelrod, R.; Hamilton, W. (1981). "The evolution of cooperation". Science 211 (4489): 1390–1396. doi:10.1126/science.7466396 [19]. PMID 7466396 [20]. with Robert Axelrod • Hamilton, W.; Zuk, M. (1982). "Heritable true fitness and bright birds: A role for parasites?". Science 218 (4570): 384–387. doi:10.1126/science.7123238 [21]. PMID 7123238 [22].

Notes [1] Obituary by Richard Dawkins - The Independent - 10 March 2000 (http:/ / www. edge. org/ 3rd_culture/ hamilton/ hamilton_index. html) [2] BBC Radio 4 - Great Lives - 2 Feb 2010 (http:/ / www. bbc. co. uk/ iplayer/ console/ b00qc2hn) [3] The Red Queen Hypothesis (http:/ / www. indiana. edu/ ~curtweb/ Research/ Red_Queen hyp. html) at Indiana University. Quote: "W.D. Hamilton and John Jaenike were among the earliest pioneers of the idea." [4] http:/ / etext. lib. virginia. edu/ etcbin/ toccer-new2?id=CarGlas. sgm& images=images/ modeng& data=/ texts/ english/ modeng/ parsed& tag=public& part=2& division=div1 [5] His path to Erdős: William D. Hamilton coauthored with Robert M. May, Robert M. May coauthored with Yoh Iwasa, Yoh Iwasa coauthored with Eugene Seneta, Eugene Seneta coauthored with Janos Galambos, Janos Galambos coauthored with Paul Erdős. [6] Hamilton, W.D. (2000) My intended burial and why, Ethology Ecology and Evolution 12 111-122 PDF (http:/ / ejour-fup. cilea. it/ index. php/ eee/ article/ viewFile/ 859/ 805) [7] http:/ / global. oup. com/ academic/ product/ natures-oracle-9780198607274?q=Nature%27s%20oracle& lang=en& cc=us [8] http:/ / dx. doi. org/ 10. 1016%2F0022-5193%2864%2990038-4 [9] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 5875341 [10] http:/ / dx. doi. org/ 10. 1016%2F0022-5193%2864%2990039-6 [11] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 5875340 [12] http:/ / dx. doi. org/ 10. 1016%2F0022-5193%2866%2990184-6 [13] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 6015424 [14] http:/ / dx. doi. org/ 10. 1126%2Fscience. 156. 3774. 477 [15] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 6021675 [16] http:/ / dx. doi. org/ 10. 1016%2F0022-5193%2871%2990189-5 [17] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 5104951 [18] http:/ / www. majorityrights. com/ docs/ Hamilton75. pdf [19] http:/ / dx. doi. org/ 10. 1126%2Fscience. 7466396 [20] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 7466396 [21] http:/ / dx. doi. org/ 10. 1126%2Fscience. 7123238 [22] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 7123238

References • Edwards A.W.F. (1998) Notes and Comments. Edwards, A.  W.  F. (1998). "Natural Selection and the Sex Ratio: Fisher's Sources". The American Naturalist 151 (6): 564–569. doi: 10.1086/286141 (http://dx.doi.org/10. 1086/286141). PMID  18811377 (http://www.ncbi.nlm.nih.gov/pubmed/18811377). • Fisher R.A. (1930). The Genetical Theory of Natural Selection. Clarendon Press, Oxford. • Ford E.B. (1945) New Naturalist 1: Butterflies. Collins: London. • Maynard Smith J. and G.R. Price (1973) The logic of animal conflict. Nature 146: 15—18. • Dawkins R. (1989) The Selfish Gene, 2nd ed. Oxford University Press. • Madsen E.A., Tunney R. Fieldman, G. Plotkin H.C., Dunbar R.I.M., Richardson J.M. & McFarland D. (2006) Kinship and altruism: a cross-cultural experimental study. British Journal of Psychology: http://www. ingentaconnect.com/content/bpsoc/bjp/pre-prints/218320

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External links • Obituaries and reminiscences (http://www.unifr.ch/biol/ecology/hamilton/hamilton.html) • Royal Society citation (http://www.royalsoc.ac.uk/DServe/dserve.exe?dsqIni=Dserve.ini& dsqApp=Archive&dsqCmd=Show.tcl&dsqSearch=RefNo=='EC/1980/12'&dsqDb=Catalog) • Truth and Science: Bill Hamilton's legacy (http://www.uow.edu.au/~/bmartin/dissent/documents/AIDS/ Bozzi03.pdf) • Centro Itinerante de Educação Ambiental e Científica Bill Hamilton (The Bill Hamilton Itinerant Centre for Environmental and Scientific Education) (in Portuguese) (http://www.mamiraua.org.br/4-2-1.html) • Non-mathematical excerpts from Hamilton 1964 (http://www.serpentfd.org/a/hamilton1964.html) • "If you have a simple idea, state it simply" a 1996 interview with Hamilton (http://www.froes.dds.nl/ HAMILTON.htm) • London Review of Books book review (http://www.lrb.co.uk/v25/n03/andrew-berry/ reasons-for-being-nice-and-having-sex) • W.D. Hamilton's work in game theory (http://www.iiasa.ac.at/Publications/Documents/IR-02-019.pdf)

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Robert Trivers

126

Robert Trivers Robert Trivers Born

February 19, 1943 Washington DC

Nationality

American

Fields

Biology

Institutions

Rutgers University

Known for

Research on sociobiology, evolutionary psychology, Darwinian social science, and behavioral ecology

Influences

Charles Darwin, W. D. Hamilton

Influenced

Steven Pinker, E. O. Wilson, Richard Dawkins

Notable awards

2007 Crafoord Prize in Biosciences

Spouse

Lorna Staples (1974-1988); Debra Dixon (1997-2004)

Children

three daughters with Staples: Natasha and Natalia (twins), Alelia, one son with Staples: Jonathan; one daughter with Dixon: Aubrey Website http:/ / roberttrivers. com

Robert L. Trivers (/ˈtrɪvərz/; born February 19, 1943) is an American evolutionary biologist and sociobiologist, who is a Professor of Anthropology and Biological Sciences at Rutgers University. Trivers proposed the theories of reciprocal altruism (1971), parental investment (1972), facultative sex ratio determination (1973), and parent-offspring conflict (1974). He has also contributed by explaining self-deception as an adaptive evolutionary strategy (first described in 1976) and discussing intragenomic conflict.

Education Trivers graduated from Phillips Academy in 1961 and intended to study mathematics at Harvard University. However, he ended studying American history, preparing to become a lawyer. He graduated in 1965. He took a psychology class after suffering a breakdown, and was very unimpressed with the state of psychology.[citation needed] He was prevented from getting into Yale law school by his breakdown, and so took a job writing social science textbooks for children that were never published.[citation needed] According to The Guardian's Andrew Brown, the breakdown occurred because Trivers stayed up "all night, night after night" reading Ludwig Wittgenstein. This landed him in a hospital where he was "treated with the first generation of effective anti-psychotic drugs" and, as part of his therapy, he took art classes. He then got a job illustrating and later writing a "series of textbooks for high schools".[citation needed] While recovering, he took courses in art, and was hired to illustrate, and then to write, a series of textbooks for high schools. Despite his history degree, it was obvious to his supervisors that he knew little about human biology, so he was given the animals to write about, and started to learn modern Darwinian biology.[1] This exposure to evolutionary theory led him to do graduate work with Ernst Mayr at Harvard from 1968 to 1972. He earned his Ph.D. in biology on June 15, 1972, also from Harvard. The second half of his first major paper, "The Evolution of Reciprocal Altruism" was published in 1971.[2]

Robert Trivers

Career Trivers was on faculty at Harvard from 1973 to 1978, and then moved to the University of California, Santa Cruz where he was a faculty member 1978 to 1994. He is currently a Rutgers University notable faculty member. In the 2008–09 academic year, he was a Fellow at the Institute for Advanced Study in Berlin (Wissenschaftskolleg zu Berlin). Trivers was recently awarded the 2007 Crafoord Prize in Biosciences for "his fundamental analysis of social evolution, conflict and cooperation". Trivers wrote the original foreword to Richard Dawkins' book The Selfish Gene in which Trivers first proposed his adaptive theory of self-deception. Trivers met Huey P. Newton, Chairman of the Black Panther Party, in 1978 when Newton applied while in prison to do a reading course with Trivers as part of a graduate degree in History of Consciousness at UC Santa Cruz. Trivers and Newton became close friends: Newton was godfather to one of Trivers' daughters.[3] Trivers joined the Black Panther Party in 1979. Trivers and Newton published an analysis of the role of self-deception by the flight crew in the crash of Air Florida Flight 90.[4]

Influence Trivers is arguably one of the most influential evolutionary theorists alive today. Steven Pinker considers Trivers to be "one of the great thinkers in the history of Western thought", who has:[5] inspired an astonishing amount of research and commentary in psychology and biology—the fields of sociobiology, evolutionary psychology, Darwinian social science, and behavioral ecology are in large part attempt to test and flesh out Trivers' ideas. It is no coincidence that E. O. Wilson's Sociobiology and Richard Dawkins' The Selfish Gene were published in 1975 and 1976 respectively, just a few years after Trivers' seminal papers. Both bestselling authors openly acknowledged that they were popularizing Trivers' ideas and the research they spawned. Likewise for the much-talked-about books on evolutionary psychology in the 1990s—The Adapted Mind, The Red Queen, Born to Rebel, The Origins of Virtue, The Moral Animal, and my own How the Mind Works. Each of these books is based in large part on Trivers' ideas and the explosion of research they inspired (involving dozens of animal species, mathematical and computer modeling, and human social and cognitive psychology).

Bibliography Significant papers • Trivers, R. L. (1971). "The Evolution of Reciprocal Altruism". The Quarterly Review of Biology 46 (1): 35–57. doi:10.1086/406755 [6]. JSTOR 2822435 [7]. • Trivers, R. L. (1972) Parental investment and sexual selection. [8] In B. Campbell (Ed.) Sexual selection and the descent of man, 1871-1971 (pp 136–179). Chicago, Aldine. • Trivers, RL; Willard, DE (1973). "Natural selection of parental ability to vary the sex ratio of offspring". Science 179 (4068): 90–2. Bibcode:1973Sci...179...90T [9]. doi:10.1126/science.179.4068.90 [10]. JSTOR 1734960 [11]. PMID 4682135 [12]. • Trivers, R. L. (1974). "Parent-Offspring Conflict". American Zoologist 14 (1): 249–264. doi:10.1093/icb/14.1.249 [13] . • Trivers, R. L.; Hare, H. (1976). "Haploidploidy and the evolution of the social insect". Science 191 (4224): 249–63. Bibcode:1976Sci...191..249T [14]. doi:10.1126/science.1108197 [15]. PMID 1108197 [16]. • Trivers, R. L. (1991) Deceit and self-deception: The relationship between communication and consciousness. In: M. Robinson and L. Tiger (eds.) Man and Beast Revisited, Smithsonian, Washington, DC, pp. 175–191.

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Books • Trivers, R. L. (1985) Social Evolution. Benjamin/Cummings, Menlo Park, CA. • Trivers, R. L. (2002) Natural Selection and Social Theory: Selected Papers of Robert L. Trivers. (Evolution and Cognition Series) Oxford University Press, Oxford. ISBN 0-19-513062-6 • Burt, A. & Trivers, R. L. (2006) Genes in Conflict : The Biology of Selfish Genetic Elements. Belknap Press, Harvard. ISBN 0-674-01713-7 • Trivers R, Palestis BG, Zaatari D. (2009) The Anatomy of a Fraud: Symmetry and Dance TPZ Publishers ISBN 978-0-615-28756-0 • Trivers R (2011) The Folly of Fools: The Logic of Deceit and Self-Deception in Human Life Basic Books ISBN 978-0-465-02755-2

References [1] The Kindness of Strangers (http:/ / www. guardian. co. uk/ books/ 2005/ aug/ 27/ featuresreviews. guardianreview9) [2] The Evolutionary Revolutionary (http:/ / www. boston. com/ news/ globe/ ideas/ articles/ 2005/ 03/ 27/ the_evolutionary_revolutionary?pg=full) [3] The Evolutionary Revolutionary (http:/ / www. boston. com/ news/ globe/ ideas/ articles/ 2005/ 03/ 27/ the_evolutionary_revolutionary?pg=full), The Boston Globe, 27 March 2005 [4] Trivers, R.L. & Newton, H.P. Science Digest 'The crash of flight 90: doomed by self-deception?' November 1982, pp 66,67,111. [5] A Full-Force Storm with Gale Winds Blowing (http:/ / www. edge. org/ 3rd_culture/ trivers04/ trivers04_index. html) [6] http:/ / dx. doi. org/ 10. 1086%2F406755 [7] http:/ / www. jstor. org/ stable/ 2822435 [8] http:/ / www1. appstate. edu/ ~kms/ classes/ psy2664/ Documents/ trivers. pdf [9] http:/ / adsabs. harvard. edu/ abs/ 1973Sci. . . 179. . . 90T [10] http:/ / dx. doi. org/ 10. 1126%2Fscience. 179. 4068. 90 [11] http:/ / www. jstor. org/ stable/ 1734960 [12] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 4682135 [13] http:/ / dx. doi. org/ 10. 1093%2Ficb%2F14. 1. 249 [14] http:/ / adsabs. harvard. edu/ abs/ 1976Sci. . . 191. . 249T [15] http:/ / dx. doi. org/ 10. 1126%2Fscience. 1108197 [16] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 1108197

External links • Rutgers University faculty listing (http://anthro.rutgers.edu/index.php?option=com_content&task=view& id=102&Itemid=136) • http://roberttrivers.com

128

George C. Williams

129

George C. Williams George C. Williams Born

May 12, 1926 Charlotte, North Carolina

Died

September 8, 2010 (aged 84)

Nationality

American

Fields

Biology

Institutions

Stony Brook University

Alma mater

UCLA

Known for

theories of natural selection

Influences

Charles Darwin

Influenced

Richard Dawkins

Notable awards Daniel Giraud Elliot Medal (1992) Crafoord Prize (1999)

George Christopher Williams (May 12, 1926 – September 8, 2010) was an American evolutionary biologist.[1] Williams was a professor emeritus of biology at the State University of New York at Stony Brook. He was best known for his vigorous critique of group selection, though later in life he recognized that it did sometimes occur. The work of Williams in this area, along with W. D. Hamilton, John Maynard Smith and others led to the development of a gene-centric view of evolution in the 1960s.

Academic work Williams' 1957 paper Pleiotropy, Natural Selection, and the Evolution of Senescence is one of the most influential in 20th century evolutionary biology, and contains at least 3 foundational ideas. The central hypothesis of antagonistic pleiotropy remains the prevailing evolutionary explanation of senescence. In this paper Williams was also the first to propose that senescence should be generally synchronized by natural selection. According to this original formulation "if the adverse genic effects appeared earlier in one system than any other, they would be removed by selection from that system more readily than from any other. In other words, natural selection will always be in greatest opposition to the decline of the most senescence-prone system." This important concept of synchrony of senescence was taken up a short time later by John Maynard Smith, and the origin of the idea is often misattributed to him, including in his obituary in the journal, Nature. This paper also contains the first basic outline of the so-called "grandmother hypothesis", which states that natural selection might select for menopause and post-reproductive life in females, although Williams does not explicitly mention grandchildren or the inclusive fitness contribution of grandparenting. In his first book, Adaptation and Natural Selection, Williams advocated a "ground rule - or perhaps doctrine would be a better term - ... that adaptation is a special and onerous concept that should only be used where it is really necessary",[2] and, that, when it is necessary, selection among genes or individuals would in general be the preferable explanation for it. He elaborated this view in later books and papers, which contributed to the development of a gene-centered view of evolution; Richard Dawkins built on Williams' ideas in this area in the book The Selfish Gene. Williams was also well known for his work on the evolution of sex, and was an advocate of evolutionary medicine.

George C. Williams In later books, including Natural Selection: Domains, Levels and Challenges, Williams softened his views on group selection, recognizing that clade selection, trait group selection and multilevel selection did sometimes occur in nature, something he had earlier thought to be so unlikely it could be safely ignored.[3] Williams became convinced that the genic neo-Darwinism of his earlier years, while essentially correct as a theory of microevolutionary change, could not account for evolutionary phenomena over longer time scales, and was thus an "utterly inadequate account of the evolution of the Earth’s biota" (1992, p. 31). In particular, he became a staunch advocate of clade selection – a generalisation of species selection to monophyletic clades of any rank – which could potentially explain phenomena such as adaptive radiations, long-term phylogenetic trends, and biases in rates of speciation/extinction. In Natural Selection (1992), Williams argued that these phenomena cannot be explained by selectively-driven allele substitutions within populations, the evolutionary mechanism he had originally championed over all others. This book thus represents a substantial departure from the position of Adaptation and Natural Selection.

Academic career Williams received a Ph.D. in biology from the University of California at Los Angeles in 1955. At Stony Brook he taught courses in marine vertebrate zoology, and he often used ichthyological examples in his books. In 1992 Williams was awarded the Daniel Giraud Elliot Medal from the National Academy of Sciences. He won the Crafoord Prize for Bioscience jointly with Ernst Mayr and John Maynard Smith in 1999. Dawkins describes Williams as "one of the most respected of American evolutionary biologists".

Books • • • •

Williams, G.C. 1966. Adaptation and Natural Selection. Princeton University Press, Princeton, N.J. Williams, G.C., ed. 1971. Group Selection. Aldine-Atherton, Chicago. Williams, G.C. 1975. Sex and Evolution. Princeton University Press, Princeton, N.J. Paradis, J. and G.C. Williams. 1989. T.H. Huxley’s Evolution and Ethics : with New Essays on its Victorian and Sociobiological Context. Princeton University Press, Princeton, N.J. • Williams, G.C. 1992. Natural Selection: Domains, Levels, and Challenges. Oxford University Press, New York. • Nesse, R.M. and G.C. Williams. 1994. Why We Get Sick : the New Science of Darwinian Medicine. Times Books, New York. • Williams, G.C. 1996. Plan and Purpose in Nature. Weidenfeld & Nicolson, London (published in the U.S. in 1997 as The Pony Fish’s Glow : and Other Clues to Plan and Purpose in Nature. Basic Books, New York).

Selected papers • Williams, G. C. 1957. Pleiotropy, Natural Selection, and the Evolution of Senescence. Evolution 11;4: 398-411 • Taylor, P. O. and G. C. Williams. 1984. Demographic parameters at evolutionary equilibrium. Canadian Journal of Zoology 62: 2264-2271. • Williams, G. C. 1985. A defense of reductionism in evolutionary biology. Oxford Surveys in Evolutionary Biology 2: 127. • Williams, G. C. 1988. Huxley's Evolution and Ethics in sociobiological perspective. Zygon 23: 383-438. • Williams, G. C. 1995. A package of information [4] In J. Brockman, ed., The Third Culture, New York: Touchstone, pp. 38–50.

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References [1] [2] [3] [4]

http:/ / chronicle. com/ blogPost/ George-C-Williams-1926-2010/ 26821/ Adaptation and Natural Selection (http:/ / www. amazon. com/ dp/ 0691026157) p4 George C. Williams, Natural Selection: Domains, Levels and Challenges, (Oxford University Press, 1992), 23-55 http:/ / www. edge. org/ documents/ ThirdCulture/ h-Ch. 1. html

External links • Official website (http://life.bio.sunysb.edu/ee/people/williamsindex.html) Wikipedia:Link rot • Article from Science by [[Carl Zimmer (http://life.bio.sunysb.edu/ee/upload/williams.pdf)]] • A Conversation With George C. Williams by Frans Roes (http://www.stephenjaygould.org/library/ williams_interview.html) • Obituary by Richard Dawkins, October 1, 2010 (http://richarddawkins.net/articles/ 527711-george-c-williams-1926–2010)

Sarah Blaffer Hrdy Sarah Hrdy (née Blaffer; born July 11, 1946) is an American anthropologist and primatologist who has made several major contributions to evolutionary psychology and sociobiology. She has been selected as one of the 21 “Leaders in Animal Behavior”.[1]

Biography Early life Sarah Blaffer was born on July 11, 1946, in Dallas, Texas. She was raised in Houston, and attended St. John's School there.

