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  1. Marshall Abrams (2009). Fitness “Kinematics”: Biological Function, Altruism, and Organism–Environment Development. Biology and Philosophy 24 (4):487-504.
    It’s recently been argued that biological fitness can’t change over the course of an organism’s life as a result of organisms’ behaviors. However, some characterizations of biological function and biological altruism tacitly or explicitly assume that an effect of a trait can change an organism’s fitness. In the first part of the paper, I explain that the core idea of changing fitness can be understood in terms of conditional probabilities defined over sequences of events in an organism’s life. The result (...)
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  2. Mahesh Ananth (2005). Psychological Altruism Vs. Biological Altruism: Narrowing the Gap with the Baldwin Effect. Acta Biotheoretica 53 (3).
    This paper defends the position that the supposed gap between biological altruism and psychological altruism is not nearly as wide as some scholars (e.g., Elliott Sober) insist. Crucial to this defense is the use of James Mark Baldwin's concepts of “organic selection”and “social heredity” to assist in revealing that the gap between biological and psychological altruism is more of a small lacuna. Specifically, this paper argues that ontogenetic behavioral adjustments, which are crucial to individual survival and reproduction, are also crucial (...)
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  3. Matthew Barrett & Peter Godfrey-smith (2002). Group Selection, Pluralism, and the Evolution of Altruism. [REVIEW] Philosophy and Phenomenological Research 65 (3):685–691.
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  4. Jonathan Birch (2014). Gene Mobility and the Concept of Relatedness. Biology and Philosophy 29 (4):445-476.
    Cooperation is rife in the microbial world, yet our best current theories of the evolution of cooperation were developed with multicellular animals in mind. Hamilton’s theory of inclusive fitness is an important case in point: applying the theory in a microbial setting is far from straightforward, as social evolution in microbes has a number of distinctive features that the theory was never intended to capture. In this article, I focus on the conceptual challenges posed by the project of extending Hamilton’s (...)
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  5. Jonathan Birch (2014). How Cooperation Became the Norm. [REVIEW] Biology and Philosophy 29 (3):433-444.
    Most of the contributions to Cooperation and Its Evolution grapple with the distinctive challenges presented by the project of explaining human sociality. Many of these puzzles have a ‘chicken and egg’ character: our virtually unparalleled capacity for large-scale cooperation is the product of psychological, behavioural, and demographic changes in our recent evolutionary history, and these changes are linked by complex patterns of reciprocal dependence. There is much we do not yet understand about the timing of these changes, and about the (...)
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  6. Jonathan Birch (2014). Hamilton's Rule and Its Discontents. British Journal for the Philosophy of Science 65 (2):381-411.
    In an incendiary 2010 Nature article, M. A. Nowak, C. E. Tarnita, and E. O. Wilson present a savage critique of the best-known and most widely used framework for the study of social evolution, W. D. Hamilton’s theory of kin selection. More than a hundred biologists have since rallied to the theory’s defence, but Nowak et al. maintain that their arguments ‘stand unrefuted’. Here I consider the most contentious claim Nowak et al. defend: that Hamilton’s rule, the core explanatory principle (...)
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  7. Jonathan Birch (2013). Samir Okasha and Ken Binmore (Eds) Evolution and Rationality: Decisions, Cooperation, and Strategic Behaviour. [REVIEW] British Journal for the Philosophy of Science 64 (3):669-673.
  8. Jonathan Birch (2012). Social Revolution. [REVIEW] Biology and Philosophy 27 (4):571-581.
    Andrew Bourke’s Principles of Social Evolution identifies three stages that characterize an evolutionary transition in individuality and deploys inclusive fitness theory to explain each stage. The third stage, social group transformation, has hitherto received relatively little attention from inclusive fitness theorists. In this review, I first discuss Bourke’s “virtual dominance” hypothesis for the evolution of the germ line. I then contrast Bourke’s inclusive fitness approach to the major transitions with the multi-level approach developed by Richard Michod, Samir Okasha and others. (...)
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  9. Jonathan Birch (2012). Collective Action in the Fraternal Transitions. Biology and Philosophy 27 (3):363-380.
    Inclusive fitness theory was not originally designed to explain the major transitions in evolution, but there is a growing consensus that it has the resources to do so. My aim in this paper is to highlight, in a constructive spirit, the puzzles and challenges that remain. I first consider the distinctive aspects of the cooperative interactions we see within the most complex social groups in nature: multicellular organisms and eusocial insect colonies. I then focus on one aspect in particular: the (...)
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  10. Jonathan Birch & James A. R. Marshall (2014). Queller's Separation Condition Explained and Defended. American Naturalist 184 (4):531-540.
    The theories of inclusive fitness and multilevel selection provide alternative perspectives on social evolution. The question of whether these perspectives are of equal generality remains a divisive issue. In an analysis based on the Price equation, Queller argued (by means of a principle he called the separation condition) that the two approaches are subject to the same limitations, arising from their fundamentally quantitative-genetical character. Recently, van Veelen et al. have challenged Queller’s results, using this as the basis for a broader (...)
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  11. Jonathan Birch & Samir Okasha (forthcoming). Kin Selection and Its Critics. BioScience.
    Hamilton’s theory of kin selection is the best-known framework for understanding the evolution of social behavior but has long been a source of controversy in evolutionary biology. A recent critique of the theory by Nowak, Tarnita, and Wilson sparked a new round of debate, which shows no signs of abating. In this overview, we highlight a number of conceptual issues that lie at the heart of the current debate. We begin by emphasizing that there are various alternative formulations of Hamilton’s (...)
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  12. Christopher Boehm (1999). The Natural Selection of Altruistic Traits. Human Nature 10 (3):205-252.
    Proponents of the standard evolutionary biology paradigm explain human “altruism” in terms of either nepotism or strict reciprocity. On that basis our underlying nature is reduced to a function of inclusive fitness: human nature has to be totally selfish or nepotistic. Proposed here are three possible paths to giving costly aid to nonrelatives, paths that are controversial because they involve assumed pleiotropic effects or group selection. One path is pleiotropic subsidies that help to extend nepotistic helping behavior from close family (...)
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  13. Alan carter (2005). Evolution and the Problem of Altruism. Philosophical Studies 123 (3):213-230.
    Genuine altruism would appear to be incompatible with evolutionary theory. And yet altruistic behavior would seem to occur, at least on occasion. This article first considers a game-theoretical attempt at solving this seeming paradox, before considering agroup selectionist approach. Neither approach, as they stand, would seem to render genuine, as opposed to reciprocal, altruism compatible with the theory of evolution. The article concludes by offering an alternative game-theoretical solution to the problem of altruism.
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  14. Gustavo Cevolani & Roberto Festa (2012). Giochi di altruismo. L'approccio evoluzionistico alla cooperazione umana. In Matt Ridley (ed.), Le Origini della Virtù. IBL Libri. 7--38.
    This is the introductory essay to the Italian translation of Matt Ridley's "The origins of virtue", surveying the game-theoretic and evolutionary approaches to the emergence and evolution of cooperation and altruism.
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  15. Christine Clavien & Michel Chapuisat (2013). Altruism Across Disciplines: One Word, Multiple Meanings. Biology and Philosophy 28 (1):125-140.
    Altruism is a deep and complex phenomenon that is analysed by scholars of various disciplines, including psychology, philosophy, biology, evolutionary anthropology and experimental economics. Much confusion arises in current literature because the term altruism covers variable concepts and processes across disciplines. Here we investigate the sense given to altruism when used in different fields and argumentative contexts. We argue that four distinct but related concepts need to be distinguished: (a) psychological altruism , the genuine motivation to improve others’ interests and (...)
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  16. Christine Clavien & Michel Chapuisat (2012). Altruism - a Philosophical Analysis. eLS.