Education At age 18, Sarah attended her mother's alma mater, Wellesley College in Massachusetts. Her intended philosophy major led her into research on Mayan folklore, leading to a transfer to Radcliffe College and major in anthropology. She graduated summa cum laude and Phi Beta Kappa in 1969 with that B.A.. (Her undergraduate thesis became the basis for The Black Man of Zincantan, published in 1972.) She took film-making courses at Stanford, hoping to make films on health care for developing countries. Finding herself disappointed with those classes, she entered Harvard in 1970.

Family life Blaffer met Daniel Hrdy at Harvard. They married in 1972 in Kathmandu. They have three children: • daughter Katrinka, born to Hrdy at age 31 • daughter Sasha, born 1982, a week before Hrdy was scheduled to present a paper at Cornell University • son Niko, born 1986, when she was 41 She lives with her husband in Northern California, where they operate the Citrona Farms walnut plantation.[2] She is Professor Emeritus of Anthropology at the University of California at Davis, where she remains involved with the Animal Behavior Graduate Group.

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Main research The Langurs of Abu Sarah Hrdy first became interested in langurs during an undergraduate primate behavior class taught by anthropologist Irven DeVore in 1968. Here, a remark by DeVore regarding the relationship between crowding and the killing of infants would forever change her life. After graduation, she returned to Harvard for graduate studies, with the goal of decoding this phenomenon of infanticide in langur colonies. Working under the supervision of DeVore and the evolutionary biologist Robert L. Trivers provided Hrdy with an introduction to a newly emerging outlook on the social world—that of sociobiology—which crystallized at Harvard during the early 1970s and shaped Hrdy's perspective on primatology in an enduring manner. Hrdy's PhD thesis tested the hypothesis that overcrowding causes infanticide in langur colonies. She went to Mount Abu in India to study Hanuman Langurs, and concluded that infanticide was independent of overcrowding—it was possibly an evolutionary tactic: When an outside male takes over a group, he usually proceeds to kill all infants. This postulated tactic would be very advantageous to the male langurs who practiced infanticide. Turnover in a langur tribe occurs approximately every 27 months. The male who is taking over has a very small window of opportunity to pass on his genes, and if the females are already nursing infants, it's likely that they won't ovulate again for another year. Killing their dependent infants makes the females once again receptive to mating. Female choice is subverted, as females are put under pressure to ovulate and are forced to breed with the infanticidal males. This is where the idea of sexual counter-strategies comes into play. Hrdy theorized that by mating with as many males as possible, particularly outside males who are not part of the colony, mothers are able to successfully protect their young, as males were unlikely to kill an infant if there was the slightest chance that it might be their own. That gives an "illusion of paternity", as Trivers put it. The goal of the male langur is to maximize the proportion of his offspring, and according to Hrdy, a male who attacks his own offspring is rapidly selected against. While infanticide has been seemingly preserved across primate orders, Hrdy found no evidence to suggest that the human species has a 'genetic imperative' for infanticide. In 1975 Hrdy was awarded her PhD for her research on langurs. In 1977 it was published in her second book, The Langurs of Abu: Female and Male Strategies of Reproduction. The controversy in the anthropology realm that her research sparked was not surprising—the classic belief that primates act for the good of the group was discarded, and the field of sociobiology gained increasing support. Many mistakenly assumed that she implied existence of an 'infanticidal gene' that could be conserved across primates. Today, her results and conclusions are widely received. Even Trivers, who once dismissed her apparently illogical convictions, admits that her theory regarding female sexual strategies has "worn well".

The Woman That Never Evolved Hrdy's third book came out in 1981: The Woman That Never Evolved. She begins chapter one with a sentence indicating that the results of her work suggest females should be given a lot more credibility than previously thought. "Biology, it is sometimes thought, has worked against women." Here, Hrdy expands upon female primate strategies. This book is one of the New York Times' Notable Books of 1981. In 1984 Hrdy co-edited Infanticide: Comparative and Evolutionary Perspectives. It was selected as one of the 1984–1985 "Outstanding Academic Books" by Choice, the Journal of the Association of College and Research Libraries.

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Mother Nature In 1999, Hrdy published Mother Nature - Maternal Instincts and How They Shape the Human Species. She places a sociobiological twist on maternal instinct, and places "human mothers and infants in a broader comparative and evolutionary framework" offering a new perspective on mother-infant interdependence. She discusses how mothers are continually making trade-offs between quality and quantity" and weighing the best possible actions for both her and her infant. Hrdy's view is that there is no defined 'maternal instinct', as it depends on a number of variables, and is therefore not innate, as once thought. She also stands by her view that humans evolved as cooperative breeders, making them essentially unable to raise offspring without a helper. This is where the concept of allomothering comes in—relatives other than the mother, such as the father, grandparents, and older siblings, as well as genetically unrelated helpers, such as nannies, nurses, and child care groups, who spend time with an infant, leaving the mother with more free time to meet her own needs. She is a strong advocate for making affordable child care a priority.

Significant works Books • 1972 - The Black-man of Zinacantan: A Central American Legend. The Texas Pan American Series. Austin: University of Texas Press. ISBN 0-292-70701-0. • 1977 - The Langurs of Abu: Female and Male Strategies of Reproduction. Cambridge: Harvard University Press. ISBN 0-674-51058-5. • 1981 - The Woman that Never Evolved. Cambridge: Harvard University Press. (Chosen by the New York Times Book Review as one of Notable Books of the Year in Science and Social Science.) 1982, Japanese edition, Tokyo: Shisaku-sha Publishing; 1984, 5th printing of paperback edition, Cambridge; 1984, 1st French edition, Des guenons et des femmes. Paris: Editions Tierce, in press, 2nd French edition, Paris: Payot et Rivage; 1985, Italian edition, La Donna Che Non si E'evoluta, Franco Angeli Editore. ISBN 0-674-95539-0. • 1984 - Hausfater, G. and S. Hrdy, eds. Infanticide: Comparative and Evolutionary Perspectives. New York: Aldine Publishing Co. (Selected as one of the 1984-85 "Outstanding Academic Books" by Choice, the Journal of the Association of College and Research Libraries.) ISBN 0-202-36221-3. • 1999 - Mother Nature: A History of Mothers, Infants and Natural Selection. New York: Pantheon. A BOMC Alternative Selection; Selected by both Publisher's Weekly and by Library Journal as one of Best Books of 1999 and a finalist for PEN USA West 2000 Literary Award for Research Nonfiction. Won the Howells Prize for Outstanding Contribution to Biological Anthropology. (Published in UK as Mother Nature: Natural selection and the female of the species. London: Chatto and Windus); also translated into Chinese, Dutch, French, German, Italian, Portuguese, Spanish, Japanese, Korean and Polish. ISBN 0-679-44265-0. • 2001 - "The Past, Present, and Future of the Human Family." The Tanner Lectures on Human Values, Delivered at University of Utah February 27 and 28, 2001. • 2005 - The 92nd Dahlem Workshop Report, "Attachment and Bonding: A New Synthesis." Edited by C. S. Carter, L. Ahnert, K. E. Grossmann, S. B. Hrdy, M. E. Lamb, S. W. Porges, and N. Sachser. ©MIT Press. ISBN 0-262-03348-8. • 2009 - Mothers and Others: The Evolutionary Origins of Mutual Understanding. Cambridge: Harvard University Press. ISBN 0-674-03299-3. • 2010 -Myths, monkeys and motherhood: An intellectual autobiography. In Lee Drickamer and Donald Dewsbury (eds.), Leaders in Animal Behavior: The Second Generation. Cambridge: Cambridge University Press, pp. 343–344

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Films • 1977 - Hrdy, S., D. B. Hrdy and John Melville Bishop. Stolen copulations; Play and Kidnapped, 16 mm, color. • 1980 - Hrdy, S., Vishnu Mathur and William Whitehead. "Hanuman langur: Monkey of India," 30 minutes, color. Canadian Broadcasting Corporation. Available on video cassette: CBC Enterprises, P. O. Box 500, Terminal A, Toronto, Ontario, Canada M5W 1E6. • 1983 - "Treatment for film on reproductive strategies of female primates," for BBC Natural History Unit, Bristol, UK. • 1988 - "Monkeys of Abu." National Geographic Explorer. May 1988. • 1990 - Nature Advisory Board, Channel Thirteen New York for series on the natural history of sex. • 1990 - Consultant for "Human Nature" for the British Broadcasting Corporation, Bristol, UK. • 2001 - Advisor for PBS series "Evolution".

Awards • Guggenheim Fellow • Member, National Academy of Sciences • Member, American Academy of Arts and Sciences • Member, California Academy of Sciences • 2003 recipient, University of California Panunzio award (honoring outstanding scholarly work and service achievements since retirement) • NYT Notable Books of 1981, The Woman That Never Evolved • Publisher's Weekly, "Best Books of 1999", Mother Nature • Library Journal, "Best Books of 1999", Mother Nature • Howells Prize for Outstanding Contributions to Biological Anthropology, Mother Nature

References [1] Drickamer L. Dewsbury S. (Dec 2009) Leaders in Animal Behaviour: The Second Generation. Cambridge University Press. [1 UNIQ-nowiki-0-fa2f87de963a2a49-QINU (http:/ / www. cambridge. org/ us/ academic/ subjects/ life-sciences/ animal-behaviour/ leaders-animal-behavior-second-generation) [2] Citrona Farms (http:/ / www. citrona. com)

• "Scientists' Nightstand: Interview with Sarah Hrdy" (http://www.americanscientist.org/bookshelf/pub/ sarah-blaffer-hrdy), American Scientist, Nov. 25, 2003. • With a Little Help from My Friends (http://www.americanscientist.org/bookshelf/pub/ with-a-little-help-from-my-friends), American Scientist • Susan Caba, "She loves me, she loves me not" (http://www.salon.com/mwt/feature/1999/12/09/maternal/), Salon.com, Dec. 9, 1999 (reviewing and discussing Mother Nature, and interviewing Hrdy) • Claudia Glenn Dowling, "The Hardy Sarah Blaffer Hrdy" (http://discovermagazine.com/2003/mar/feathrdy), Discover Magazine, March 1, 2003 (interview, discussion of Mother Nature and other work)

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Richard Machalek

Richard Machalek Richard Machalek (born April 12, 1946) is a social theorist, sociobiologist, and professor of sociology. A student and colleague of sociobiologist E.O. Wilson, Machalek is best known for using traditional sociological frameworks and theories to explain complex social behavior and structures in non-human societies, with a special emphasis on ant populations. Machalek has repeatedly – and only half jokingly – called for a “one-hundred year moratorium on studying humans within the field of sociology” and he thus believes that the bedrock of sociological knowledge lies in explaining social phenomena that are exhibited across many different types of species. He can be, and often is, considered a radical sociological theorist in this regard. As such, Machalek also applies knowledge from the fields of evolutionary theory, zoology, and biology and is especially concerned with the trans-species social behaviors of cheating, cooperation, and division of labor, among others. Besides teaching at a number of universities, in 1986 he was a visiting professor at Harvard University’s Museum of Comparative Zoology under the tutelage of E.O. Wilson, and currently teaches sociology at the University Of Wyoming.

External links • Machalek's University of Wyoming Faculty page [1] - Including Vita, publications, and contact info.

References [1] http:/ / uwacadweb. uwyo. edu/ machalek/

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Steven Pinker

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Steven Pinker Steven Pinker

Pinker in 2011 Born

Steven Arthur Pinker September 18, 1954 Montreal, Quebec, Canada

Residence

United States

Citizenship

Canadian

Fields

Evolutionary psychology, experimental psychology, cognitive science, linguistics, visual cognition

Alma mater

Dawson College, McGill University, Harvard University

Known for

How the Mind Works, The Blank Slate, The Better Angels of Our Nature

Influences

Noam Chomsky, Thomas Sowell, Leda Cosmides, John Tooby, Richard Dawkins, Thomas Schelling

[1]

Notable awards Troland Award [2] (1993, National Academy of Sciences), Henry Dale Prize (2004, Royal Institution), Walter P. Kistler Book Award (2005), Humanist of the Year award (2006, issued by the AHA), George Miller Prize (2010, Cognitive Neuroscience Society) Spouse

Nancy Etcoff (1980–1992; divorced) Ilavenil Subbiah (1995–?; divorced) Rebecca Goldstein (?-present) Website http:/ / www. stevenpinker. com/

Steven Arthur Pinker (born September 18, 1954) is a Canadian-born experimental psychologist, cognitive scientist, linguist, and popular science author. He is a Harvard College Professor and the Johnstone Family Professor in the Department of Psychology at Harvard University,[3] and is known for his advocacy of evolutionary psychology and the computational theory of mind. Pinker's academic specializations are visual cognition and psycholinguistics. His experimental subjects include mental imagery,

Pinker in Göttingen in 2010

Steven Pinker shape recognition, visual attention, children's language development, regular and irregular phenomena in language, the neural bases of words and grammar, and the psychology of innuendo and euphemism. He published two technical books which proposed a general theory of language acquisition and applied it to children's learning of verbs. In his popular books, he has argued that language is an "instinct" or biological adaptation shaped by natural selection. He is the author of six books for a general audience: The Language Instinct (1994), How the Mind Works (1997), Words and Rules (2000), The Blank Slate (2002), The Stuff of Thought (2007), and The Better Angels of Our Nature (2011).

Biography Early life, education and career Pinker was born in Montreal, Quebec, Canada in 1954, to a middle-class Jewish family. His parents were Roslyn and Harry Pinker. He graduated from Dawson College in 1971. He received a Bachelor of Arts in psychology from McGill University in 1976, and then went on to earn his Doctor of Philosophy in experimental psychology at Harvard University in 1979. He did research at the Massachusetts Institute of Technology (MIT) for a year, after which he became an assistant professor at Harvard and then Stanford University. From 1982 until 2003, Pinker taught at the Department of Brain and Cognitive Sciences at MIT, and eventually became the director of the Center for Cognitive Neuroscience, taking a one-year sabbatical at the University of California, Santa Barbara, in 1995–96. As of 2003, he is the Johnstone Family Professor of Psychology at Harvard; from 2008 to 2013 he also held the title of Harvard College Professor in recognition of his dedication to teaching. In June 2011 it was announced he would give lectures as a visiting professor at the New College of the Humanities, a private college in London.[4] Pinker was named one of Time Magazine's 100 most influential scientists and thinkers in the world in 2004[5] and one of Prospect and Foreign Policy's 100 top public intellectuals in both years the poll was carried out, 2005[6] and 2008; in 2010 and 2011 he was named by Foreign Policy magazine to its list of top global thinkers. His research in cognitive psychology has won the Early Career Award (1984) and Boyd McCandless Award (1986) from the American Psychological Association, the Troland Research Award (1993) from the National Academy of Sciences, the Henry Dale Prize (2004) from the Royal Institution of Great Britain, and the George Miller Prize (2010) from the Cognitive Neuroscience Society. He has also received honorary doctorates from the universities of Newcastle, Surrey, Tel Aviv, McGill, and the University of Tromsø, Norway. He was twice a finalist for the Pulitzer Prize, in 1998 and in 2003. In January 2005, Pinker defended Lawrence Summers, President of Harvard University, whose comments about a gender gap in mathematics and science angered much of the faculty. Pinker noted that Summers's remarks, properly understood, were hypotheses about overlapping statistical distributions of men’s and women’s talents and tastes, and that in a university such hypotheses ought to be the subject of empirical testing rather than dogma and outrage.[7] On May 13, 2006, Pinker received the American Humanist Association's Humanist of the Year award for his contributions to public understanding of human evolution. In 2009, Pinker wrote a mixed review of Malcolm Gladwell's essays in The New York Times criticizing his analytical methods. Gladwell published a rebuttal in the Times regarding Pinker's comments about the importance of IQ on teaching performance and by analogy, the effect, if any, of draft order on quarterback performance in the National Football League. Pinker then responded to Gladwell's rebuttal. The exchange prompted Advanced NFL Stats to step in and address the issue statistically, siding with Pinker in that draft order is indeed correlated with quarterback performance. The differences in Gladwell and Pinker's respective methodologies caused the disagreement. Gladwell evaluated a performance on a per-play basis, whereas Pinker and Advanced NFL Stats evaluated performance based on total overall productivity based on the assumption that starters were superior players to backups.

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Steven Pinker Pinker has served on the editorial boards of journals such as Cognition, Daedalus, and PLOS One, and on the advisory boards of institutions for scientific research (e.g., the Allen Institute for Brain Science), free speech (e.g., the Foundation for Individual Rights in Education), the popularization of science (e.g., the World Science Festival and the Committee for Skeptical Inquiry), peace (e.g., the Peace Research Endowment), and secular humanism (e.g., the Freedom from Religion Foundation and the Secular Coalition for America). Since 2008, he has chaired the Usage Panel of the American Heritage Dictionary, and wrote the essay on usage for the fifth edition of the Dictionary, which was published in 2011.

Personal life His father, a lawyer, first worked as a manufacturer's representative, while his mother was first a home-maker then a guidance counselor and high-school vice-principal. He has two younger siblings. His brother is a policy analyst for the Canadian government. His sister, Susan Pinker, is a psychologist and writer, author of The Sexual Paradox.[8] Pinker married Nancy Etcoff in 1980 and they divorced in 1992; he married Ilavenil Subbiah in 1995 and they too divorced.[9] His current wife, whom he married in 2007, is the novelist and philosopher Rebecca Goldstein.[10] He has two stepdaughters: the novelist Yael Goldstein Love and the poet Danielle Blau. He has said, "I was never religious in the theological sense... I never outgrew my conversion to atheism at 13, but at various times was a serious cultural Jew."[11] As a teenager, he says he considered himself an anarchist until he witnessed civil unrest following a police strike in 1969.[12] Pinker identifies himself as a feminist, specifically an equity feminist, and in The Blank Slate defines equity feminism as "a moral doctrine about equal treatment that makes no commitments regarding open empirical issues in psychology or biology".[13] He has reported the result of a test of his political orientation that characterized him as "neither leftist nor rightist, more libertarian than authoritarian."[14] Pinker confesses to having "experienced a primitive tribal stirring" after his genes were shown to trace back to the Middle East. In June 2013 he was the guest on UK BBC Radio 4's Desert Island Discs. His choices were "Further On Up the Road" by Eric Clapton, "Der Alter Bulgar" by the Andy Statman Klezmer Orchestra (with Itzhak Perlman), "God’s Comic" by Elvis Costello, "Olé" by John Coltrane, "Yellow Moon" by The Neville Brothers, "I'm Your Man" by Leonard Cohen, "Machine Gun" by Jimi Hendrix and "Lullaby of Birdland" by Sarah Vaughan.

Research and theory Pinker’s research on visual cognition, begun in collaboration with his thesis adviser Stephen Kosslyn, showed that mental images represent scenes and objects as they appear from a specific vantage point (rather than capturing their intrinsic three-dimensional structure), and thus correspond to the neuroscientist David Marr’s theory of a “two-and-a-half-dimensional sketch.”[15] He also showed that this level of representation is used in visual attention, and in object recognition (at least for asymmetrical shapes), contrary to Marr’s theory that recognition uses viewpoint-independent representations. In psycholinguistics, Pinker became known early in his career for promoting computational learning theory as a way to understand language acquisition in children. He wrote a tutorial review of the field followed by two books that advanced his own theory of language acquisition, and a series of experiments on how children acquire the passive, dative, and locative constructions. In 1989 Pinker and Alan Prince published an influential critique of a connectionist model of the acquisition of the past tense (a textbook problem in language acquisition), followed by a series of studies of how people use and acquire the past tense. This included a monograph on children’s regularization of irregular forms, and a popular 1999 book, Words and Rules: The Ingredients of Language, in which he argued that regular and irregular phenomena were products of computation and memory lookup, respectively, and that language could be understood as an interaction between the two.