    Altruism is a malleable notion that is understood differently in various disciplines. The common denominator of most definitions of altruism is the idea of unidirectional helping behaviour. However, a closer examination reveals that the term altruism sometimes refers to the outcomes of a helping behaviour for the agent and its neighbours – i.e. reproductive altruism – and sometimes to what motivates the agent to help others – i.e. psychological altruism. Since these perspectives on altruism are crucially different, it is important (...)
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  17. Jay Elliott (2011). Stag Hunts and Committee Work: Cooperation and the Mutualistic Paradigm. [REVIEW] Review of Philosophy and Psychology 2 (2):245-260.
    Contemporary philosophers and psychologists seek the roots of ethically sound forms of behavior, including altruism and a sense of fairness, in the basic structure of cooperative action. I argue that recent work on cooperation in both philosophy and psychology has been hampered by what I call “the mutualistic paradigm.” The mutualistic paradigm treats one kind of cooperative situation—what I call a “mutualistic situation”—as paradigmatic of cooperation in general. In mutualistic situations, such as the primeval stag hunt described by Brian Skyrms, (...)
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  18. Peter Gildenhuys (2003). The Evolution of Altruism: The Sober/Wilson Model. Philosophy of Science 70 (1):27-48.
    In what follows, I critique the interpretation that Sober and Wilson offer of their group selection model in Unto Others. Sober and Wilson mistakenly claim that their model operates as an example of Simpson's paradox and defend an interpretation of their model according to which groups are operated upon by natural selection. In the place of their interpretation, I offer one that parallels the mathematical calculation of the model's outcome and does not depend on the postulation of a force of (...)
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  19. Peter Godfrey-Smith (2002). Group Selection, Pluralism, and the Evolution of Altruism. [REVIEW] Philosophy and Phenomenological Research 65 (3):685 - 691.
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  20. Michael Gurven (2004). To Give and to Give Not: The Behavioral Ecology of Human Food Transfers. Behavioral and Brain Sciences 27 (4):543-559.
    The transfer of food among group members is a ubiquitous feature of small-scale forager and forager-agricultural populations. The uniqueness of pervasive sharing among humans, especially among unrelated individuals, has led researchers to evaluate numerous hypotheses about the adaptive functions and patterns of sharing in different ecologies. This article attempts to organize available cross-cultural evidence pertaining to several contentious evolutionary models: kin selection, reciprocal altruism, tolerated scrounging, and costly signaling. Debates about the relevance of these models focus primarily on the extent (...)
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  21. Oren Harman (2011). Helical Biography and the Historical Craft: The Case of Altruism and George Price. [REVIEW] Journal of the History of Biology 44 (4):671 - 691.
    The life of George Price (1922-1975), the eccentric polymath genius and father of the Price equation, is used as a prism and counterpoint through which to consider an age-old evolutionary conundrum: the origins of altruism. This biographical project, and biography and history more generally, are considered in terms of the possibility of using form to convey content in particular ways. Closer to an art form than a science, this approach to scholarship presents both a unique challenge and promise.
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  22. William Harms (1999). Biological Altruism in Hostile Environments. Complexity 5 (2):23-28.
    The evolution of economic altruism is one of the most vigorous areas of study at the intersection of biology, economics, and philosophy. The basic problem is easily understood. Biological organisms, be they people or paramecia, have ample opportunity to confer benefits on others at relatively low cost to themselves. If conferring such benefits becomes common, the overall productivity of the population in which it occurs is increased. Presumably, there is no advantage to refusing such benefits, but it is also the (...)
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  23. Joseph Heath (2002). The Robustness of Altruism as an Evolutionary Strategy. Biology and Philosophy 17 (4):567-590.
    Kin selection, reciprocity and group selection are widely regarded as evolutionary mechanisms capable of sustaining altruism among humans andother cooperative species. Our research indicates, however, that these mechanisms are only particular examples of a broader set of evolutionary possibilities.In this paper we present the results of a series of simple replicator simulations, run on variations of the 2–player prisoner's dilemma, designed to illustrate the wide range of scenarios under which altruism proves to be robust under evolutionary pressures. The set of (...)
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  24. Hope Hollocher, Agustin Fuentes, Charles H. Pence, Grant Ramsey, Daniel John Sportiello & Michelle M. Wirth (2011). On the Origin of Stories: Evolution, Cognition, and Fiction. [REVIEW] Quarterly Review of Biology 86 (2):137-138.
  25. Klaus Jaffe (2004). Altruism, Altruistic Punishment and Social Investment. Acta Biotheoretica 52 (3).
    The concept of altruism is used in very different forms by computer scientists,economists, philosophers, social scientists, psychologists and biologists. Yet, in order to be useful in social simulations, the concept altruism requires a more precise meaning. A quantitative formulation is proposed here, based on the cost/benefit analysis of the altruist and of society at large. This formulation is applied in the analysis of the social dynamic working of behaviors that have been called altruistic punishments, using the agent based computer model (...)
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  26. Benjamin Kerr, Peter Godfrey-Smith & Marcus W. Feldman, What is Altruism?
    Altruism is generally understood to be behavior that benefits others at a personal cost to the behaving individual. However, within evolutionary biology, different authors have interpreted the concept of altruism differently, leading to dissimilar predictions about the evolution of altruistic behavior. Generally, different interpretations diverge on which party receives the benefit from altruism and on how the cost of altruism is assessed. Using a simple trait-group framework, we delineate the assumptions underlying different interpretations and show how they relate to one (...)
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  27. Philip Kitcher (1993). The Evolution of Human Altruism. Journal of Philosophy 60 (10):497-516.
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  28. B. Linsky & M. Mathen (eds.) (1988). New Essays on Philosophy and Biology (Canadian Journal of Philosophy Supp. Vol. 14). University of Calgary Press.
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  29. Randolph M. Nesse (2007). Runaway Social Selection for Displays of Partner Value and Altruism. Biological Theory 2 (2):143-155.
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  30. Leonard Nunney (2000). Altruism, Benevolence and Culture. Journal of Consciousness Studies 7 (1-2):1-2.
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  31. Samir Okasha (2002). Genetic Relatedness and the Evolution of Altruism. Philosophy of Science 69 (1):138-149.
    In their recent book, Elliott Sober and David Wilson (1998) argue that evolutionary biologists have wrongly regarded kinship as the exclusive means by which altruistic behavior can evolve, at the expense of other mechanisms. I argue that Sober and Wilson overlook certain genetical considerations which suggest that kinship is likely to be a more powerful means for generating complex altruistic adaptations than the alternative mechanisms they propose.
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  32. Charles H. Pence & Lara Buchak (2012). Oyun: A New, Free Program for Iterated Prisoner’s Dilemma Tournaments in the Classroom. Evolution Education and Outreach 5 (3):467-476.
    Evolutionary applications of game theory present one of the most pedagogically accessible varieties of genuine, contemporary theoretical biology. We present here Oyun (OY-oon, http://charlespence.net/oyun), a program designed to run iterated prisoner’s dilemma tournaments, competitions between prisoner’s dilemma strategies developed by the students themselves. Using this software, students are able to readily design and tweak their own strategies, and to see how they fare both in round-robin tournaments and in “evolutionary” tournaments, where the scores in a given “generation” directly determine contribution (...)
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  33. John D. Pitts (1998). The Discovery of Metabolic Co-Operation. Bioessays 20 (12):1047-1051.
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  34. Angela Potochnik (2012). Modeling Social and Evolutionary Games. Studies in History and Philosophy of Science Part C 43 (1):202-208.
    When game theory was introduced to biology, the components of classic game theory models were replaced with elements more befitting evolutionary phenomena. The actions of intelligent agents are replaced by phenotypic traits; utility is replaced by fitness; rational deliberation is replaced by natural selection. In this paper, I argue that this classic conception of comprehensive reapplication is misleading, for it overemphasizes the discontinuity between human behavior and evolved traits. Explicitly considering the representational roles of evolutionary game theory brings to attention (...)
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  35. Howard Rachlin (2002). Altruism and Selfishness. Behavioral and Brain Sciences 25 (2):239-250.