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Steven Pinker The Language Instinct (1994) was the first of several books that combined cognitive science with behavioral genetics and evolutionary psychology. In it he introduced the science of language and popularized Noam Chomsky's theory that language is an innate faculty of mind, with the twist that this faculty evolved by natural selection as a Darwinian adaptation for communication (both ideas remain controversial; see below). Two other books, How the Mind Works (1997) and The Blank Slate (2002), broadly surveyed the mind and defended the idea of a complex human nature which comprises many mental faculties that are adaptive (and is an ally of Daniel Dennett and Richard Dawkins in many disputes surrounding adaptationism). Another major theme in Pinker's theories is that human cognition works, in part, by combinatorial symbol-manipulation, not just associations among sensory features, as in many connectionist models. In 2011 Pinker published The Better Angels of Our Nature, which argued that violence has decreased over multiple scales of time and magnitude, including tribal warfare, homicide, cruel punishments, child abuse, animal cruelty, domestic violence, lynching, pogroms, and international and civil wars. He tried to explain these declines as a change in the interaction between violent impulses (such as dominance, sadism and revenge) and peaceful impulses (such as self-control, empathy and reason) brought on by historical forces such as government, trade and literacy. Pinker's books, The Language Instinct, How the Mind Works, Words and Rules, The Blank Slate, and The Stuff of Thought combine cognitive science with behavioral genetics and evolutionary psychology.

Music and cognition A number of theories about the evolutionary origins of language have argued that linguistic cognition might have evolved from earlier musical cognition. Pinker is a critic of this view, as he sees language as being tied primarily to the capacity for logical reasoning. In How the Mind Works (1997) Pinker reiterates an argument from Immanuel Kant positing that music is not in itself an important cognitive phenomenon, but that music simply happens to stimulate important auditory and spatio-motoric cognitive functions. Pinker compares music to "auditory cheesecake", and states that "As far as biological cause and effect is concerned, music is useless". This argument has been rejected by a number of experts in music cognition who argue that music has had an important role in the evolution of human cognition, for example Daniel Levitin and Joseph Carroll.[16][17][18][19] Levitin argues against Pinker’s view of music and cognition in his book This Is Your Brain On Music.[20]

Criticism The Language Instinct has been criticized by Geoffrey Sampson in his book, The 'Language Instinct' Debate. The assumptions underlying the nativist view have also been subject to sustained criticism in Jeffrey Elman's Rethinking Innateness: A Connectionist Perspective on Development (Neural Networks and Connectionist Modeling), which defends the connectionist approach that Pinker has criticized. David Shenk has criticized Pinker for an article he wrote in The New York Times which addressed the nature versus nurture debate. He criticized him for siding with the "nature" argument and for "never once acknowledg[ing] gene-environment interaction or epigenetics." Shenk contends that because of these factors the debate over nature versus nurture has been "rendered obsolete."[21] Pinker responded to a question about epigenetics as a possibility for the decline in violence in a lecture for the BBC World Service. Pinker said it was unlikely since the decline in violence happened too rapidly to be explained by genetic changes.[22] Edward S. Herman and David Peterson have criticized The Better Angels of Our Nature. In an extensive essay, they accuse Pinker of having ventured outside of his field of expertise, and of cherry-picking historical data to fit his thesis.

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Bibliography Books • Language Learnability and Language Development (1984) ISBN 978-0-674-51055-5 • Visual Cognition (1985) ISBN 978-0-262-66178-2 • Connections and Symbols (1988) ISBN 978-0-262-66064-8 • Learnability and Cognition: The Acquisition of Argument Structure (1989) ISBN 978-0-262-66073-0 • Lexical and Conceptual Semantics (1992) ISBN 978-1-55786-354-6 • The Language Instinct (1994) ISBN 978-0-06-097651-4 • How the Mind Works (1997) ISBN 978-0-393-31848-7 • Words and Rules: The Ingredients of Language (1999) ISBN 978-0-465-07269-9 • The Blank Slate: The Modern Denial of Human Nature (2002) ISBN 978-0-670-03151-1 • The Best American Science and Nature Writing (editor and introduction author, 2004) ISBN 978-0-618-24698-4 • Hotheads (an extract from How the Mind Works, 2005) ISBN 978-0-14-102238-3

Steven Pinker in 2005

• The Stuff of Thought: Language as a Window into Human Nature (2007) ISBN 978-0-670-06327-7 • The Better Angels of Our Nature: Why Violence Has Declined (2011) ISBN 978-0-670-02295-3

Articles and essays • Pinker, S. (1991). "Rules of Language". Science 253 (5019): 530–535. doi:10.1126/science.1857983 [23]. • Ullman, M.; Corkin, S.; Coppola, M.; Hickok, G.; Growdon, J. H.; Koroshetz, W. J.; Pinker, S. (1997). "A neural dissociation within language: Evidence that the mental dictionary is part of declarative memory, and that grammatical rules are processed by the procedural system". Journal of Cognitive Neuroscience 9: 289–299. • Pinker, S. (2003) "Language as an adaptation to the cognitive niche" In M. Christiansen & S. Kirby (Eds.), Language evolution: States of the Art New York: Oxford University Press. • Pinker, S. (2005). "So How Does the Mind Work?". Mind and Language 20 (1): 1–24. doi:10.1111/j.0268-1064.2005.00274.x [24]. • Jackendoff, R.; Pinker, S. (2005). "The nature of the language faculty and its implications for evolution of language" (Reply to Fitch, Hauser, & Chomsky)". Cognition 97 (2): 211–225. doi:10.1016/j.cognition.2005.04.006 [25]. • S. Pinker (2007), "In Defense of Dangerous Ideas" Chicago Sun-Times, July 15, 2007 [26] • Pinker, S. (2012). The False Allure of Group Selection [27]. Edge, Jun 19, 2012. • A number of Pinker's articles at Harvard [28] • Pinker, S. (2013). Science Is Not Your Enemy [29]. The New Republic, Aug 6, 2013.

Steven Pinker

References [1] C-SPAN | BookTV "In Depth with Steven Pinker" (http:/ / www. c-spanvideo. org/ program/ 282181-1) November 2nd 2008 [2] http:/ / www. nasonline. org/ about-nas/ awards/ troland-research-awards. html [3] Steven Pinker – About. Department of Psychology Harvard University (http:/ / pinker. wjh. harvard. edu/ about/ index. html) Accessed 2010-02-28 [4] "The professoriate" (http:/ / www. nchum. org/ who-we-are/ the-professoriate), New College of the Humanities, accessed June 8, 2011. [5] "Steven Pinker: How Our Minds Evolved" by Robert Wright (http:/ / pinker. wjh. harvard. edu/ about/ media/ 2004_04_26_time. htm) Time Magazine Accessed 2006-02-08 [6] "The Prospect/FP Top 100 Public Intellectuals" (http:/ / www. foreignpolicy. com/ story/ cms. php?story_id=3249) Foreign Policy (free registration required) Accessed 2006-082-08 [7] "PSYCHOANALYSIS Q-and-A: Steven Pinker" (http:/ / www. thecrimson. com/ article. aspx?ref=505366) The Harvard Crimson Accessed 2006-02-08 [8] "Steven Pinker: the mind reader" (http:/ / www. guardian. co. uk/ Archive/ Article/ 0,4273,3926387,00. html) The Guardian Accessed 2006-11-25 [9] Biography for Steven Pinker at imdb (http:/ / www. imdb. com/ name/ nm0684348/ bio) Accessed 2007-09-12 [10] "How Steven Pinker Works" by Kristin E. Blagg (http:/ / pinker. wjh. harvard. edu/ about/ media/ 2005_11_04_harvardcrimson. html) The Harvard Crimson Accessed 2006-02-03 [11] "Steven Pinker: the mind reader" by Ed Douglas (http:/ / www. guardian. co. uk/ Archive/ Article/ 0,4273,3926387,00. html) The Guardian Accessed 2006-02-03 [12] "As a young teenager in proudly peaceable Canada during the romantic 1960s, I was a true believer in Bakunin's anarchism. I laughed off my parents' argument that if the government ever laid down its arms all hell would break loose. Our competing predictions were put to the test at 8:00 A.M. on October 17, 1969, when the Montreal police went on strike. ... This decisive empirical test left my politics in tatters (and offered a foretaste of life as a scientist)." – Pinker, Steven (2002), The Blank Slate: The Modern Denial of Human Nature, Penguin Putnam, ISBN 0-670-03151-8. [13] Pinker, Steven, The Blank Slate: The Modern Denial of Human Nature (Viking, 2002), p. 341 [14] "My Genome, My Self" by Steven Pinker (http:/ / www. nytimes. com/ 2009/ 01/ 11/ magazine/ 11Genome-t. html?_r=1& scp=1& sq=my genome myself& st=cse) The New York Times Sunday Magazine Accessed 2010-04-10 [15] The nature of the language faculty and its implications for evolution of language [16] Levitin, D. J. and Tirovolas, A. K. (2009), Current Advances in the Cognitive Neuroscience of Music. Annals of the New York Academy of Sciences, 1156: 211–231. [17] Perlovsky L. Music. Cognitive Function, Origin, And Evolution Of Musical Emotions . WebmedCentral PSYCHOLOGY 2011;2(2):WMC001494 [18] Alison Abbott. 2002. Neurobiology: Music, maestro, please! Nature 416, 12–14 (7 March 2002) | doi:10.1038/416012a [19] Cross, I. (1999). Is music the most important thing we ever did? Music, development and evolution. [preprint (html)] [preprint (pdf)] In Suk Won Yi (Ed.), Music, mind and science (pp10-39), Seoul: Seoul National University Press. [20] Levitin, Daniel. 2006. This Is Your Brain On Music: The Science of a Human Obsession, New York: Dutton/Penguin. [21] " Steven Pinker's "probabilistic" genes (http:/ / geniusblog. davidshenk. com/ 2009/ 01/ steven-pinkers-probabilistic-genes. html)," David Shenk [22] " Exchanges At The Frontier 2011 (http:/ / www. bbc. co. uk/ programmes/ p00m98km)", BBC. [23] http:/ / dx. doi. org/ 10. 1126%2Fscience. 1857983 [24] http:/ / dx. doi. org/ 10. 1111%2Fj. 0268-1064. 2005. 00274. x [25] http:/ / dx. doi. org/ 10. 1016%2Fj. cognition. 2005. 04. 006 [26] http:/ / richarddawkins. net/ article,1449,In-defense-of-dangerous-ideas,Steven-Pinker) [27] http:/ / edge. org/ conversation/ the-false-allure-of-group-selection [28] http:/ / pinker. wjh. harvard. edu/ articles/ [29] http:/ / www. newrepublic. com/ article/ 114127/ science-not-enemy-humanities

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External links • Official website (http://stevenpinker.com/) Interviews • • • •

In Depth interview with Pinker, November 2, 2008 (http://www.c-spanvideo.org/program/282181-1) Steven Pinker on The Hour (http://www.cbc.ca/thehour/video.php?id=1918) Steven Pinker Multimedia (http://www.reitstoen.com/pinker.php). Extensive lists of audio and video files Biology vs. the Blank Slate (http://reason.com/0210/fe.rb.biology.shtml) Reason magazine interview with Pinker • Live interview with Steven Pinker on The Blank Slate (http://www.booktalk.org/transcripts/transcript5.php) • Online video interview with Pinker (http://video.google.com/videoplay?docid=3554279466299738997) TED talks • Chalking it up to the blank slate (http://www.ted.com/index.php/talks/ steven_pinker_chalks_it_up_to_the_blank_slate.html), 2003 • The stuff of thought (http://www.ted.com/index.php/talks/view/id/164), 2005 • A brief history of violence (http://www.ted.com/index.php/talks/view/id/163), 2007 Filmed Talks • The Better Angels of our Nature (http://richannel.org/steven-pinker-the-better-angels-of-our-nature), Royal Institution, November 2011 Debates • The Two Steves (http://www.edge.org/3rd_culture/pinker_rose/pinker_rose_p1.html) Debate with Steven Rose • The Science of Gender and Science (http://www.edge.org/3rd_culture/debate05/debate05_index.html) Debate with Elizabeth Spelke

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Frans de Waal

143

Frans de Waal Frans de Waal

Frans de Waal Born

October 29, 1948 's-Hertogenbosch, North Brabant, Netherlands

Occupation

Primatologist, Ethologist

Years active 1975–present

Franciscus Bernardus Maria "Frans" de Waal, PhD (born 29 October 1948) is a Dutch primatologist and ethologist. He is the Charles Howard Candler professor of Primate Behavior in the Emory University psychology department in Atlanta, Georgia, and director of the Living Links Center at the Yerkes National Primate Research Center and author of numerous books including Chimpanzee Politics and Our Inner Ape. His research centers on primate social behavior, including conflict resolution, cooperation, inequity aversion, and food-sharing. He is a Member of the United States National Academy of Sciences and the Royal Netherlands Academy of Sciences.

Early life and education De Waal was born in 's-Hertogenbosch. De Waal studied at the Dutch universities of Radboud University Nijmegen, University of Groningen, and Utrecht. In 1977, De Waal received his doctorate in biology from Utrecht University after training as a zoologist and ethologist with Professor Jan van Hooff, a well-known expert of emotional facial expressions in primates. His dissertation research concerned aggressive behavior and alliance formation in macaques.[1]

Career In 1975, De Waal began a six-year project on the world's largest captive colony of chimpanzees at the Arnhem Zoo. The study resulted in many scientific papers, and resulted in publication of his first book, Chimpanzee Politics, in 1982. This book offered the first description of primate behavior explicitly in terms of planned social strategies. De Waal was first to introduce the thinking of Machiavelli to primatology, leading to the label "Machiavellian Intelligence" that later became associated with it. In his writings, De Waal has never shied away from attributing emotions and intentions to his primates, and as such his work inspired the field of primate cognition that, three decades later, flourishes around themes of cooperation, altruism, and fairness. His early work also drew attention to deception and conflict resolution, nowadays two major areas of research. Initially, all of this was highly controversial. Thus, the label of "reconciliation", which De Waal introduced for

Frans de Waal reunions after fights, was questioned at first, but is now fully accepted with respect to animal behavior. Recently, De Waal's work has emphasized non-human animal empathy and even the origins of morality. His most widely cited paper, [2] written with his former student Stephanie Preston, concerns the evolutionary origin and neuroscience of empathy, not just in primates, but in mammals in general. De Waal's name is also associated with Bonobo, the "make love – not war" primate that he has made popular. But even his Bonobo studies are secondary to the larger goal of understanding what binds primate societies together rather than how competition structures them. Competition is not ignored in his work: the original focus of De Waal's research, before he was well known, was aggressive behavior and social dominance. Whereas his science focuses on the behavior of nonhuman primates (mostly chimpanzees, bonobos, macaques, and capuchin monkeys), his popular books have given De Waal worldwide visibility by relating the insights he has gained from monkey and ape behavior to human society. With his students, he has also worked on elephants, which are increasingly featured in his writings. His research into the innate capacity for empathy among primates has led De Waal to the conclusion that non-human great apes and humans are simply different types of apes, and that empathic and cooperative tendencies are continuous between these species. His belief is illustrated in the following quote from The Age of Empathy: “We start out postulating sharp boundaries, such as between humans and apes, or between apes and monkeys, but are in fact dealing with sand castles that lose much of their structure when the sea of knowledge washes over them. They turn into hills, leveled ever more, until we are back to where evolutionary theory always leads us: a gently sloping beach.” This is quite opposite to the view of some economists and anthropologists, who love to postulate differences between humans and other animals. However, recent work on prosocial tendencies in apes and monkeys supports De Waal's position. See, for example, the research of Felix Warneken, a psychologist at the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany. In 2011, De Waal and his co-workers were the first to report that chimpanzees given a free choice between helping only themselves or helping themselves plus a partner, prefer the latter. In fact, De Waal does not believe these tendencies to be restricted to humans and apes, but views empathy and sympathy as universal mammalian characteristics, a view that over the past decade has gained support from studies on rodents and other mammals, such as dogs. He and his students have also extensively worked on fairness, leading to a video that went viral on inequity aversion among capuchin monkeys. The most recent work in this area was the first demonstration that given a chance to play the Ultimatum Game, chimpanzees respond in the same way as children and human adults by preferring the equitable outcome.[3] In 1981, De Waal moved to the United States for a position at the Wisconsin National Primate Research Center, and took his current position at Emory and the Yerkes National Primate Research Center in 1991. He became an American citizen in 2008. His 2005 book Our Inner Ape examines human behavior through the eyes of a primatologist, using the behavior of common chimpanzees and bonobos as metaphors for human psychology. He also writes a column for Psychologie, a popular Dutch monthly magazine. It was announced on August 30, 2013 that De Waal would be appointed professor at Utrecht University.

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Frans de Waal

Quotes "The possibility that empathy resides in parts of the brain so ancient that we share them with rats should give pause to anyone comparing politicians with those poor, underestimated creatures." "I've argued that many of what philosophers call moral sentiments can be seen in other species. In chimpanzees and other animals, you see examples of sympathy, empathy, reciprocity, a willingness to follow social rules. Dogs are a good example of a species that have and obey social rules; that's why we like them so much, even though they're large carnivores." "To endow animals with human emotions has long been a scientific taboo. But if we do not, we risk missing something fundamental, about both animals and us."[4] "Atheism will need to be combined with something else, something more constructive than its opposition to religion, to be relevant to our lives. The only possibility is to embrace morality as natural to our species." "The Bonobo and the Atheist," 2013.

Awards • 2013 Doctor Honoris Causa, Utrecht University (Netherlands) • 2012 Ig Nobel Prize winner, in the Anatomy category • • • • • • • • • • • • •

2011 Discover magazine's "47 (all time) Great Minds of Science" 2011 Doctor Honoris Causa, Colgate University (USA) 2010 Ridder (knighted), Orde van de Nederlandse Leeuw (Order of the Netherlands' Lion) 2009 Medal, Società di Medicina & Scienze Naturali, Parma (Italy) 2009 Medal, Ariëns Kappers (Netherlands' Institute for Neuroscience) 2009 Doctor Honoris Causa, University for Humanistics (Netherlands) 2008 Fellow of the American Academy of Arts & Sciences (AAAS) 2007 Time Magazine 100 World’s Most Influential People Today 2005 Member of the American Philosophical Society (APS) 2005 Arthur W. Staats Award, American Psychological Foundation 2004 Member of the (US) National Academy of Sciences (NAS) 1993 Royal Netherlands Academy of Arts and Sciences (KNAW) 1989 Los Angeles Times Book Award for Peacemaking among Primates

Selected bibliography Books • The Bonobo and the Atheist [5], 2013. ISBN 978-0393073775 • The Age of Empathy: Nature's Lessons for a Kinder Society [6], 2009. ISBN 978-0-307-40776-4 (reviewed in American Scientist [7]) • Primates and Philosophers: How Morality Evolved [8], 2006. ISBN 0-691-12447-7 • Our Inner Ape. New York: Riverhead Books, 2005. ISBN 1-57322-312-3 • Animal Social Complexity: Intelligence, Culture, and Individualized Societies [9], Edited with Peter L. Tyack. Cambridge: Harvard University Press, 2003. ISBN 0-674-00929-0. • My Family Album, Thirty Years of Primate Photography 2003. • Tree of Origin: What Primate Behavior Can Tell Us about Human Social Evolution [10], Harvard University Press, 2001. ISBN 0-674-00460-4. • The Ape and the Sushi Master, Cultural reflections by a primatologist. New York: Basic Books, 2001. ISBN 0-465-04175-2

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Frans de Waal • Chimpanzee Politics: Power and Sex Among Apes [11] (25th Anniversary ed.). Baltimore, MD: JHU Press; 2007. ISBN 978-0-8018-8656-0. • Natural Conflict Resolution. 2000 (with Filippo Aureli) • Bonobo: The Forgotten Ape. Berkeley: University of California Press, 1997. ISBN 0-520-20535-9 (with Frans Lanting) • Good Natured: The Origins of Right and Wrong in Humans and Other Animals [12]. Cambridge: Harvard University Press, 1996. ISBN 0-674-35660-8 • Chimpanzee Cultures [13], Edited with Richard Wrangham, W.C. McGrew, and Paul Heltne. Foreword by Jane Goodall. Cambridge: Harvard University Press, 1994. ISBN 0-674-11662-3. • Peacemaking Among Primates [14]. Cambridge: Harvard University Press, 1989. ISBN 0-674-65920-1

Articles • 2010 "Opinion piece about God and morality in The New York Times" [15] • 2010 "Towards a bottom-up perspective on animal and human cognition" [16], Trends in Cognitive Sciences 201-207. May 2010 • 2009, "Darwin's last laugh" [17], Essay, Nature 460, 175 (9 July 2009) • 2008 "Putting the Altruism Back into Altruism: The Evolution of Empathy" [18], Annual Review of Psychology, Vol. 59: 279-300 • 2007, "Bonobos, Left & Right" [19] Skeptic, (8 August 2007). • 2006, "Self-recognition in an Asian elephant" [20], PNAS, vol 103, no 45, 17053-17057 • 2001, "Do Humans Alone 'Feel Your Pain'?" (Chronicle.com [21], October 26, 2001) • 1999, "The End of Nature Versus Nurture", Scientific American, vol 281, no 6, p 94-99 • 1995, "Bonobo Sex and Society The behavior of a close relative challenges assumptions about male supremacy in human evolution [22]", Scientific American, vol 272, no 3, p 82-88

References [1] [2] [3] [4]

Living Links Bio Page (http:/ / www. emory. edu/ LIVING_LINKS/ dewaal. html) http:/ / citeseerx. ist. psu. edu/ viewdoc/ download?doi=10. 1. 1. 120. 7176& rep=rep1& type=pdf http:/ / www. emory. edu/ LIVING_LINKS/ publications/ articles/ Proctor_etal_2013. pdf Frans de Waal (1997-07). "Are We in Anthropodenial?" (http:/ / discovermagazine. com/ 1997/ jul/ areweinanthropod1180). DISCOVER magazine, pp. 50–53. Retrieved 2011-07-30. [5] http:/ / books. wwnorton. com/ books/ detail-inside. aspx?ID=24800& CTYPE=G/ [6] http:/ / www. emory. edu/ LIVING_LINKS/ empathy/ [7] http:/ / www. americanscientist. org/ bookshelf/ pub/ fellow-feeling [8] http:/ / press. princeton. edu/ titles/ 8240. html [9] http:/ / www. hup. harvard. edu/ catalog. php?isbn=9780674018235 [10] http:/ / www. hup. harvard. edu/ catalog. php?isbn=9780674010048 [11] http:/ / books. google. com/ books?id=XsrhU2vV5PIC [12] http:/ / www. hup. harvard. edu/ catalog. php?isbn=9780674356610 [13] http:/ / www. hup. harvard. edu/ catalog. php?isbn=9780674116634 [14] http:/ / www. hup. harvard. edu/ catalog. php?isbn=9780674659216 [15] http:/ / opinionator. blogs. nytimes. com/ 2010/ 10/ 17/ morals-without-god/ [16] http:/ / www. sciencedirect. com/ science?_ob=ArticleURL& _udi=B6VH9-4YRVY96-4& _user=10& _coverDate=05%2F31%2F2010& _rdoc=6& _fmt=high& _orig=browse& _srch=doc-info%28%23toc%236061%232010%23999859994%231931692%23FLA%23display%23Volume%29& _cdi=6061& _sort=d& _docanchor=& _ct=9& _acct=C000050221& _version=1& _urlVersion=0& _userid=10& md5=568c2e04758a0b1691b43073e3a8ff91 [17] http:/ / www. emory. edu/ LIVING_LINKS/ Nature_essay. htm [18] http:/ / arjournals. annualreviews. org/ doi/ abs/ 10. 1146/ annurev. psych. 59. 103006. 093625 [19] http:/ / www. skeptic. com/ eskeptic/ 07-08-08. html [20] http:/ / www. pnas. org/ cgi/ content/ abstract/ 103/ 45/ 17053?maxtoshow=& HITS=10& hits=10& RESULTFORMAT=& fulltext=frans+ waal& searchid=1& FIRSTINDEX=0& resourcetype=HWCIT [21] http:/ / chronicle. com/ free/ v48/ i09/ 09b00701. htm

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Frans de Waal [22] http:/ / songweaver. com/ info/ bonobos. html

External links • Frans de Waal on Big Think about God and morality. (http://www.youtube.com/watch?v=sEQuIDqY6Cc) • Frans de Waal: Do animals have morals (http://www.ted.com/talks/frans_de_waal_do_animals_have_morals. html) • Frans de Waal's official FaceBook site (https://www.facebook.com/pages/Frans-de-Waal-Public-Page/ 99206759699) • De Waal on The Colbert Report, in 2008 (http://www.colbertnation.com/the-colbert-report-videos/148996/ january-30-2008/frans-de-waal) • Interview with Frans de Waal about empathy (http://www.youtube.com/watch?v=UONxT4Tb3C0) • Interview with De Waal on BigThink about morality (http://bigthink.com/ideas/15501) • De Waal, Wrangham, and Pinker discuss warfare (http://www.youtube.com/watch?v=ru8tHB6yZjk) • TED talk by De Waal on the origins of morality (http://www.ted.com/talks/ frans_de_waal_do_animals_have_morals.html) (Includes clips of experiments.) • Living Links Center (http://www.emory.edu/LIVING_LINKS/index.html), currently directed by Frans de Waal • Emory.edu (http://www.psychology.emory.edu/nab/dewaal/) - 'Frans B. M. de Waal, C. H. Candler Professor of Primate Behavior', Emory University faculty homepage • 92Y.org 'Talking Primates with Dr. Frans de Waal' (August 25, 2005 Blog) (http://blog.92y.org/index.php/ weblog/item/dr_frans_de_waal/) • AmericanScientist.org (http://www.americanscientist.org/template/ScientistNightstandTypeDetail/assetid/ 27334) - 'The Bookshelf talks with Frans de Waal', American Scientist (2001) • ITConversations.com (http://www.itconversations.com/shows/detail231.html) - 'Frans de Waal, Comparative Primatologist' (includes mp3 audio clip; October 21, 2004) • PaulaGordon.com (http://www.paulagordon.com/shows/waal/) - 'Natural Goodness', Paula Gordon • PNAS.org (http://www.pnas.org/cgi/content/full/102/32/11137) - 'Profile of Frans B. M. de Waal', Regina Nuzz, Proceedings of the National Academy of Sciences (PNAS) • De Waal's editorial contribution to social psychology in Greater Good magazine (http://greatergoodmag.org) • BookTalk.org: online reading group discussion of Frans de Waal's "Our Inner Ape" (http://www.booktalk.org/ our-inner-ape-by-frans-de-waal-f86.html) • BookTalk.org: book discussion of Frans de Waal's "Primates and Philosophers: How Morality Evolved" (http:// www.booktalk.org/primates-and-philosophers-how-morality-evolved-by-frans-de-waal-f110.html) • Video (with mp3 available) of discussion with De Waal about aspects of primate behavior (http://bloggingheads. tv/diavlogs/19818) on Bloggingheads.tv • "On Atheists and Bonobos - A conversation with Frans de Waal" (http://www.ideasroadshow.com/issues/ frans-de-waal-2013-07-26), Ideas Roadshow, 2013

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Richard Lewontin

148

Richard Lewontin Richard Lewontin Born

29 March 1929 New York City, New York, U.S.

Nationality

American

Fields

Genetics Evolutionary biology Population genetics

Institutions

Harvard University North Carolina State University University of Rochester University of Chicago Columbia University

Alma mater

Harvard University Columbia University

Thesis

The Effects of Population Density and Composition on Viability in Drosophila melanogaster

[1]

(1955)

Academic advisors Theodosius Dobzhansky[citation needed] Doctoral students

Joseph Felsenstein Jerry Coyne Martin Kreitman

Known for

Evolutionary biology Population genetics

Richard Charles "Dick" Lewontin (born March 29, 1929) is an American evolutionary biologist, geneticist, academic and social commentator. A leader in developing the mathematical basis of population genetics and evolutionary theory, he pioneered the application of techniques from molecular biology, such as gel electrophoresis, to questions of genetic variation and evolution. In a pair of 1966 papers co-authored with J.L. Hubby in the journal Genetics, Lewontin helped set the stage for the modern field of molecular evolution. In 1979 he and Stephen Jay Gould introduced the term "spandrel" into evolutionary theory. From 1973 to 1998, he held an endowed chair in zoology and biology at Harvard University, and since 2003 has been a research professor there. Lewontin strongly opposes genetic determinism, especially as allegedly propounded by researchers in behavioral genetics.

Early life and education Lewontin was born in New York City to parents descended from late 19th-century Eastern European Jewish immigrants. He attended Forest Hills High School and the École Libre des Hautes Études in New York. In 1951 he graduated from Harvard College (BS, biology). In 1952, Lewontin received a master's degree in mathematical statistics, followed by a doctorate in zoology in 1954, both from Columbia University, where he was a student of Theodosius Dobzhansky. He held faculty positions at North Carolina State University, the University of Rochester, and the University of Chicago. In 1973 Lewontin was appointed as Alexander Agassiz Professor of Zoology and Professor of Biology at Harvard University, holding the position until 1998.

Richard Lewontin

Career Work in population genetics Lewontin has worked in both theoretical and experimental population genetics. A hallmark of his work has been an interest in new technology. He was the first person to do a computer simulation of the behavior of a single locus (previous simulation work having been of models with multiple loci). In 1960 he and Ken-Ichi Kojima were the first population geneticists to give the equations for change of haplotype frequencies with interacting natural selection at two loci.[2] This set off a wave of theoretical work on two-locus selection in the 1960s and 1970s. Their paper gave a theoretical derivation of the equilibria expected, and also investigated the dynamics of the model by computer iteration. Lewontin later introduced the D' measure of linkage disequilibrium.[3] (An achievement that he would be less happy to claim is that he introduced the name "linkage disequilibrium" itself, one about which many population geneticists have been unenthusiastic). In 1966, he and Jack Hubby published a paper that revolutionized population genetics. They used protein gel electrophoresis to survey dozens of loci in Drosophila pseudoobscura, and reported that a large fraction of the loci were polymorphic, and that at the average locus there was about a 15% chance that the individual was heterozygous. (Harry Harris reported similar results for humans at about the same time). Previous work with gel electrophoresis had been reports of variation in single loci and did not give any sense of how common variation was. Lewontin and Hubby's paper also discussed the possible explanation of the high levels of variability by either balancing selection or neutral mutation. Although they did not commit themselves to advocating neutrality, this was the first clear statement of the neutral theory for levels of variability within species. Lewontin and Hubby's paper had great impact—the discovery of high levels of molecular variability gave population geneticists ample material to work on, and gave them access to variation at single loci. The possible theoretical explanations of this rampant polymorphism became the focus of most population genetics work thereafter. Martin Kreitman was later to do a pioneering survey of population-level variability in DNA sequences while a Ph.D. student in Lewontin's lab.

Work on human genetic diversity In a landmark paper, in 1972 Lewontin identified that most of the variation (80–85%) within human populations is found within local geographic groups and differences attributable to traditional "race" groups are a minor part of human genetic variability (1–15%). He argued that the probability of racial misclassification of an individual based on variation in a single genetic locus is approximately 30% and the misclassification probability becomes close to zero if enough loci are studied. In a 2003 paper, A.W.F. Edwards criticized Lewontin's conclusion that race is an invalid taxonomic construct, terming it Lewontin's fallacy.

Critique of orthodox evolutionary biology In 1975, when E. O. Wilson's book Sociobiology proposed evolutionary explanations for human social behaviors, a number of biologists responded negatively, including Lewontin, Stephen Jay Gould, Ruth Hubbard, and others.[4] Lewontin and his late Harvard colleague Stephen Jay Gould introduced the term spandrel to evolutionary biology, inspired by the architectural term "spandrel", in an influential 1979 paper, "The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme." "Spandrels" were described as features of an organism that exist as a necessary consequence of other (perhaps adaptive) features, but do not directly improve fitness (and thus are not necessarily adaptive). The relative frequency of spandrels versus adaptations continues to stir controversy in evolutionary biology. Lewontin was an early proponent of a hierarchy of levels of selection in his article, "The Units of Selection". He has been a major influence on philosophers of biology, notably William C. Wimsatt (who taught with Lewontin and Richard Levins at the University of Chicago), Robert Brandon and Elisabeth Lloyd (who studied with Lewontin as graduate students), Philip Kitcher, and Elliott Sober. Lewontin briefly argued for the historical nature of biological

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Richard Lewontin causality in "Is Nature Probable or Capricious?"[citation needed] In "Organism and Environment" in Scientia, and in more popular form in the last chapter of Biology as Ideology, Lewontin argued that while traditional Darwinism has portrayed the organism as a passive recipient of environmental influences, a correct understanding should emphasize the organism as an active constructor of its own environment. Niches are not pre-formed, empty receptacles into which organisms are inserted, but are defined and created by organisms. The organism-environment relationship is reciprocal and dialectical. M.W. Feldman, K.N. Laland, and F.J. Odling-Smee, among others, have developed Lewontin's conception in more detailed models.[citation needed]

In the adaptationist view of evolution, the organism is a function of both the organism and environment, while the environment is only a function of itself. The environment is seen as autonomous and unshaped by the organism. Lewontin instead believed in a constructivist view, in which the organism is a function of the organism and environment, with the environment being a function of the organism and environment as well. This means that the organism shapes the environment as the environment shapes the organism. The organism shapes the environment for future generations.[5] Lewontin has long been a critic of traditional neo-Darwinian approaches to adaptation. In his article "Adaptation" in the Italian Enciclopedia Einaudi, and in a toned-down version in Scientific American, he emphasized the need to give an engineering characterization of adaptation separate from measurement of number of offspring, rather than simply assuming organs or organisms are at adaptive optima. Lewontin has said that his more general, technical criticism of adaptationism grew out of his recognition that the fallacies of sociobiology reflect fundamentally flawed assumptions of adaptiveness of all traits in much of the modern evolutionary synthesis. Lewontin accused neo-Darwinists of telling Just So Stories when they try to show how natural selection explains such novelties as long-necked giraffes.[6] Sociobiology and evolutionary psychology Along with others, such as Gould, Lewontin has been a persistent critic of some themes in neo-Darwinism. Specifically, he has criticised proponents of sociobiology and evolutionary psychology such as Edward O. Wilson and Richard Dawkins, who attempt to explain animal behaviour and social structures in terms of evolutionary advantage or strategy. He and others criticize this approach when applied to humans, as he sees it as genetic determinism. In his writing, Lewontin suggests a more nuanced view of evolution is needed, which requires a more careful understanding of the context of the whole organism as well as the environment.[citation needed] Such concerns about what he views as the oversimplification of genetics has led Lewontin to be a frequent participant in debates, and an active life as a public intellectual. He has lectured widely to promote his views on evolutionary biology and science. In books such as Not in Our Genes (co-authored with Steven Rose and Leon J. Kamin) and numerous articles, Lewontin has questioned much of the claimed heritability of human behavioral traits, such as intelligence as measured by IQ tests.[citation needed] Some academics have criticized him for rejecting sociobiology for non-scientific reasons. Edward Wilson (1995) suggested that Lewontin's political beliefs affected his scientific view. Lewontin has at times identified himself as Marxist, and admitted that his ideological views have affected his scientific work (Levins and Lewontin 1985). Others such as Kitcher (1985) have countered that Lewontin's criticisms of sociobiology are genuine scientific concerns about the discipline. He wrote that attacking Lewontin's motives amounts to an ad hominem argument.[citation needed]

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Richard Lewontin Agribusiness Lewontin has written on the economics of agribusiness. He has contended that hybrid corn was developed and propagated not because of its superior quality, but because it allowed agribusiness corporations to force farmers to buy new seed each year rather than plant seed produced by their previous crop of corn. Lewontin testified in an unsuccessful suit in California challenging the state's financing of research to develop automatic tomato pickers. This favored the profits of agribusiness over the employment of farm workers.[citation needed]

Personal life As of 2003, Lewontin was the Alexander Agassiz Research Professor at Harvard. He has worked with and had great influence on many philosophers of biology, including William C. Wimsatt, Elliott Sober, Philip Kitcher, Elisabeth Lloyd, Peter Godfrey-Smith, Sahotra Sarkar, and Robert Brandon, often inviting them to work in his lab.

Recognition • 1961: Fulbright Fellowship • 1961: National Science Foundation Senior Postdoctoral Fellow • 1970s: Membership of the National Academy of Sciences (later resigned) • 1994: Sewall Wright Award from the American Society of Naturalists

Bibliography • Lewontin, R. C.; Kojima, K. (Dec., 1960). "The Evolutionary Dynamics of Complex Polymorphisms" [7]. Evolution (Society for the Study of Evolution) 14 (4): 458–472. doi:10.2307/2405995 [8]. JSTOR 2405995 [7]. • Richard C. Lewontin (Jan., 1966). "Is Nature Probable or Capricious?" [9]. BioScience (University of California Press) 16 (1, Logic in Biological Investigation): 25–27. doi:10.2307/1293548 [10]. JSTOR 1293548 [9]. • Lewontin, R. C. (1970). "The Units of Selection". Annual Review of Ecology and Systematics 1: 1–0. doi:10.1146/annurev.es.01.110170.000245 [11]. • "The Apportionment of Human Diversity," Evolutionary Biology, vol. 6 (1972) pp. 391–398. • Lewontin, R. C. (1974). The genetic basis of evolutionary change. New York: Columbia University Press. ISBN 0-231-03392-3. • "Adattamento," Enciclopedia Einaudi, (1977) vol. 1, 198–214. • "Adaptation," Scientific American, vol. 239, (1978) 212–228. • Gould, S. J.; Lewontin, R. C. (1979). "The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme". Proceedings of the Royal Society B: Biological Sciences 205 (1161): 581. doi:10.1098/rspb.1979.0086 [12]. • Lewontin, R. C. (1995). Human diversity (2nd ed.). New York: Scientific American Library. ISBN 0-7167-6013-4. • "The Organism as Subject and Object of Evolution," Scientia vol. 188 (1983) 65–82. • Not in Our Genes: Biology, Ideology and Human Nature (with Steven Rose and Leon J. Kamin) (1984) ISBN 0-394-72888-2 • The Dialectical Biologist (with Richard Levins), Harvard University Press (1985) ISBN 0-674-20283-X • Biology as Ideology: The Doctrine of DNA (1991) ISBN 0-06-097519-9 • The Triple Helix: Gene, Organism, and Environment, Harvard University Press (2000) ISBN 0-674-00159-1 • It Ain't Necessarily So: The Dream of the Human Genome and Other Illusions, New York Review of Books (2000) • Biology Under The Influence: Dialectical Essays on the Coevolution of Nature and Society (with Richard Levins), (2007)

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References [1] [2] [3] [4]

http:/ / search. proquest. com/ docview/ 301991815 Lewontin, R. C. and K. Kojima (1960). "The evolutionary dynamics of complex polymorphisms". Evolution 14: 458-472. Lewontin, R. C. (1964). "The interaction of selection and linkage. I. General considerations; heterotic models". Genetics 49: 49-67. Elizabeth Allen et al., 1975, "Against 'Sociobiology'" (http:/ / www. nybooks. com/ articles/ archives/ 1975/ nov/ 13/ against-sociobiology/ ), The New York Review of Books, 13 November 1975 [5] Richard Levins and Richard Lewontin, The Dialectical Biologist, 1985 [6] "Science Contra Darwin", Newsweek, 8 April 1985, p.80 [7] http:/ / www. jstor. org/ stable/ 2405995 [8] http:/ / dx. doi. org/ 10. 2307%2F2405995 [9] http:/ / www. jstor. org/ stable/ 1293548 [10] http:/ / dx. doi. org/ 10. 2307%2F1293548 [11] http:/ / dx. doi. org/ 10. 1146%2Fannurev. es. 01. 110170. 000245 [12] http:/ / dx. doi. org/ 10. 1098%2Frspb. 1979. 0086

Further reading • Philip Kitcher (1985). Vaulting Ambition : Sociobiology and the Quest for Human Nature. MIT Press. ISBN 0-262-11109-8. • Steven Pinker (2002). The Blank Slate: The Denial of Human Nature in Modern Intellectual Life. Penguin Press. • Rama S. Singh, Costas Krimbas, Diane Paul, and John Beatty (2001). Thinking about Evolution. Cambridge University Press. - a two volume Festschrift for Lewontin with a full bibliography • Edward O. Wilson (1995). "Science and ideology" (http://lrainc.com/swtaboo/taboos/wilson01.html). Academic Questions 8. • Richard Lewontin (2004). The Triple Helix: Gene, Organism and Environment (http://video.google.com/ videoplay?docid=7320126698292379909). MIT Press.