    Many situations in human life present choices between (a) narrowly preferred particular alternatives and (b) narrowly less preferred (or aversive) particular alternatives that nevertheless form part of highly preferred abstract behavioral patterns. Such alternatives characterize problems of self-control. For example, at any given moment, a person may accept alcoholic drinks yet also prefer being sober to being drunk over the next few days. Other situations present choices between (a) alternatives beneficial to an individual and (b) alternatives that are less beneficial (...)
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  36. I. I. I. Rolston (2011). SuperCooperators: Altruism, Evolution, and Why We Need Each Other to Succeed by Martin A. Nowak, with Roger Highfield. Zygon 46 (4):1003-1005.
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  37. Alejandro Rosas (2010). Beyond Inclusive Fitness? On A Simple And General Explanation For The Evolution of Altruism. Philosophy and Theory in Biology 2 (20130604).
    Altruism is a central concept in evolutionary biology. Evolutionary biologists still disagree about its meaning (E.O. Wilson 2005; Fletcher et al. 2006; D.S. Wilson 2008; Foster et al. 2006a, b; West et al. 2007a, 2008). Semantic disagreement appears to be quite robust and not easily overcome by attempts at clarification, suggesting that substantive conceptual issues lurk in the background. Briefly, group selection theorists define altruism as any trait that makes altruists losers to selfish traits within groups, and makes groups of (...)
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  38. Alejandro Rosas (2008). Multilevel Selection and Human Altruism. Biology and Philosophy 23 (2):205-215.
    Views on the evolution of altruism based upon multilevel selection on structured populations pay little attention to the difference between fortuitous and deliberate processes leading to assortative grouping. Altruism may evolve when assortative grouping is fortuitously produced by forces external to the organism. But when it is deliberately produced by the same proximate mechanism that controls altruistic responses, as in humans, exploitation of altruists by selfish individuals is unlikely and altruism evolves as an individually advantageous trait. Groups formed with altruists (...)
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  39. Alejandro Rosas (2005). La Moral y Sus Sombras: La Racionalidad Instrumental y la Evolución de Las Normas de Equidad (Morality and its Shadows: Instrumental Rationality and the Evolution of Fairness Norms). Critica 37 (110):79 - 104.
    Los sociobiólogos han defendido una posición "calvinista" que se resume en la siguiente fórmula: si la selección natural explica las actitudes morales, no hay altruismo genuino en la moral; si la moral es altruista, entonces la selección natural no puede explicarla. En este ensayo desenmascaro los presupuestos erróneos de esta posición y defiendo que el altruismo como equidad no es incompatible con la selección natural. Rechazo una concepción hobbesiana de la moral, pero sugiero su empleo en la interpretación de la (...)
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  40. Mary Rousseau (1997). The Evolution of Altruism and the Ordering of Love. American Catholic Philosophical Quarterly 71 (1):133-136.
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  41. Hannes Rusch, A Threshold for Biological Altruism in Public Goods Games Played in Groups Including Kin. MAGKS Discussion Paper Series in Economics.
    Phenomena like meat sharing in hunter-gatherers, altruistic self-sacrifice in intergroup conflicts, and contribution to the production of public goods in laboratory experiments have led to the development of numerous theories trying to explain human prosocial preferences and behavior. Many of these focus on direct and indirect reciprocity, assortment, or (cultural) group selection. Here, I investigate analytically how genetic relatedness changes the incentive structure of that paradigmatic game which is conventionally used to model and experimentally investigate collective action problems: the public (...)
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  42. Hannes Rusch (2014). The Evolutionary Interplay of Intergroup Conflict and Altruism in Humans: A Review of Parochial Altruism Theory and Prospects for its Extension. Proceedings of the Royal Society of London B 281 (1794): 20141539.
    Drawing on an idea proposed by Darwin, it has recently been hypothesised that violent intergroup conflict might have played a substantial role in the evolution of human cooperativeness and altruism. The central notion of this argument, dubbed ‘parochial altruism’, is that the two genetic or cultural traits, aggressiveness against out-groups and cooperativeness towards the in-group, including self-sacrificial altruistic behaviour, might have coevolved in humans. This review assesses the explanatory power of current theories of ‘parochial altruism’. After a brief synopsis of (...)