External links • Quotations related to Richard Lewontin at Wikiquote • an interview given at Berkeley in 2003 (http://globetrotter.berkeley.edu/people3/Lewontin/lewontin-con0. html) • a profile and bibliography (http://authors.library.caltech.edu/5456/1/hrst.mit.edu/hrs/evolution/public/ profiles/lewontin.html) • Gene, Organism and Environment: Bad Metaphors and Good Biology - RealAudio stream of Hitchcock lecture on UCTV (http://webcast.ucsd.edu:8080/ramgen/UCSD_TV/8393.rm) • The Concept of Race: The Confusion of Social and Biological Reality - RealAudio stream of Hitchcock lecture on UCTV (http://webcast.ucsd.edu:8080/ramgen/UCSD_TV/8456.rm) • Internalism and Externalism in Biology (http://internalism.blip.tv/file/812402/), lecture delivered at Harvard university on December 13, 2007.

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Steven Rose

Steven Rose Steven Peter Russell Rose (born 4 July 1938 in London, United Kingdom)[1] is a Professor of Biology and Neurobiology at the Open University and University of London.

Life Rose studied biochemistry at King's College, Cambridge, and neurobiology at Cambridge and the Institute of Psychiatry, King's College London.[2] When he was appointed to the professorship of biology at the newly instituted Open University in 1969 he was Britain's youngest full professor and chair of department. At the Open University he established the Brain Research Group, within which he and his colleagues focussed on the biological processes involved in memory formation and treatments for Alzheimer's Disease on which he has published some 300 research papers and reviews. He has written several popular science books and regularly writes for The Guardian newspaper. From 1999 to 2002, he gave public lectures as Professor of Physic at Gresham College, London. His work has won him numerous medals and prizes including the Biochemical Society medal for communication in science and the prestigious Edinburgh Medal. His book The Making of Memory won the Science Book Prize in 1993.[3] In 2012 the British Neuroscience Association gave him a lifetime award for "Outstanding contributions to neuroscience." His younger brother is Nikolas Rose, professor of sociology at the LSE. He is married to the sociologist Hilary Rose with whom he shared the Gresham professorship, and with whom he has written and edited a number of books including Alas Poor Darwin: Arguments against Evolutionary Psychology and, in 2012, Genes, Cells and Brains: the Promethean promises of the new biology, (Verso) described by Guardian reviewer Steven Poole as 'fascinating, lucid and angry' with a 'lethally impressive hit ratio.' Novelist Margaret Atwood tweeted that it was on her 'must read' list. Growing up as an Orthodox Jew, Rose says that he decided to become an atheist when he was eight years old.

Research and scientific controversies With Richard Lewontin and Leon Kamin, Rose championed the "radical science movement." The three criticized sociobiology, evolutionary psychology, and adaptationism, most prominently in the book Not in Our Genes (1984), laying out their opposition to Sociobiology (E. O. Wilson, 1975), The Selfish Gene (Richard Dawkins, 1976), and other works promoting an evolutionary explanation for human social behaviour. Not in Our Genes described Dawkins as "the most reductionist of sociobiologists". In retort, Dawkins wrote that the book practices reductionism by distorting arguments in terms of genetics to "an idiotic travesty (that the properties of a complex whole are simply the sum of those same properties in the parts)", and accused the authors of giving "ideology priority over truth". Rose replied in the 2nd edition of his book Lifelines. Rose wrote further works in this area; in 2000 he jointly edited with the sociologist Hilary Rose, a critique of evolutionary psychology: Alas, Poor Darwin: Arguments Against Evolutionary Psychology. In 2006 he wrote a paper dismissing classical heritability estimates as useful scientific measures in respect of human populations especially in the context of IQ. Rose was for several years a regular panelist on BBC Radio 4's ethics debating series The Moral Maze. Rose is a Distinguished Supporter of the British Humanist Association. He is presently a member of the Nuffield Council on Bioethics Working Party on Novel Neurotechnologies.[4]

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Steven Rose

Political views The Guardian described Rose as a "polemicist on the left." His friend and collaborator Patrick Bateson wrote that Rose "may be the last of the Marxist radical scientists," adding that "Steven is not always right; but he has been very brave in some of the things he has said." Steven Rose, together with sociologist Hilary Rose has been instrumental in calling to boycott Israeli academic institutions for as long as Israel continues its illegal occupation of the Palestinian Territories, on the grounds of Israel academics' close relationship with the IDF. An open letter initiated by Steven and Hilary Rose, and also signed by 123 other academics was published in The Guardian on 6 April 2002. In 2004 he was one of the founding members of the British Committee for Universities of Palestine.[5]

Bibliography • • • •

Chemical and Biological Warfare 1968, Chambers Harrap Publishers, ISBN 024559485X Science and Society with Hilary Rose, Penguin, 1969 The Conscious Brain 1973, ISBN 0-394-46066-9 Radicalisation of Science with Hilary Rose, 1976, Macmillan, ISBN 0333211413

• Political Economy of Science: Ideology of/in the Natural Science Editor with Hilary Rose, 1976, Macmillan, ISBN 0333211383 • Towards a Liberatory Biology (Editor) 1981, Allison & Busby, ISBN 0850314259 • Against Biological Determinism (Editor), 1982, Schocken, ISBN 0805281126 • Not in Our Genes (With Richard Lewontin & Leon Kamin) 1984, ISBN 0-394-72888-2 • No Fire, No Thunder: Threat of Chemical and Biological Weapons with Sean Murphy and Alistair Hay, Pluto, ISBN 0861047389 • The Chemistry of Life 1991 (first published in 1966), ISBN 0-14-027273-9 • The Making Of Memory 1992, ISBN 0-593-01990-3 • Alas, Poor Darwin: Arguments against Evolutionary Psychology with Hilary Rose, 2000, ISBN 0-609-60513-5 • Lifelines 2005, ISBN 0-09-946863-8 • The 21st Century Brain 2005, ISBN 0-224-06254-9 • The Future of the Brain: The Promise and Perils of Tomorrow's Neuroscience 2005, ISBN 0-19-515420-7 • Genes, Cells and Brains: Bioscience's Promethean Promises with Hilary Rose, 2012, Verso, ISBN 1844678814

References [1] ‘ROSE, Prof. Steven Peter Russell’, Who's Who 2013, A & C Black, an imprint of Bloomsbury Publishing plc, 2013; online edn, Oxford University Press, Dec 2012 ; online edn, Nov 2012 accessed 6 Aug 2013 (http:/ / www. ukwhoswho. com/ view/ article/ oupww/ whoswho/ U33166) [2] Biography at The Moral Maze. (http:/ / www. bbc. co. uk/ radio4/ religion/ steven_rose. shtml) [3] http:/ / www. stevenroseonline. net/ SPRR/ Long_bio_. html [4] http:/ / www. nuffieldbioethics. org/ neurotechnology/ neurotechnology-about-working-party [5] http:/ / www. bricup. org. uk/ December09tour. html

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External links • • • • • •

Steven Rose Online (http://stevenroseonline.net/SPRR/Welcome.html) Profile on The Third Culture (http://www.edge.org/3rd_culture/bios/rose.html) Debate with (http://www.edge.org/3rd_culture/pinker_rose/pinker_rose_p1.html) Steven Pinker Biography at Counterbalance (http://www.counterbalance.net/bio/srose-body.html) Steven Rose articles at The Guardian (http://www.guardian.co.uk/profile/stevenrose?INTCMP=SRCH) Poole, Steven (19 December 2012). "Genes, Cells and Brains by Hilary Rose and Steven Rose - review" (http:// www.guardian.co.uk/books/2012/dec/19/genes-cells-and-brains-hilary-steven-rose-review). The Guardian.

Leon Kamin Leon J. Kamin (born December 29, 1927 in Taunton, Massachusetts) is an American psychologist known for his contributions to learning theory and his critique of estimates of the heritability of IQ. He studied under Richard Solomon at Harvard and discovered several important facts about conditioning, including the "Kamin Effect" and the "blocking effect". Kamin was blacklisted during the McCarthy era (Kamin, 2005) and had to find employment in Canada, where he chaired the Psychology Department at McMaster University in Ontario, Canada (1957–68). When he was removed from the blacklist in 1968, he returned to the US and chaired Princeton University's Department of Psychology and later the Psychology Department at Northeastern University in Boston, MA. Kamin's most well-known contribution to learning theory was his discovery and analysis of the "blocking effect" (1969). He showed that conditioning an animal to associate a salient conditioned stimulus (CSb), such as a bright light, with a salient unconditioned stimulus (US), like a shock, is "blocked" when CSb is presented simultaneously with another conditioned stimulus (CSa) that was already conditioned to the US. (Kamin used rats in most of his research, but the effect has been found in many animals). The blocking effect is one of the hallmark effects in the study of associative learning in animals, including humans. Kamin has long opposed the idea that significant personal traits are largely heritable. He became skeptical of the claims of Cyril Burt regarding the heritability of IQ, and published his findings in a 1974 book The Science and Politics of IQ. He co-authored the controversial book Not in Our Genes (1984) with geneticist Richard Lewontin and neurobiologist Steven Rose. This book, championing the "radical science movement," attacked sociobiology and evolutionary psychology. Kamin is known in some circles for his position that the heritability of IQ could be zero (Mackintosh, 1998) He is honorary professor of psychology at the University of Cape Town in South Africa.

Criticism In Race Differences in Intelligence, Loehlin, Lindzey & Spuhler wrote (p. 85): Kamin's more radical assertion of zero heritability, if substantiated, might not render the present book entirely meaningless, but it would certainly require a considerable revision of its language and point of view. In Appendix H we have taken a second look in the light of Kamin's critique at some of the data that provide the main focus for his misgivings, namely the studies of adoptive families and separated identical twins. We do not find these data to be quite as fragile as does Kamin, and we find Kamin's analysis to suffer from a number of statistical and logical problems. and also (p. 299): On the whole, then, although Kamin's critique emphasizes some methodological points worthy of attention in conducting and evaluating studies in this area, it does not, in our view, constitute an unbiased survey of the data, and it suffers from enough logical and statistical difficulties that Kamin's

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Leon Kamin "reasonably prudent man" will want to think twice before accepting his conclusions.

Bibliography • The Science and Politics of IQ (1974) • Hans Jürgen Eysenck and Leon J. Kamin (1981). Intelligence: the Battle for the Mind. MacMillan. ISBN 0-330-26399-4. • Richard Lewontin, Steven Rose and Leon J. Kamin (1984). Not in Our Genes.

References • Kamin, L. J. (1969). Predictability, surprise, attention, and conditioning. In B. A. Campbell & R. M . Church (Eds.), Punishment and aversive behavior (pp. 279–296). New York: Appleton-Century-Crofts. • Kamin, L.J. (2005). Letter to the Editor, New York Review of Books, May 26. • Mackintosh, N. (1998). IQ and Human Intelligence. Oxford: University Press. pp. 78–79. • Loehlin, Lindzey & Spuhler (Freeman, 1975). Race Differences in Intelligence (ISBN 0-7167-0754-3)

External links • Indiana University profile [1]

References [1] http:/ / www. indiana. edu/ ~intell/ kamin. shtml

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Stephen Jay Gould

157

Stephen Jay Gould Stephen Jay Gould

Born

September 10, 1941 Bayside, New York, United States

Died

May 20, 2002 (aged 60) Manhattan, New York, United States

Nationality

American

Fields

Paleontology, Evolutionary biology, History of Science

Institutions

Harvard University, American Museum of Natural History, New York University

Alma mater

Antioch College, Columbia University

Known for

Punctuated equilibrium, Non-overlapping magisteria

Notable awards Linnean Society of London's Darwin–Wallace Medal (2008) Paleontological Society Medal (2002) Charles Schuchert Award (1975) Phi Beta Kappa Award in Science (twice – 1983, 1990) MacArthur Fellowship National Book Award National Book Critics Circle Award Spouse

Deborah Lee (1965–?; divorced; 2 children) Rhonda Roland Shearer (1995–2002; his death; 2 stepchildren) Signature

Stephen Jay Gould (September 10, 1941 – May 20, 2002) was an American paleontologist, evolutionary biologist, and historian of science. He was also one of the most influential and widely read writers of popular science of his generation. Gould spent most of his career teaching at Harvard University and working at the American Museum of

Stephen Jay Gould Natural History in New York. In the later years of his life, Gould also taught biology and evolution at New York University. Gould's most significant contribution to evolutionary biology was the theory of punctuated equilibrium, which he developed with Niles Eldredge in 1972.[1] The theory proposes that most evolution is marked by long periods of evolutionary stability, which is punctuated by rare instances of branching evolution. The theory was contrasted against phyletic gradualism, the popular idea that evolutionary change is marked by a pattern of smooth and continuous change in the fossil record. Most of Gould's empirical research was based on the land snail genera Poecilozonites and Cerion. He also contributed to evolutionary developmental biology, and has received wide praise for his book Ontogeny and Phylogeny. In evolutionary theory he opposed strict selectionism, sociobiology as applied to humans, and evolutionary psychology. He campaigned against creationism and proposed that science and religion should be considered two distinct fields (or "magisteria") whose authorities do not overlap.[2] Gould was known by the general public mainly from his 300 popular essays in the magazine Natural History, and his books written for a non-specialist audience. In April 2000, the US Library of Congress named him a "Living Legend".

Biography Stephen Jay Gould was born and raised in the community of Bayside, a neighborhood of the northeastern section of Queens in New York City. His father, Leonard, was a court stenographer, and his mother, Eleanor, was an artist whose parents were Jewish immigrants living and working in the city's Garment District. When Gould was five years old, his father took him to the Hall of Dinosaurs in the American Museum of Natural History, where he first encountered Tyrannosaurus rex. "I had no idea there were such things—I was awestruck," Gould once recalled.[3] It was in that moment that he decided to become a paleontologist. Raised in a secular Jewish home, Gould did not formally practice religion and preferred to be called an agnostic. Though he "had been brought up by a Marxist father", he stated that his father's politics were "very different" from his own.[4] In describing his own political views, he has said they "tend to the left of center".[5] According to Gould the most influential political books he read were C. Wright Mills' The Power Elite and the political writings of Noam Chomsky. While attending Antioch College in the early 1960s, Gould was active in the civil rights movement and often campaigned for social justice. When he attended the University of Leeds as a visiting undergraduate, he organized weekly demonstrations outside a Bradford dance hall which refused to admit Blacks. Gould continued these demonstrations until the policy was revoked.[6] Throughout his career and writings, he spoke out against cultural oppression in all its forms, especially what he saw as the pseudoscience used in the service of racism and sexism.[7] Interspersed throughout his scientific essays for Natural History magazine, Gould frequently referred to his nonscientific interests and pastimes. As a boy he collected baseball cards and remained a New York Yankees fan throughout his life. As an adult he was fond of science fiction movies, but often lamented their mediocrity (not just in their presentation of science, but in their storytelling as well).[8] His other interests included singing in the Boston Cecilia, and he was a great aficionado of Gilbert and Sullivan operettas. He collected rare antiquarian books and textbooks. He often traveled to Europe, and spoke French, German, Russian, and Italian. He admired Renaissance architecture. When discussing the Judeo-Christian tradition, he usually referred to it simply as "Moses". He sometimes alluded ruefully to his tendency to put on weight.[9]

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Marriage and family Gould was married twice. His first marriage was to artist Deborah Lee on October 3, 1965. Gould met Lee while they were students together at Antioch College. They had two sons, Jesse and Ethan.[10] His second marriage, in 1995, was to artist and sculptor Rhonda Roland Shearer, who is the mother of two children, Jade and London Allen, stepchildren of Gould.

First bout with cancer In July 1982, Gould was diagnosed with peritoneal mesothelioma, a deadly form of cancer affecting the abdominal lining and frequently found in people who have been exposed to asbestos or rock dust. After a difficult two-year recovery, Gould published a column for Discover magazine, entitled, "The Median Isn't the Message", which discusses his reaction to discovering that mesothelioma patients had a median lifespan of only eight months after diagnosis.[11] He then describes the true significance behind this number, and his relief upon realizing that statistical averages are just useful abstractions, and do not encompass the full range of variation. The median is the halfway point, which means that 50% of patients will die before eight months, but the other half will live longer, potentially much longer. He then needed to determine where his personal characteristics placed him within this range. Considering that the cancer was detected early, the fact he was young, optimistic, and had the best treatments available, Gould figured that he should be in the favorable half of the upper statistical range. After an experimental treatment of radiation, chemotherapy, and surgery, Gould made a full recovery, and his column became a source of comfort for many cancer patients. Gould was also an advocate of medical marijuana. During his bout with cancer, he smoked the illegal drug to alleviate the nausea associated with his medical treatments. According to Gould, his use of marijuana had a "most important effect" on his eventual recovery.[12] In 1998, he testified in the case of Jim Wakeford, a Canadian medical-marijuana user and activist.

Final illness and death Gould survived for 20 years until another cancer ended his life. Gould died on May 20, 2002, from a metastatic adenocarcinoma of the lung, a form of cancer which had spread to his brain.[13] This cancer was unrelated to his abdominal cancer. He died in his home "in a bed set up in the library of his SoHo loft, surrounded by his wife Rhonda, his mother Eleanor, and the many books he loved."[14]

Scientific career Gould began his higher education at Antioch College, graduating with a double major in geology and philosophy in 1963.[15] During this time, he also studied at the University of Leeds in the United Kingdom.[16] After completing graduate work at Columbia University in 1967 under the guidance of Norman Newell, he was immediately hired by Harvard University where he worked until the end of his life (1967–2002). In 1973, Harvard promoted him to Professor of Geology and Curator of Invertebrate paleontology at the institution's Museum of Comparative Zoology; he very often described himself as a taxonomist. In 1982 Harvard awarded him the title of Alexander Agassiz Professor of Zoology. The following year, 1983, he was awarded fellowship of the American Association for the Advancement of Science, where he later served as president (1999–2001). The AAAS news release cited his "numerous contributions to both scientific progress and the public understanding of science". He also served as president of the Paleontological Society (1985–1986) and of the Society for the Study of Evolution (1990–1991). In 1989 Gould was elected into the body of the National Academy of Sciences. Through 1996–2002 Gould was Vincent Astor Visiting Research Professor of Biology at New York University. In 2001, the American Humanist Association named him the Humanist of the Year for his lifetime of work. In 2008, he was posthumously awarded

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Stephen Jay Gould the Darwin-Wallace Medal, along with 12 other recipients. (Until 2008 this medal had been awarded every 50 years by the Linnean Society of London.[17])

Punctuated equilibrium Early in his career, Gould and Niles Eldredge developed the theory of punctuated equilibrium, according to which evolutionary change occurs relatively rapidly, alternating with longer periods of relative evolutionary stability. Although Gould suggested the term itself, the basic concept was first presented in Eldredge's doctoral dissertation on Devonian trilobites and in an article published the previous year on allopatric speciation.[18] According to Gould, punctuated equilibrium revised a key pillar "in the central logic of Darwinian theory". Some evolutionary biologists have argued that while punctuated equilibrium was "of great interest to biology",[19] it merely modified neo-Darwinism in a manner that was fully compatible with what had been known before. For example, George Gaylord Simpson, in Tempo and Mode in Evolution (1941), described evolutionary history as being characterized by long periods of stasis (bradytely) or gradual change (horotely), punctuated by short bursts of rapid change (tachytely). This description of evolutionary change, however, is not a theory of the process(es) that produces it. Punctuated equilibrium is such a theory. Others have emphasized the theoretical novelty of punctuated equilibrium, and argued that evolutionary stasis had been "unexpected by most evolutionary biologists" and "had a major impact on paleontology and evolutionary biology".[20] Some critics jokingly referred to the theory as "evolution by jerks",[21] which elicited Gould to respond in kind by describing gradualism as "evolution by creeps".[22]

Evolutionary developmental biology Gould made significant contributions to evolutionary developmental biology,[23] especially in his work Ontogeny and Phylogeny. In this book he emphasized the process of heterochrony, which encompasses two distinct processes: pedomorphosis and terminal additions. Pedomorphosis is the process where ontogeny is slowed down and the organism does not reach the end of its development. Terminal addition is the process by which an organism adds to its development by speeding and shortening earlier stages in the developmental process. Gould's influence in the field of evolutionary developmental biology continues to be seen in such areas as the study of evolution of feathers.[24]

Selectionism and sociobiology Gould championed biological constraints such as the limitations of developmental pathways on evolutionary outcomes, as well as other non-selectionist forces in evolution. In particular, he considered many higher functions of the human brain to be the unintended side consequence or by-product of natural selection, rather than direct adaptations. To describe such co-opted features he coined the term exaptation with Elisabeth Vrba. Gould believed this understanding undermines an essential premise of human sociobiology and evolutionary psychology. Against "Sociobiology" In 1975, Gould's Harvard colleague E. O. Wilson introduced his analysis of animal behavior (including human behavior) based on a sociobiological framework that suggested that many social behaviors have a strong evolutionary basis.[25] In response, Gould, Richard Lewontin, and others from the Boston area wrote the subsequently well-referenced letter to The New York Review of Books entitled, "Against 'Sociobiology'". This open letter criticized Wilson's notion of a "deterministic view of human society and human action".[26] But Gould did not rule out sociobiological explanations for many aspects of animal behavior, and later wrote: "Sociobiologists have broadened their range of selective stories by invoking concepts of inclusive fitness and kin selection to solve (successfully I think) the vexatious problem of altruism—previously the greatest stumbling block

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to a Darwinian theory of social behavior... Here sociobiology has had and will continue to have success. And here I wish it well. For it represents an extension of basic Darwinism to a realm where it should apply."[27] Spandrels and the Panglossian Paradigm With Richard Lewontin, Gould wrote an influential 1979 paper entitled, "The Spandrels of San Marco and the panglossian paradigm",[28] which introduced the architectural term "spandrel" into evolutionary biology. In architecture, a spandrel is a curved area of masonry which exists between arches supporting a dome. Spandrels, also called pendentives in this context, are found particularly in Gothic churches. When visiting Venice in 1978, Gould noted that the spandrels of the San Marco cathedral, while quite beautiful, were not spaces planned by the architect. Rather the spaces arise as "necessary architectural byproducts of mounting a dome on rounded arches." Gould and Lewontin thus defined "spandrels" in the evolutionary biology context, A spandrel from the Holy Trinity Church in to mean any biological feature of an organism that arises as a necessary Fulnek, Czech Republic. side consequence of other features, which is not directly selected for by natural selection. Proposed examples include the "masculinized genitalia in female hyenas, exaptive use of an umbilicus as a brooding chamber by snails, the shoulder hump of the giant Irish deer, and several key features of human mentality."[29] In Voltaire's Candide, Dr. Pangloss is portrayed as a clueless scholar who, despite the evidence, insists that "all is for the best in this best of all possible worlds". Gould and Lewontin asserted that it is Panglossian for evolutionary biologists to view all traits as atomized things that had been naturally selected for, and criticised biologists for not granting theoretical space to other causes, such as phyletic and developmental constraints. The relative frequency of spandrels, so defined, versus adaptive features in nature, remains a controversial topic in evolutionary biology.[30] An illustrative example of Gould's approach can be found in Elisabeth Lloyd's case study suggesting that the female orgasm is a by-product of shared developmental pathways.[31] Gould also wrote on this topic in his essay "Male Nipples and Clitoral Ripples",[32] prompted by Lloyd's earlier work.

Evolutionary progress Gould favored the argument that evolution has no inherent drive towards long-term "progress". Uncritical commentaries often portray evolution as a ladder of progress, leading towards bigger, faster, and smarter organisms, the assumption being that evolution is somehow driving organisms to get more complex and ultimately more like humankind. Gould argued that evolution's drive was not towards complexity, but towards diversification. Because life is constrained to begin with a simple starting point ( like bacteria), any diversity resulting from this start, by random walk, will have a skewed distribution and therefore be perceived to move in the direction of higher complexity. But life, Gould argued, can also easily adapt towards simplification, as is often the case with parasites.[33] In a review of Full House, Richard Dawkins approved of Gould's general argument, but suggested that he saw evidence of a "tendency for lineages to improve cumulatively their adaptive fit to their particular way of life, by increasing the numbers of features which combine together in adaptive complexes. ... By this definition, adaptive evolution is not just incidentally progressive, it is deeply, dyed-in-the-wool, indispensably progressive."

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Cladistics Gould never embraced cladistics as a method of investigating evolutionary lineages and process, possibly because he was concerned that such investigations would lead to neglect of the details in historical biology, which he considered all-important. In the early 1990s this led him into a debate with Derek Briggs, who had begun to apply quantitative cladistic techniques to the Burgess Shale fossils, about the methods to be used in interpreting these fossils.[34] Around this time cladistics rapidly became the dominant method of classification in evolutionary biology. Inexpensive but increasingly powerful personal computers made it possible to process large quantities of data about organisms and their characteristics. Around the same time the development of effective polymerase chain reaction techniques made it possible to apply cladistic methods of analysis to biochemical and genetic features as well.[35]

Technical work on land snails Most of Gould's empirical research pertained to land snails. He focused his early work on the Bermudian genus Poecilozonites, while his later work concentrated on the West Indian genus Cerion. According to Gould "Cerion is the land snail of maximal diversity in form throughout the entire world. There are 600 described species of this single genus. In fact, they're not really species, they all interbreed, but the names exist to express a real phenomenon which is this incredible morphological diversity. Some are shaped like golf balls, some are shaped like pencils.…Now my main subject is the evolution of form, and the problem of how it is that you can get this diversity amid so little genetic difference, so far as we can tell, is a very interesting one. And if we could solve this we'd learn something general about the evolution of form."[36] Given Cerion's extensive geographic diversity, Gould later lamented that if Christopher Columbus had only cataloged a single Cerion it would have ended the scholarly debate about which island Columbus had first set foot on in America.[37]

Influence Gould is one of the most frequently cited scientists in the field of evolutionary theory. His 1979 "spandrels" paper has been cited more than 4,000 times.[38] In Paleobiology—the flagship journal of his own speciality—only Charles Darwin and George Gaylord Simpson have been cited more often.[39] Gould was also a considerably respected historian of science. Historian Ronald Numbers has been quoted as saying: "I can't say much about Gould's strengths as a scientist, but for a long time I've regarded him as the second most influential historian of science (next to Thomas Kuhn)."

The Structure of Evolutionary Theory Shortly before his death, Gould published a long treatise recapitulating his version of modern evolutionary theory: The Structure of Evolutionary Theory (2002).

As a public figure Gould became widely known through his popular essays on evolution in the Natural History magazine. His essays were published in a series titled This View of Life (a phrase from the concluding paragraph of Charles Darwin's Origin of Species) starting from January 1974 and ended in January 2001, amounting to a continuous publication of 300 essays. Many of his essays were reprinted in collected volumes that became bestselling books such as Ever Since Darwin and The Panda's Thumb, Hens' Teeth and Horses' Toes, and The Flamingo's Smile. A passionate advocate of evolutionary theory, Gould wrote prolifically on the subject, trying to communicate his understanding of contemporary evolutionary biology to a wide audience. A recurring theme in his writings is the history and development of pre-evolutionary and evolutionary thought. He was also an enthusiastic baseball fan and sabermetrician, and made frequent reference to the sport in his essays. Many of his baseball essays were

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Stephen Jay Gould anthologized in his posthumously published book Triumph and Tragedy in Mudville (2003).[40] Although a proud Darwinist, Gould's emphasis was less gradualist and reductionist than most neo-Darwinists. He fiercely opposed many aspects of sociobiology and its intellectual descendant evolutionary psychology. He devoted considerable time to fighting against creationism, creation science, and intelligent design. Most notably, Gould provided expert testimony against the equal-time creationism law in McLean v. Arkansas. Gould later developed the term "non-overlapping magisteria" (NOMA) to describe how, in his view, science and religion could not comment on each other's realm. Gould went on to develop this idea in some detail, particularly in the books Rocks of Ages (1999) and The Hedgehog, the Fox, and the Magister's Pox (2003). In a 1982 essay for Natural History Gould wrote: Our failure to discern a universal good does not record any lack of insight or ingenuity, but merely demonstrates that nature contains no moral messages framed in human terms. Morality is a subject for philosophers, theologians, students of the humanities, indeed for all thinking people. The answers will not be read passively from nature; they do not, and cannot, arise from the data of science. The factual state of the world does not teach us how we, with our powers for good and evil, should alter or preserve it in the most ethical manner.[41] The anti-evolution petition A Scientific Dissent From Darwinism spawned the National Center for Science Education's pro-evolution counterpart Project Steve, which is named in Gould's honor.[42] Gould also became a noted public face of science, often appearing on television. In 1984 Gould received his own NOVA special on PBS.[43] Other appearances included interviews on CNN's Crossfire, NBC's The Today Show, and regular appearances on the Charlie Rose show. Gould was also a guest in all seven episodes of the Dutch talk series A Glorious Accident, in which he appeared with Oliver Sacks.[44] Gould was featured prominently as a guest in Ken Burns's PBS documentary Baseball, as well as PBS's Evolution series. Gould was also on the Board of Advisers to the influential Children's Television Workshop television show 3-2-1 Contact, where he made frequent guest appearances. In 1997 he voiced a cartoon version of himself on the television series The Simpsons. In the episode "Lisa the Skeptic", Lisa finds a skeleton that many people believe is an apocalyptic angel. Lisa contacts Gould and asks him to test the skeleton's DNA. The fossil is discovered to be a marketing gimmick for a new mall.[45] During production the only phrase Gould objected to was a line in the script that introduced him as the "world's most brilliant paleontologist".[46] In 2002 the show paid tribute to Gould after his death, dedicating the season 13 finale to his memory. Gould had died two days before the episode aired.

Controversy Gould received many accolades for his scholarly work and popular expositions of natural history,[47] but was not immune from criticism by biologists who felt his public presentations were out of step with mainstream evolutionary theory.[48] The public debates between Gould's supporters and detractors have been so quarrelsome that they have been dubbed "The Darwin Wars" by several commentators.[49][50][51][52][53] John Maynard Smith, an eminent British evolutionary biologist, was among Gould's strongest critics. Maynard Smith thought that Gould misjudged the vital role of adaptation in biology, and was critical of Gould's acceptance of species selection as a major component of biological evolution.[54] In a review of Daniel Dennett's book Darwin's Dangerous Idea, Maynard Smith wrote that Gould "is giving non-biologists a largely false picture of the state of evolutionary theory."[55] But Maynard Smith has not been consistently negative, writing in a review of The Panda's Thumb that "Stephen Gould is the best writer of popular science now active... Often he infuriates me, but I hope he will go right on writing essays like these."[56] Maynard Smith was also among those who welcomed Gould's reinvigoration of evolutionary paleontology. One reason for criticism was that Gould appeared to be presenting his ideas as a revolutionary way of understanding evolution, and argued for the importance of mechanisms other than natural selection, mechanisms which he believed

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Stephen Jay Gould had been ignored by many professional evolutionists. As a result, many non-specialists sometimes inferred from his early writings that Darwinian explanations had been proven to be unscientific (which Gould never tried to imply). Along with many other researchers in the field, Gould's works were sometimes deliberately taken out of context by creationists as "proof" that scientists no longer understood how organisms evolved.[57] Gould himself corrected some of these misinterpretations and distortions of his writings in later works.[58] As documented by Kim Sterelny among others, Gould disagreed with Richard Dawkins about the importance of gene selection in evolution. Dawkins argued that evolution is best understood as competition among genes (or replicators), while Gould advocated the importance of multi-level selection, including selection amongst genes, cell lineages, organisms, demes, species, and clades. Dawkins also found that Gould deliberately played down the difference between rapid gradualism and macromutation in his theory of punctuated equilibrium.[59] Criticism of Gould and his theory of punctuated equilibrium can be found in Dawkins' The Blind Watchmaker and Unweaving the Rainbow, as well as chapter 10 of Dennett's Darwin's Dangerous Idea.

Cambrian fauna Gould's interpretation of the Cambrian Burgess Shale fossils in his book Wonderful Life emphasized the striking morphological disparity (or "weirdness") of the Burgess Shale fauna, and the role of chance in determining which members of this fauna survived and flourished. He used the Cambrian fauna as an example of the role of contingency in the broader pattern of evolution. His view was criticized by Simon Conway Morris in his 1998 book The Crucible of Creation. Conway Morris stressed those members of the Cambrian fauna that resemble modern taxa. He also promoted convergent evolution as a mechanism producing similar forms in similar environmental circumstances, and argued in a subsequent book that the appearance of human-like animals is likely. Paleontologists Derek Briggs and Richard Fortey have also argued that much of the Cambrian fauna may be regarded as stem groups of living taxa,[60] though this is still a subject of intense research and debate, and the relationship of many Cambrian taxa to modern phyla has not been established in the eyes of many palaeontologists. Paleontologist Richard Fortey noted that prior to the release of Wonderful Life, Conway Morris shared many of Gould's sentiments and views. It was only after publication of Wonderful Life that Conway Morris revised his interpretation and adopted a more progressive stance towards the history of life.[61] Richard Dawkins also disagreed with Gould's interpretation of the Burgess Shale, arguing: The extreme Gouldian view—certainly the view inspired by his rhetoric, though it is hard to tell from his own words whether he literally holds it himself—is radically different from and utterly incompatible with the standard neo-Darwinian model.... For a new body plan—a new phylum—to spring into existence, what actually has to happen on the ground is that a child is born which suddenly, out of the blue, is as different from its parents as a snail is from an earthworm. No zoologist who thinks through the implications, not even the most ardent saltationist, has ever supported any such notion.[62]

Opposition to sociobiology and evolutionary psychology Gould also had a long-running public feud with E. O. Wilson and other evolutionary biologists about human sociobiology and its later descendant evolutionary psychology (which Gould, Lewontin, and Maynard Smith opposed, but which Richard Dawkins, Daniel Dennett, and Steven Pinker advocated).[63] These debates reached their climax in the 1970s, and included strong opposition from groups like the Sociobiology Study Group and Science for the People.[64] Pinker accuses Gould, Lewontin, and other opponents of evolutionary psychology of being "radical scientists", whose stance on human nature is influenced by politics rather than science. Gould stated that he made "no attribution of motive in Wilson's or anyone else's case" but cautioned that all human beings are influenced, especially unconsciously, by our personal expectations and biases. He wrote:

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Stephen Jay Gould I grew up in a family with a tradition of participation in campaigns for social justice, and I was active, as a student, in the civil rights movement at a time of great excitement and success in the early 1960s. Scholars are often wary of citing such commitments. … [but] it is dangerous for a scholar even to imagine that he might attain complete neutrality, for then one stops being vigilant about personal preferences and their influences—and then one truly falls victim to the dictates of prejudice. Objectivity must be operationally defined as fair treatment of data, not absence of preference.[65] Gould's primary criticism held that human sociobiological explanations lacked evidential support, and argued that adaptive behaviors are frequently assumed to be genetic for no other reason than their supposed universality, or their adaptive nature. Gould emphasized that adaptive behaviors can be passed on through culture as well, and either hypothesis is equally plausible.[66] Gould did not deny the relevance of biology to human nature, but reframed the debate as "biological potentiality vs. biological determinism". Gould stated that the human brain allows for a wide range of behaviors. Its flexibility "permits us to be aggressive or peaceful, dominant or submissive, spiteful or generous… Violence, sexism, and general nastiness are biological since they represent one subset of a possible range of behaviors. But peacefulness, equality, and kindness are just as biological—and we may see their influence increase if we can create social structures that permit them to flourish."

The Mismeasure of Man Gould was the author of The Mismeasure of Man (1981), a history and inquiry of psychometrics and intelligence testing. Gould investigated the methods of nineteenth century craniometry, as well as the history of psychological testing. Gould claimed that both theories developed from an unfounded belief in biological determinism, the view that "social and economic differences between human groups—primarily races, classes, and sexes—arise from inherited, inborn distinctions and that society, in this sense, is an accurate reflection of biology."[67] It was reprinted in 1996 with the addition of a new foreword and a critical review of The Bell Curve. The Mismeasure of Man has generated perhaps the greatest controversy of all of Gould's books. It has received both widespread praise[68] and extensive criticism,[69] including claims of misrepresentation. In 2011, a study conducted by six anthropologists reanalyzed Gould's claim that Samuel Morton unconsciously manipulated his skull measurements,[70] and concluded that Gould's analysis was poorly supported and incorrect. They praised Gould for his "staunch opposition to racism" but concluded, "we find that Morton's initial reputation as the objectivist of his era was well-deserved."[71] Ralph Holloway, one of the co-authors of the study, commented, "I just didn't trust Gould...I had the feeling that his ideological stance was supreme. When the 1996 version of 'The Mismeasure of Man' came and he never even bothered to mention Michael's study, I just felt he was a charlatan."[72] The group's paper was reviewed in the journal Nature, which recommended a degree of caution, stating "the critique leaves the majority of Gould's work unscathed," and notes that "because they couldn't measure all the skulls, they do not know whether the average cranial capacities that Morton reported represent his sample accurately."[73] The journal stated that Gould's opposition to racism may have biased his interpretation of Morton's data, but also noted that "Lewis and his colleagues have their own motivations. Several in the group have an association with the University of Pennsylvania, and have an interest in seeing the valuable but understudied skull collection freed from the stigma of bias."

Non-overlapping magisteria In his book Rocks of Ages (1999), Gould put forward what he described as "a blessedly simple and entirely conventional resolution to...the supposed conflict between science and religion."[74] He defines the term magisterium as "a domain where one form of teaching holds the appropriate tools for meaningful discourse and resolution." The non-overlapping magisteria (NOMA) principle therefore divides the magisterium of science to cover "the empirical realm: what the Universe is made of (fact) and why does it work in this way (theory). The magisterium of religion extends over questions of ultimate meaning and moral value. These two magisteria do not overlap, nor do they

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encompass all inquiry." He suggests that "NOMA enjoys strong and fully explicit support, even from the primary cultural stereotypes of hard-line traditionalism" and that NOMA is "a sound position of general consensus, established by long struggle among people of goodwill in both magisteria." However, this view has not been without criticism. For example, in his book The God Delusion, Richard Dawkins argues that the division between religion and science is not as simple as it seems, as few religions exist without claiming the existence of miracles, which "by definition, violate the principles of science".[75] Dawkins also opposes the idea that religion has anything meaningful to say about ethics and values, and therefore has no authority to claim a magisterium of its own.

Gould's publications Articles Gould's publications were numerous. One review of his publications between 1965 and 2000 noted 479 peer-reviewed papers, 22 books, 300 essays, and 101 "major" book reviews.[76] A select number of his papers are listed online [77].

Books The following is a list of books either written or edited by Stephen Jay Gould, including those published posthumously, after his death in 2002. While some books have been republished at later dates, by multiple publishers, the list below comprises the original publisher and publishing date. •

1977. Ontogeny and Phylogeny, Cambridge MA: Belknap Press • of Harvard University Press, ISBN 0-674-63940-5 online [78] preview

1995. Dinosaur in a Haystack, New York: Harmony Books, ISBN 0-517-70393-9

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1977. Ever Since Darwin, New York: W. W. Norton, ISBN 978-0-393-06425-4

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1996. Full House: The Spread of Excellence From Plato to Darwin, New York: Harmony Books, ISBN 0-517-70394-7

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1980. The Panda's Thumb, New York: W. W. Norton, ISBN 0-393-01380-4

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1997. Questioning the Millennium: A Rationalist's Guide to a Precisely Arbitrary Countdown, New York: Harmony Books, ISBN 0-609-60541-0

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1980. Gould, Stephen Jay (1980-12), The Evolution of [79] Gryphaea , New York: Arno Press, ISBN 0-405-12751-0

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1998. Leonardo's Mountain of Clams and the Diet of Worms, N.Y.: Harmony Books, ISBN 0-609-60141-5

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1981. The Mismeasure of Man, New York: W. W. Norton, ISBN 978-0-393-31425-0

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1999. Rocks of Ages: Science and Religion in the Fullness of Life, New York: Ballantine Books, ISBN 0-345-43009-3

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1983. Hen's Teeth and Horse's Toes, New York: W. W. Norton, • ISBN 978-0-393-01716-8

2000. The Lying Stones of Marrakech, New York: Harmony Books, ISBN 0-609-60142-3

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1985. The Flamingo's Smile, New York: W. W. Norton, ISBN 0-393-02228-5

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2000. Crossing Over: Where Art and Science Meet, New York: Three Rivers Press, ISBN 0-609-80586-X

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1987. Time's Arrow, Time's Cycle, Cambridge MA: Harvard [80] Univ. Press, ISBN 0-674-89198-8 online preview

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2002. The Structure of Evolutionary Theory, Cambridge MA: Belknap Press of Harvard University Press, ISBN 978-0-674-00613-3 online [81] preview

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1987. An Urchin in the Storm: Essays about Books and Ideas, N.Y.: W. W. Norton, ISBN 0-393-02492-X

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2002. I Have Landed: The End of a Beginning in Natural History, New York: Harmony Books, ISBN 0-609-60143-1

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1987. (with Rosamond Wolff Purcell) Illuminations: A Bestiary, • N.Y.: W. W. Norton, ISBN 0-393-30436-1

2003. Triumph and Tragedy in Mudville: A Lifelong Passion for Baseball, New York: W. W. Norton, ISBN 0-393-05755-0

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1989. Wonderful Life: The Burgess Shale and the Nature of History, New York: W. W. Norton, ISBN 0-393-02705-8. 347 pp.