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  43. Hannes Rusch (2014). The Two Sides of Warfare: An Extended Model of Altruistic Behavior in Ancestral Human Intergroup Conflict. Human Nature 25 (3):359-377.
    Building on and partially refining previous theoretical work, this paper presents an extended simulation model of ancestral warfare. This model (1) disentangles attack and defense, (2) tries to differentiate more strictly between selfish and altruistic efforts during war, (3) incorporates risk aversion and deterrence, and (4) pays special attention to the role of brutality. Modeling refinements and simulation results yield a differentiated picture of possible evolutionary dynamics. The main observations are: (i) Altruism in this model is more likely to evolve (...)
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  44. Hannes Rusch (2013). Asymmetries in Altruistic Behavior During Violent Intergroup Conflict. Evolutionary Psychology 11 (5):973-993.
    Recent theoretical and experimental investigations of altruistic behavior in intergroup conflict in humans frequently make use of the assumption that warfare can be modeled as a symmetrical n-person prisoner’s dilemma, abstracting away the strategic differences between attack and defense. In contrast, some empirical studies on intergroup conflict in hunter-gatherer societies and chimpanzees indicate that fitness relevant risks and potential benefits of attacks and defenses might have differed substantially under ancestral conditions. Drawing on these studies, it is hypothesized that the success (...)
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  45. Tom Settle (1993). 'Fitness' and 'Altruism': Traps for the Unwary, Bystander and Biologist Alike. [REVIEW] Biology and Philosophy 8 (1):61-83.
    At one level, this paper is a lament and a warning. I lament biologists borrowing well-known terms and then drastically and awkwardly changing their meanings, and I warn about the mischief this does. Biology''s public image is at stake, as is its general usefulness. At another level, I attempt to clarify the misnamed concepts, beyond what has been achieved in recent philosophical writings. This helps to account for the mischief, and to see how it might be avoidable. But the most (...)
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  46. Pouwel Slurink (1993). Ecological Dominance and the Final Sprint in Hominid Evolution. Human Evolution.
    In contrast to many other models of human evolution the "balance of power" theory of Alexander has a clear answer to the question why a runaway selection process for unique social and moral capacities occurred in our ancestry only and not in other species: "ecological dominance" is hypothesized to have diminished the effects of "extrinsic" forces of natural selection such that within-species, intergroup competition increased (Alexander, 1989). Alexander seems to be wrong, however, in his claim that already the common HUCHIBO (...)
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  47. Harry Smit (2010). The Development of Altruistic Behavior Out of Reactive Crying. Biological Theory 5 (1):79-86.
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  48. Thomas S. Smith & Gregory T. Stevens (2002). Hyperstructures and the Biology of Interpersonal Dependence: Rethinking Reciprocity and Altruism. Sociological Theory 20 (1):106-130.
    Fluctuations in endogenous opioid activity in the brain, controlled under ordinary conditions by attachment, are capable of producing patterns of dependence in social behavior resembling those appearing in substance abusers. Withdrawal symptoms arising in relation to these fluctuations, short of producing dependence, ordinarily fuel everyday social interaction, and interaction then serves to modulate opioid activity within a range associated with comfort. Comfort-constraints in this sense operate in all settings of social interaction, part of an innate caregiving mechanism conserved by evolution (...)
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  49. Elliott Sober (1992). The Evolution of Altruism: Correlation, Cost, and Benefit. [REVIEW] Biology and Philosophy 7 (2):177-187.
    A simple and general criterion is derived for the evolution of altruism when individuals interact in pairs. It is argued that the treatment of this problem in kin selection theory and in game theory are special cases of this general criterion.
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  50. Elliott Sober (1988). What is Evolutionary Altruism? In B. Linsky & M. Mathen (eds.), New Essays on Philosophy and Biology (Canadian Journal of Philosophy Supp. Vol. 14). University of Calgary Press.
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