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2003. The Hedgehog, the Fox, and the Magister's Pox, New York: Harmony Books, ISBN 0-609-60140-7

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1991. Bully for Brontosaurus, New York: W. W. Norton, ISBN 978-0-393-02961-1. 540 pp.

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2006. The Richness of Life: the Essential Stephen Jay Gould, London: Jonathan Cape, ISBN 978-0-09-948867-5 This is an anthology of Gould's writings edited by Paul McGarr and Steven Rose, introduced by Steven Rose.

Stephen Jay Gould •

1992. (with Rosamond Wolff Purcell) Finders, Keepers: Eight Collectors, New York: W. W. Norton, ISBN 978-0-393-03054-9

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1993. Eight Little Piggies, New York: W. W. Norton, ISBN 0-393-03416-X

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1993. The Book of Life. Preface, pp. 6–21. New York: W. W. Norton (S. J. Gould general editor, 10 contributors). ISBN [83] 0-393-05003-3 review citing original publishing date

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2007. Punctuated Equilibrium, Cambridge MA: Belknap Press of Harvard [82] University Press, ISBN 0-674-02444-3 Book review

Notes and references Specific citations: [1] Eldredge, Niles, and S. J. Gould (1972). "Punctuated equilibria: an alternative to phyletic gradualism." (http:/ / www. blackwellpublishing. com/ ridley/ classictexts/ eldredge. pdf) In T.J.M. Schopf, ed., Models in Paleobiology. San Francisco: Freeman, Cooper and Company, pp. 82–115. [2] Gould, S. J. (1997). "Nonoverlapping magisteria". (http:/ / www. stephenjaygould. org/ library/ gould_noma. html) Natural History 106 (March): 16–22. [3] Green, Michelle (1986). "Stephen Jay Gould: driven by a hunger to learn and to write". (http:/ / www. stephenjaygould. org/ library/ green_sjgould. html) People 25 (June 2): 109–114. [4] Gould, S. J. (2002). The Structure of Evolutionary Theory. Cambridge: Belknap Press of Harvard University Press. ISBN 0-674-00613-5 [5] Gould, S. J. (1981). "Official Transcript for Gould’s deposition in McLean v. Arkansas". (http:/ / www. antievolution. org/ projects/ mclean/ new_site/ depos/ pf_gould_dep. htm) (Nov. 27). Under oath Gould stated: "My political views tend to the left of center. Q. Could you be more specific about your political views? A. I don't know how to be. I am not a joiner, so I am not a member of any organization. So I have always resisted labeling. But if you read my other book, The Mismeasure of Man, which is not included because it is not about evolution, you will get a sense of my political views." p. 153. [6] Gasper, Phil (2002). "Stephen Jay Gould: (http:/ / www. isreview. org/ issues/ 24/ gould. shtml) Dialectical Biologist". International Socialist Review 24 (July–August). [7] Lewontin, Richard and Richard Levins (2002). "Stephen Jay Gould—what does it mean to be a radical?" (http:/ / www. monthlyreview. org/ 1102lewontin. htm) Monthly Review 54 (Nov. 1). "The public intellectual and political life of Steve Gould was extraordinary, if not unique. First, he was an evolutionary biologist and historian of science whose intellectual work had a major impact on our views of the process of evolution. Second, he was, by far, the most widely known and influential expositor of science who has ever written for a lay public. Third, he was a consistent political activist in support of socialism and in opposition to all forms of colonialism and oppression. The figure he most closely resembled in these respects was the British biologist of the 1930s, J. B. S. Haldane, a founder of the modern genetical theory of evolution, a wonderful essayist on science for the general public, and an idiosyncratic Marxist and columnist for the Daily Worker who finally split with the Communist Party over its demand that scientific claims follow Party doctrine." [8] Gould, S. J. (1993). "Dinomania". (http:/ / www. nybooks. com/ articles/ 2483) New York Review of Books 40 (August 12): 51–56. [9] Gould, S. J. (1983). Hen's Teeth and Horse's Toes. New York: W. W. Norton & Company. ISBN 0-393-31103-1. [10] Carol Kaesuk Yoon (2002). "Stephen Jay Gould, 60, Is Dead; Enlivened Evolutionary Theory," (http:/ / www. nytimes. com/ 2002/ 05/ 21/ us/ stephen-jay-gould-60-is-dead-enlivened-evolutionary-theory. html) New York Times May 21, 2002. [11] Gould, S. J. (1985). "The Median Isn't the Message". (http:/ / www. cancerguide. org/ median_not_msg. html) Discover 6 (June): 40–42. [12] Bakalar, James and Lester Grinspoon (1997). Marihuana, the Forbidden Medicine. New Haven: Yale University Press, pp. 39–41. (http:/ / www. stephenjaygould. org/ library/ gould_marijuana. html) [13] Harvard News Office (2002). "Paleontologist, author Gould dies at 60". (http:/ / www. news. harvard. edu/ gazette/ 2002/ 05. 16/ 99-gould. html) The Harvard Gazette. (May 20). Retrieved on June 4, 2009. [14] Krementz, Jill (2002). "Jill Krementz Photo Journal". (http:/ / www. asrlab. org/ archive/ jillPage. htm) New York Social Diary. Retrieved on June 4, 2009. [15] Allen, Warren (2008). "The Structure of Gould". (http:/ / books. google. com/ books?id=6iVOHEVeSFMC& pg=PA3) In Warren Allen et al. Stephen Jay Gould: Reflections on His View of Life. Oxford: Oxford University Press, p. 24, 59. [16] Masha, Etkin (2002). "A Tribute to Stephen Jay Gould '63". (http:/ / antiochians. org/ antiochdocs/ antiochian/ archive/ Antiochian_wi02/ winter2002/ book_gould. html) Antiochian (Winter ed.). Retrieved on June 4, 2009. [17] Linnean Society of London (2008). "The Darwin–Wallace Medal". (http:/ / linnean. org/ index. php?id=344l) Retrieved on June 4, 2009. [18] Eldredge, Niles (1971). "The Allopatric Model and Phylogeny in Paleozoic Invertebrates." Evolution Vol. 25, No. 1 (Mar. 1971), pp. 156–167. [19] Dawkins, Richard (1999). The Extended Phenotype. Oxfordshire: Oxford University Press. p. 101. ISBN 0-19-288051-9. [20] Mayr, Ernst (1992). "Speciational Evolution or Punctuated Equilibria". (http:/ / www. stephenjaygould. org/ library/ mayr_punctuated. html) In Steven Peterson and Albert Somit. The Dynamics of Evolution. Ithaca: Cornell University Press, pp. 21–48. Special:BookSources/0801497639ISBN 0-8014-9763-9.

Stephen Jay Gould [21] Turner, John (1984). "Why we need evolution by jerks." (http:/ / books. google. com/ books?id=6idhDboGmZoC& pg=PA34) New Scientist 101 (Feb. 9): 34–35. [22] Gould, S. J. and Steven Rose, ed. (2007). The Richness of Life: The Essential Stephen Jay Gould. New York: W. W. Norton & Co., p. 6. (http:/ / books. google. com/ books?id=yfXJhKmp1wUC& lpg=PP1& pg=PA6) [23] Thomas, R.D.K. (2009). "Gould, Stephen Jay (1941–2002)". in M. Ruse and J. Travis (eds). Evolution: The First Four Billion Years. Cambridge MA: Belknap Press. pp. 611–615. [24] Prum, R.O., & Brush, A.H. (March 2003). "Which Came First, the Feather or the Bird?" Scientific American, vol.288, no.3, pp. 84–93 [25] Wilson, E. O. (1975). Sociobiology: The New Synthesis. Cambridge MA: Harvard University Press. [26] Allen, Elizabeth, et al. (1975). "Against 'Sociobiology'". (http:/ / www. nybooks. com/ articles/ 9017?sess=305fe41afae729849e1e7eb4b004bb81) [letter] New York Review of Books 22 (Nov. 13): 182, 184–186. [27] Gould, S. J. (1980). "Sociobiology and the Theory of Natural Selection". In G. W. Barlow and J. Silverberg, eds., Sociobiology: Beyond Nature/Nurture? Boulder CO: Westview Press, pp. 257–269. [28] Gould, S. J. and Richard Lewontin (1979). "The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme". (http:/ / www. aaas. org/ spp/ dser/ 03_Areas/ evolution/ perspectives/ Gould_Lewontin_1979. shtml) Proc. R. Soc. Lond., B, Biol. Sci. 205 (1161): 581–98. DOI (http:/ / dx. doi. org/ 10. 1098/ rspb. 1979. 0086) PMID (http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 42062); for background see Gould's "The Pattern of Life's History" (http:/ / www. edge. org/ documents/ ThirdCulture/ i-Ch. 2. html) in John Brockman The Third Culture (http:/ / www. edge. org/ documents/ ThirdCulture/ d-Contents. html). New York: Simon & Schuster. 1996, pp. 52–64. ISBN 0-684-82344-6. [29] Gould, S. J. (1997). "The exaptive excellence of spandrels as a term and prototype". (http:/ / www. pnas. org/ content/ 94/ 20/ 10750. full) Proc. Natl. Acad. Sci. U.S.A. 94 (20): 10750–5. DOI (http:/ / dx. doi. org/ 10. 1073/ pnas. 94. 20. 10750) PMID (http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 11038582) [30] Maynard Smith, John (1995). "Genes, Memes, & Minds". (http:/ / www. nybooks. com/ articles/ 1703) The New York Review of Books 42 (Nov. 30): 46–48. "By and large, I think their [Spandrels] paper had a healthy effect. . . . Their critique forced us to clean up our act and to provide evidence for our stories. But adaptationism remains the core of biological thinking." A similar appraisal is reflected by Ernst Mayr in his 1983 paper "How to Carry Out the Adaptationist Program?" The American Naturalist 121 (3): 324–334; and George C. Williams, Natural Selection: Domains, Levels, and Challenges. New York: Oxford University Press. 1992. [31] Lloyd, E.A. (2005). The Case of The Female Orgasm: Bias in the science of evolution. Cambridge MA: Harvard University Press. [32] Gould, S.J. (1992). "Male Nipples and Clitoral Ripples". (http:/ / books. google. com/ books?id=pzj90slTTEIC& pg=PA124) In Bully for Brontosaurus: Further Reflections in Natural History. New York: W. W. Norton. pp. 124–138. [33] Gould, S. J. (1996). Full House: The Spread of Excellence From Plato to Darwin. New York: Harmony Books. [34] Gould, S. J. (1991). "The disparity of the Burgess Shale arthropod fauna and the limits of cladistic analysis". (http:/ / webh01. ua. ac. be/ funmorph/ raoul/ macroevolutie/ Gould1991. pdf) Paleobiology 17 (October): 411–423. [35] Baron, Christian and J. T. Høeg (2005). "Gould, Scharm and the Paleontologocal Perspective in Evolutionary Biology". (http:/ / books. google. co. uk/ books?id=LalmQ4346O0C& pg=PA3) In S. Koenemann and R.A. Jenner, Crustacea and Arthropod Relationships. CRC Press. pp. 3–14. ISBN 0-8493-3498-5. [36] Wolpert, Lewis and Alison Richards (1998). A Passion For Science. Oxford: Oxford University Press, pp. 139–152. (http:/ / www. stephenjaygould. org/ library/ wolpert_sjg-interview. html) ISBN 0-19-854212-7 [37] Gould, S. J. (1996). "A Cerion for Christopher". Natural History 105 (Oct.): 22–29, 78—79. [38] Google Scholar. http:/ / scholar. google. com (http:/ / scholar. google. com/ scholar?cluster=8886524888381702203). Retrieved on 2011-6-12. [39] Prothero, Donald (2000). "Evolution Revolution: Paleontology, History, Biography". (http:/ / www. skeptic. com/ Merchant2/ merchant. mvc?Screen=PROD& Store_Code=SS& Product_Code=av093& Category_Code=) Skeptic Festschrift lecture for Stephen Jay Gould. October 7, 2000. [40] Gould, S. J. (2003). Triumph and Tragedy in Mudville (http:/ / books. google. com/ books?id=XIJ-ay4GJ_kC& pg=PA). New York: W. W. Norton & Co. See his essays: "The Streak of Streaks", (http:/ / www. nybooks. com/ articles/ 4337) , and "Baseball's reliquary: the oddly possible hybrid of shrine and university" (http:/ / findarticles. com/ p/ articles/ mi_m1134/ is_2_111/ ai_83553539/ ) [41] Gould, S. J. (1982). "Nonmoral Nature". (http:/ / www. stephenjaygould. org/ library/ gould_nonmoral. html) Natural History 91 (Feb.): 19–26. [42] NCSE Project Steve (http:/ / ncse. com/ taking-action/ project-steve) official webpage, National Center for Science Education, February 16, 2003 [43] PBS (1984). "Stephen Jay Gould: This View of Life". (http:/ / www. stephenjaygould. org/ audio/ gould_nova-special. html) NOVA. December 18. [44] Sacks, Oliver (2007). Forward. (http:/ / books. google. com/ books?id=yfXJhKmp1wUC& pg=PP17) In Steven Rose, ed. The Richness of Life. New York: W. W. Norton & Company, p. xi. Video (http:/ / www. youtube. com/ watch?v=SJzqHnwNbUY) [45] Fox. The Simpsons. "Lisa the Skeptic", November 23, 1997. Audio here. (http:/ / www. stephenjaygould. org/ audio/ thesimpsons. rm) [46] Scully, Mike (2006). The Simpsons. Season 9 DVD Commentary for "Lisa the Skeptic". DVD. 20th Century Fox. [47] Shermer, Michael (2002). "This View of Science" (http:/ / www. stephenjaygould. org/ library/ shermer_sjgould. pdf). Social Studies of Science 32 (4): 518.

• This is almost all of Schermer's note 10 (which cites "Gould's curriculum vitae, dated September 2000"):

168

Stephen Jay Gould Awards include a National Book Award for The Panda's Thumb, a National Book Critics Circle Award for The Mismeasure of Man, the Phi Beta Kappa Book Award for Hen's Teeth and Horse's Toes, and a Pulitzer Prize Finalist for Wonderful Life, on which Gould commented "close but, as they say, no cigar". Forty-four honorary degrees and 66 major fellowships, medals, and awards bear witness to the depth and scope of his accomplishments in both the sciences and humanities: Member of the National Academy of Sciences, President and Fellow of AAAS, MacArthur Foundation 'genius' Fellowship (in the first group of awardees), Humanist Laureate from the Academy of Humanism, Fellow of the Linnean Society of London, Fellow of the Royal Society of Edinburgh, Fellow of the American Academy of Arts and Sciences, Fellow of the European Union of Geosciences, Associate of the Muséum National D'Histoire Naturelle Paris, the Schuchert Award for excellence in paleontological research, Scientist of the Year from Discover magazine, the Silver Medal from the Zoological Society of London, the Gold Medal for Service to Zoology from the Linnean Society of London, the Edinburgh Medal from the City of Edinburgh, the Britannica Award and Gold Medal for dissemination of public knowledge, Public Service Award from the Geological Society of America, Anthropology in Media Award from the American Anthropological Association, Distinguished Service Award from the National Association of Biology Teachers, Distinguished Scientist Award from UCLA, the Randi Award for Skeptic of the Year from the Skeptics Society, and a Festschrift in his honour at Caltech. [48] These are the first two of 25 paragraphs, with notes, from a "Letter to the Editor of The New York Review of Books" (http:/ / cogweb. ucla. edu/ Debate/ CEP_Gould. html) by Leda Cosmides and John Tooby (July 7, 1997). They wrote in comment on two recent NYRB articles by Gould (June 12 and 26). The source (cogweb.ucla.edu, August 2002) does not say whether NYRB published the letter.

John Maynard Smith, one of the world's leading evolutionary biologists, recently summarized in the NYRB the sharply conflicting assessments of Stephen Jay Gould: "Because of the excellence of his essays, he has come to be seen by non-biologists as the pre-eminent evolutionary theorist. In contrast, the evolutionary biologists with whom I have discussed his work tend to see him as a man whose ideas are so confused as to be hardly worth bothering with, but as one who should not be publicly criticized because he is at least on our side against the creationists." (NYRB, November 30, 1995, p. 46). No one can take any pleasure in the evident pain Gould is experiencing now that his actual standing within the community of professional evolutionary biologists is finally becoming more widely known. If what was a stake was solely one man's self-regard, common decency would preclude comment. But as Maynard Smith points out, more is at stake. Gould "is giving non-biologists a largely false picture of the state of evolutionary theory"—or as Ernst Mayr says of Gould and his small group of allies—they "quite conspicuously misrepresent the views of [biology's] leading spokesmen."{1}. Indeed, although Gould characterizes his critics as "anonymous" and "a tiny coterie", nearly every major evolutionary biologist of our era has weighed in a vain attempt to correct the tangle of confusions that the higher profile Gould has inundated the intellectual world with.{2} The point is not that Gould is the object of some criticism—so properly are we all—it is that his reputation as a credible and balanced authority about evolutionary biology is non-existent among those who are in a professional position to know. ... {1} [Full reference provided by the writers.] {2} These include Ernst Mayr, John Maynard Smith, George Williams, Bill Hamilton, Richard Dawkins, E.O. Wilson, Tim Clutton-Brock, Paul Harvey, Brian Charlesworth, Jerry Coyne, Robert Trivers, John Alcock, Randy Thornhill, and many others. It should be noted that where Tooby & Cosmides quote Ernst Mayr, he does not speak of Gould in particular, and does not mention him by name, but speaks generally of the critics of the Neo-Darwinian Synthesis. • The list of experts provided by Tooby and Cosmides is also questionable. For example, Mayr, Williams, Hamilton, Dawkins, Wilson, Coyne, and Trivers have shown great respect for Gould as a scientist. In the first of the two articles that provoked Tooby & Cosmides, Gould had commented on the November 1995 review of his work by Maynard Smith, which they quoted in support (quoted in this note).

169

Stephen Jay Gould • Gould, "Darwinian Fundamentalism" (webpage 2) (http:/ / www. nybooks. com/ articles/ archives/ 1997/ jun/ 12/ darwinian-fundamentalism/?page=2), reprinted from New York Review of Books 44 (June 12, 1997): 34–37. A false fact can be refuted, a false argument exposed; but how can one respond to a purely ad hominem attack? This harder, and altogether more discouraging, task may best be achieved by exposing internal inconsistency and unfairness of rhetoric. [quotation of Smith's November 1995 NYRB quotation of Gould, later quoted by Tooby & Consides (above)] It seems futile to reply to an attack so empty of content, and based only on comments by anonymous critics; ... Instead of responding to Maynard Smith's attack against my integrity and scholarship, citing people unknown and with arguments unmentioned, let me, instead, merely remind him of the blatant inconsistency between his admirable past and lamentable present. Some sixteen years ago he wrote a highly critical but wonderfully supportive review of my early book of essays, The Panda's Thumb, stating: "I hope it will be obvious that my wish to argue with Gould is a compliment, not a criticism." He then attended my series of Tanner Lectures at Cambridge in 1984 and wrote in a report for Nature, and under the remarkable title "Paleontology at the High Table", the kindest and most supportive critical commentary I have ever received. He argued that the work of a small group of American paleobiologists had brought the entire subject back to theoretical centrality within the evolutionary sciences. ... [multiple paragraphs] So we face the enigma of a man who has written numerous articles, amounting to tens of thousands of words, about my work—always strongly and incisively critical, always richly informed (and always, I might add, enormously appreciated by me). But now Maynard Smith needs to canvass unnamed colleagues to find out that my ideas are "hardly worth bothering with". He really ought to be asking himself why he has been bothering about my work so intensely, and for so many years. ... [49] Brown, Andrew (1999). The Darwin Wars: The Scientific Battle for the Soul of Man. London: Simon & Schuster. ISBN 0-8050-7137-7 [50] Rose, Steven (2002). "Obituaries: Stephen Jay Gould". (http:/ / www. guardian. co. uk/ education/ 2002/ may/ 22/ medicalscience. internationaleducationnews) The Guardian (May 22): 20. [51] Blume, Harvey (2002). "The Origin of Specious". (http:/ / www. prospect. org/ cs/ articles?article=the_origin_of_specious) The American Prospect (September 22): 41–43. [52] Also ISBN 978-1-84046-780-2 [53] Dawkins (1998), p. 195 (http:/ / books. google. com. hk/ books?id=ZudTchiioUoC& printsec=frontcover& dq=unweaving+ the+ rainbow& source=bl& ots=vUbqz7NJs0& sig=P-FaM4jTHMMv4d-ZjM4yvd--P30& hl=zh-CN& sa=X& ei=CQsnUP38OcTtrQeTuIG4DA& ved=0CDUQ6AEwAA#v=onepage& q=gould& f=false) [54] Maynard Smith, John (1981). "Did Darwin get it right?" The London Review of Books 3 (11): 10–11; Also reprinted in Did Darwin Get it Right? New York: Chapman and Hall, 1989, pp. 148–156. [55] Maynard Smith, John (1995). "Genes, Memes, & Minds". (http:/ / www. nybooks. com/ articles/ 1703) The New York Review of Books 42 (Nov. 30): 46–48. [56] Maynard Smith, John (1981). "Review of The Panda's Thumb" The London Review of Books pp. 17–30; Reprinted as "Tinkering" in his Did Darwin Get It Right? New York: Chapman and Hall. 1989, pp. 94, 97. [57] Wright, Robert (1999). "The Accidental Creationist: Why Stephen J. Gould is bad for evolution". (http:/ / www. nonzero. org/ newyorker. htm) The New Yorker 75 (Dec. 13): 56–65. [58] Gould, S. J. (1981). "Evolution as fact and theory". (http:/ / www. stephenjaygould. org/ library/ gould_fact-and-theory. html) Discover 2 (May): 34–37. [59] "It is when we ask what happens during the sudden bursts of species formation that the confusion... arises... Gould is aware of the difference between rapid gradualism and macromutation, but he treats the matter as though it were a minor detail, to be cleared up after we have taken on board the overarching question of whether evolution is episodic rather than gradual." As seen at: Dawkins, Richard (1998). Unweaving the Rainbow, pp. 196–197 (http:/ / books. google. com/ books?id=ZudTchiioUoC& pg=PA202) [60] Abstract (http:/ / www. psjournals. org/ doi/ abs/ 10. 1666/ 0094-8373(2005)031[0094:WSSSGA]2. 0. CO;2) [61] Fortey, Richard (1998). "Shock Lobsters". (http:/ / www. lrb. co. uk/ v20/ n19/ fort01_. html) London Review of Books 20 (Oct. 1). [62] Dawkins, Richard (1998). Unweaving the Rainbow, p. 202. (http:/ / books. google. com/ books?id=ZudTchiioUoC& pg=PA202) [63] Gould, S. J. (1997). "Evolution: The pleasures of pluralism". (http:/ / www. stephenjaygould. org/ reviews/ gould_pluralism. html) The New York Review of Books 44 (June 26): 47–52. [64] Wilson, E. O. (2006). Naturalist New York: Island Press, p.337 (http:/ / books. google. com/ books?id=TZH2nHEPSjYC& pg=PA337) ISBN 1-59726-088-6.

170

Stephen Jay Gould [65] Gould S. J. (1996). The Mismeasure of Man: Revised and Expanded Edition. New York: W. W. Norton & Co., p. 36. ISBN 0-14-025824-8 [66] Gould, S. J. (1992). "Biological potentiality vs. biological determinism". (http:/ / books. google. com/ books?id=_VCnI02FwHAC& pg=PA251) In Ever Since Darwin. New York: W. W. Norton & Co., pp. 251–259. [67] Gould, S. J. (1981). The Mismeasure of Man. New York: W.W. Norton & Co. p. 20. [68] In 1981 The Mismeasure of Man won the National Book Critics Circle Award for non-fiction. It was voted as the 17th greatest science book of all time by Discover magazine vol. 27 (December 8, 2006); 9th best skeptic book by The Skeptics Society (Frank Diller, "Scientists' Nightstand" American Scientist); and ranked 24th place for the best non-fiction book by the Modern Library. [69] Blinkhorn, Steve (1982). "What Skulduggery?" (http:/ / www. skepticfiles. org/ evolut/ mismeasr. htm) Nature 296 (April 8): 506. [70] Gould, S. J. (1978). "Morton's Ranking of Races by Cranial Capacity." (http:/ / www. colorado. edu/ MCDB/ MCDB4234/ readings/ Morton-Skulls-Gould. pdf) Science 200 (May 5): 503–509. [71] Lewis, J., E., DeGusta, D. Meyer, M.R., Monge, J.M., Mann, A.E. and Holloway, R.L. (2011). "The Mismeasure of Science: Stephen Jay Gould versus Samuel George Morton on Skulls and Bias." (http:/ / www. plosbiology. org/ article/ info:doi/ 10. 1371/ journal. pbio. 1001071) Public Library of Science Biology 9 (6): e1001071. [72] Wade, Nicholas (2011). "Scientists Measure the Accuracy of a Racism Claim." (http:/ / www. nytimes. com/ 2011/ 06/ 14/ science/ 14skull. html?_r=1& ref=science) New York Times (June 14): D4. [73] Editorial (2011). "Mismeasure for mismeasure." (http:/ / www. nature. com/ nature/ journal/ v474/ n7352/ full/ 474419a. html) Nature 474 (June 23): 419. [74] Gould, S. J. (2002). Rocks of Ages: Science and Religion in the Fullness of Life. New York: Ballantine Books. [75] Dawkins, R. (2006). The God Delusion. New York: Houghton Mifflin Harcourt, p. 83. (http:/ / books. google. com/ books?id=yq1xDpicghkC& pg=PA83) [76] Shermer, Michael (2002). "This View of Science" (http:/ / www. stephenjaygould. org/ library/ shermer_sjgould. pdf) Social Studies of Science 32 (4): 496 [77] http:/ / www. stephenjaygould. org/ bibliography. html [78] http:/ / books. google. com. au/ books?id=z8tDIQi5HaUC& printsec=frontcover& dq=%22Stephen+ Jay+ Gould%22& hl=en& ei=RUHWS7fdO4uK6gP7sPmEAw& sa=X& oi=book_result& ct=result& resnum=8& ved=0CFgQ6AEwBw#v=onepage& q& f=false [79] http:/ / books. google. com/ ?id=k4ykb5el47kC& printsec=frontcover& dq=%22'The+ Evolution+ of+ Gryphaea%22& q [80] http:/ / books. google. com. au/ books?id=UeGXJ5b8Ph0C& printsec=frontcover& dq=%22Stephen+ Jay+ Gould%22& hl=en& ei=30LWS8SlC4Hk7AOF-qH8Ag& sa=X& oi=book_result& ct=result& resnum=4& ved=0CEoQ6AEwAw#v=onepage& q& f=false [81] http:/ / books. google. com. au/ books?id=nhIl7e61WOUC& printsec=frontcover& dq=%22Stephen+ Jay+ Gould%22& hl=en& ei=30LWS8SlC4Hk7AOF-qH8Ag& sa=X& oi=book_result& ct=result& resnum=1& ved=0CDsQ6AEwAA#v=onepage& q& f=false [82] http:/ / palaeo-electronica. org/ 2007_3/ books/ equal. htm [83] http:/ / palaeo-electronica. org/ 2001_1/ books/ life. pdf

General references: • Dawkins, Richard (1998). Unweaving the Rainbow: Science, Delusion and the Appetite for Wonder (http:// books.google.com/books?id=ZudTchiioUoC). Boston: Houghton Mifflin Harcourt. ISBN 0395883822. • Richard C. Lewontin sums up Gould's career in an obituary (http://www.findarticles.com/p/articles/ mi_m1132/is_6_54/ai_94142087) from findarticles.com • "Darwinian Fundamentalism" (http://www.nybooks.com/articles/1151) – Gould's response to Daniel Dennett and other critics, from the The New York Review of Books

External links • Excerpts from Gould Lectures at Stanford University (http://prelectur.stanford.edu/lecturers/gould/) • Stephen Jay Gould papers (http://www.oac.cdlib.org/findaid/ark:/13030/kt229036tr) at Stanford University Libraries • Stephen Jay Gould (http://www.charlierose.com/search/?text=Stephen+Jay+Gould) Charlie Rose interviews • Unofficial Stephen Jay Gould Archive (http://www.stephenjaygould.org/)

171

Article Sources and Contributors

Article Sources and Contributors Sociobiology  Source: http://en.wikipedia.org/w/index.php?oldid=580289066  Contributors: 217.98.151.xxx, 2404:C800:9003:8:0:0:0:12, 4ever lovin jah, AaronSw, Abu badali, Acadēmica Orientālis, Acanon, Adbarnhart, Adoniscik, Alan Liefting, Alan Peakall, Alienus, AlisterBarclay, Andrew Levine, Andycjp, Aprock, Arnold90, Arthena, Ayeaye, Azcolvin429, Bdmccray, Belltower, Bendroz, Berkeley99, Biólogo conservacionista, Br43402, Brusegadi, Brya, Bueller 007, Burningsquid, CNicol, Calaschysm, Canto2009, Cdc, Ceoil, Chealer, Chowbok, Chris Roy, Circeus, Civeel, CommonsDelinker, Conversion script, Countered, Cretog8, Cyan, Cybercobra, D6, DCDuring, DKolle, Dajamtee, Danmateju, Davewild, Delirium, Designquest10, Detsif, Dirk P Broer, Dissembly, Dj Capricorn, Dmacw6, DonSiano, Dsandeford, Duncharris, EPM, East718, Ed Poor, Eddie tejeda, Ekserevnitis, Erianna, Extremophile, FHifzaMBuhari, Fang 23, FrankCostanza, GTBacchus, Gadfium, Gaurav, Gdvorsky, Gjbloom, GoingBatty, Graham87, Grapplequip, Guillaume2303, Gwernol, Hasanisawi, Hauganm, Hermeneus, Hmains, HowieKor, Iamcuriousblue, Itai, Jabowery, Jbshaldanelion, Jeff G., JenLouise, Jeremy68, John Vandenberg, Johnuniq, Jorge Stolfi, Jossi, Juanycueva, KYPark, Kaarel, Karl-Henner, Kelseymetzger, Kernsters, Killdeer, Kirstyc90, Kris Schnee, Kruwi, Kukini, Lcgarcia, Leadwind, Levineps, Lexor, Lightmouse, Livingrm, Looxix, Lucidish, MagneticFlux, Maikstrik, Malcohol, Maprovonsha172, Maunus, Mav, Mchavez, Meclee, Meco, Michael Johnson, Michaeldsuarez, Mike Dillon, Mikewax, Minesweeper, Ming the Merciless, Miradre, MistyMorn, Mlaurenti, Moralmo, Mr.troughton, Mydreamsparrow, Nabeth, Naddy, Nectarflowed, NickCT, Nikkimaria, Noisy, Ok!, Onemanbandbjm, Orioane, Ostensibly1, Pallab1234, Pearle, Peregrine981, Pete.Hurd, Philiptdotcom, Piotrus, Polisher of Cobwebs, Presearch, Prothonotar, Psychologesetz, Puckly, Purplefeltangel, Rankiri, Ray Van De Walker, RedHouse18, RedWolf, Richard001, Rjwilmsi, Rls, Rmallott1, Roadrunner, RockMagnetist, Rovertime, Ruhrjung, Saucepan, SightWatcher, Simon_J_Kissane, Siodine, Skittleys, Spearhead, Steinsky, Steverapaport, Stillnotelf, Stimulieconomy, Stobierskik1, T@Di, Taragui, Taw, Tbhotch, Tequilastoner, Thalustan, That Guy, From That Show!, The Anome, The Devil's Advocate, Tide rolls, Tim Carter Pedrera, TimShell, Timo Honkasalo, Tito-, Tomsega, Topbanana, Tothebarricades.tk, Trucquergirl420, Uruiamme, User2004, Victimofleisure, Vincent Moon, W Pete Welch, Wapondaponda, Watonga, Wavelength, Waywethin, We used to sit, WeijiBaikeBianji, Wetman, White Shadows, Wik, Wikiborg, Wintonian, Wobble, WpZurp, Zephirum, Zoologyteacher, Zzuuzz, ‫ בלנק‬,.‫אריה ה‬, ‫ﻣﺤﺒﻮﺏ ﻋﺎﻟﻢ‬, 관인생략, 245 anonymous edits Biocultural anthropology  Source: http://en.wikipedia.org/w/index.php?oldid=558490603  Contributors: BMF81, Canto2009, Closedmouth, Coren, Cremepuff222, Elrodriguez, Howellbm, Kelsoj1, Kered77, Khongsdier, Ktr101, Newman Luke, Newsroom hierarchies, PB54, Pandelver, Pigman, Piotrus, Pwjb, Rimapacowa, Semmler, Skysmith, Whoosit, Zenauberon, 10 anonymous edits Biosocial theory  Source: http://en.wikipedia.org/w/index.php?oldid=577034892  Contributors: Akjar13, Altenmann, Benrl27, Ciche, Firsfron, Flyingtoaster1337, GeorgeLouis, GoingBatty, Hburg, Iridescent, JohnChrysostom, Kungfuadam, Mattisse, Michael Devore, Piotrus, RDBrown, Retired username, SchreiberBike, Silver seren, WhatamIdoing, Zeraeph, 17 anonymous edits Cultural selection theory  Source: http://en.wikipedia.org/w/index.php?oldid=568530753  Contributors: Ambreena, Andrewaskew, Arnold90, Arundhati bakshi, Benplowman, Chris Capoccia, Daniel Cordoba-Bahle, Fheyligh, Ghobadi07, Harro5, Jac16888, JenLouise, John of Reading, Kaizhangchina, Kkmurray, Legion fi, Loadmaster, Mak68, Nabz phoenix, Nihil novi, Pandelver, PsyGirl29, Saarahb19, Scrycer, Sir Daniel M. A. 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Dalism, Vanished user, Vanished user 19794758563875, Vegetator, Veliger, VolatileChemical, Vsmith, WAS 4.250, Waldow, Warfwar3, Wavelength, Weightofair, Whatiguana, Wilke, Willerror, Woohookitty, Wotnow, Yahel Guhan, Yerpo, Zfr, Солярист, ‫ﮐﺎﺷﻒ ﻋﻘﯿﻞ‬, 小 為, 181 anonymous edits Evolutionary developmental psychology  Source: http://en.wikipedia.org/w/index.php?oldid=553327535  Contributors: Circeus, Craigjclemson, Cybercobra, EPM, Farrajak, Gnaussor, H.sanat, JTBurman, Jackhynes, Jalalzahir, Lova Falk, Mattisse, Memills, Mpluess, Nabeth, Pengortm, Pete.Hurd, Sardanaphalus, Semmler, Twirligig, Viriditas, We used to sit, 26 anonymous edits Evolutionary neuroscience  Source: http://en.wikipedia.org/w/index.php?oldid=565506691  Contributors: Adashiel, Andycjp, Antaeus Feldspar, Bridwater, Coelacan, Cogpsych, Daniel Mietchen, Dbrodbeck, Edhubbard, Epastore, Fences and windows, Harizotoh9, Igiffin, Jinhl, KrisK, Looie496, Lord Patrick, Lova Falk, Malcolma, Megan1967, Metanoid, Mgiganteus1, Neuromere, Ombudsman, P'tit Pierre, Semmler, Tequilastoner, Utku Tanrivere, WikHead, 24 anonymous edits Evolutionary psychology  Source: http://en.wikipedia.org/w/index.php?oldid=578011735  Contributors: 2004-12-29T22:45Z, Adhib, Adoniscik, Aeusoes1, Alan Peakall, Alansohn, Allinthebrain, Altg20April2nd, AmigoNico, Andonic, AndyTheGrump, Arbor, Arkuat, Arru, Arthur Rubin, Aschwa5, AstroHurricane001, Averylongdream, Aviad.Aharonov, Azcolvin429, BBrihem, BKalesti, Bahhasg, Banzai6666, Barbaking, BarryCull, Bart133, BashBrannigan, Bbb23, Beetstra, Ben Ben, Betterworld, Bfinn, Blingbling1000, Bmcirillo, Bmeacham, Bodinagamin, Bonnieguan, Bonzsu, BrainMagMo, Bryan Derksen, Bueller 007, CNicol, CaroleHenson, Cashtastrophy, Cat Cubed, Cavrdg, Cedibciz, Cesar Tort, Cgingold, Charles Matthews, Chealer, Chet Ubetcha, Chevapdva, Chhandama, Chris.Cole.DC, ChrisGualtieri, Circeus, Clicketyclack, Closedmouth, Coffee, Conversion script, Correctia, Corrigann, Courcelles, CourtChru., Craptree, 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SimonP, Sir Nicholas de Mimsy-Porpington, Slakr, Slrubenstein, Smalljim, Snuffkin, Sonicyouth86,

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Article Sources and Contributors SpuriousQ, Star767, Stegosaur, Steinsky, Stephenb, Stevenmitchell, Stimpster, Stragermont, Sun Creator, Superweapons, Surv1v4l1st, Swidbert R Ott, Taw, Tebbe en, Tectonicura, Temtem, ThatPeskyCommoner, The Anome, The Devil's Advocate, The Thing That Should Not Be, The stuart, TheaterMarine, Thiago90ap, Thorell, Tikiwont, Tim bates, Tjnelso1, Tmh, Tomwsulcer, Tony1, Topbanana, Tothebarricades.tk, Totorotroll, Tubby, Ubiq, Urthogie, User2004, Vaughan, Versageek, Viriditas, Vlad24601, VladimirGolovin, Vrenator, W Pete Welch, WLU, Waldow, Wangi, Wapondaponda, Washburnmav, Waywethin, Whatiguana, Whispering, WikiDan61, Will Beback, Wingnut99, Winston Trechane, Wlambeth, Wmahan, Wmspriggs2008, Woodsrock, Woohookitty, Wyang, Xezbeth, Yakushima, Yath, Yeohac, Yworo, Zooeycat, Zoostin, Zoz, ‫ﮐﺎﺷﻒ ﻋﻘﯿﻞ‬, 㓟, 596 anonymous edits Human behavioral ecology  Source: http://en.wikipedia.org/w/index.php?oldid=563310998  Contributors: Bissinger, Cgingold, Chealer, Circeus, Cybercobra, DASonnenfeld, Dbrodbeck, Delusion23, Denispir, EPM, FD Johnston, Googlesalot, Hispaniensis, Hunarian, Jandalhandler, Jose nnnn, Pandelver, Richard001, Rolando, Semmler, Shot357, Shrumster, Slightsmile, Some standardized rigour, Tribaal, Vanished user 19794758563875, Viriditas, 23 anonymous edits Prisoner's dilemma  Source: http://en.wikipedia.org/w/index.php?oldid=579861132  Contributors: 129.186.19.xxx, 12mmclean, 2001:628:404:2F:706E:D02F:4794:15F2, 2607:F470:E:C:E543:4015:699E:622D, 7&6=thirteen, 848219pineapple, AdRock, Adam Conover, Aherunar, Akerans, Alai, Alaiche, AlanM1, Alex3917, Alison, AllanLee, AllanR, Alsandro, Amead, Americanhero, Andrejj, Anetode, Angelbo, Annielogue, Antimatter15, Arco74, Artoasis, Arvindn, Asaadi, Ascorbic, Ashutt92, Atlant, Audacity, AxelBoldt, Az7997, Babij, Baccyak4H, Backfromquadrangle, Badger Drink, Bando26, Barrrower, Behavioralethics, Benjamin H-W, Bhause, BigK HeX, Bigturtle, BillyH, Billyd992, Binkyuk, Blackmetalbaz, BlueNovember, 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