Group selection

We need scarcely add that the contemplation in natural science of a wider domain than the actual leads to a far better understanding of the actual.' (p. 267,The, Nature of the Physical World.)x PREFACE evolutionary theory was thus chiefly retrogressive, the mighty body of Mendelian researches throughout the world has evidently out- grown the fallacies with which it was at first fostered.As a pioneer of genetics he has done more than enough to expiate the rash polemics of his early writings.To treat Natural Selection as an agency based independently on its own foundations is not to mimimize its importance in the theory of evolution.On the contrary, as soon as we require to form opinions by other means than by comparison and analogy, such an indepen- dent deductive basis becomes a necessity.This necessity is particu- larly to be noted for mankind; since we have some knowledge of the structure of society, of human motives, and of the vital statistics of this species, the use of the deductive method can supply a more intimate knowledge of the evolutionary processes than is elsewhere possible.In addition it will be of importance for our subject to call) attention to several consequences of the principle of Natural Selection!which, since they do not consist in the adaptive modification of specific I forms, have necessarily escaped attention.The genetic phenomena of I dominance and linkage seem to offer examples of this class, the future ' investigation of which may add greatly to the scope of our subject.No efforts of mine could avail to make the book easy reading.I have endeavoured to assist the reader by giving short summaries at the ends of all chapters, except Chapter IV, which is summarized conjointly with Chapter V.Those who prefer to do so may regardChapter IV as a mathematical appendix to the corresponding part of the summary.The deductions respecting Man are strictly in- separable from the more general chapters, but have been placed together in a group commencing with Chapter VIII.I believe no one will be surprised that a large number of the points considered demand a far fuller, more rigorous, and more comprehensive treat- ment.It seems impossible that full justice should be done to the subject in this way, until there is built up a tradition of mathematical work devoted to biological problems, comparable to the researches upon which a mathematical physicist can draw in the resolution of special difficulties.

@book{doi105962bhltitle27468,
    author = "Fisher, Ronald Aylmer",
    title = "The genetical theory of natural selection",
    year = "1930",
    booktitle = "Clarendon Press eBooks",
    abstract = "We need scarcely add that the contemplation in natural science of a wider domain than the actual leads to a far better understanding of the actual.' (p. 267,The, Nature of the Physical World.)x PREFACE evolutionary theory was thus chiefly retrogressive, the mighty body of Mendelian researches throughout the world has evidently out- grown the fallacies with which it was at first fostered.As a pioneer of genetics he has done more than enough to expiate the rash polemics of his early writings.To treat Natural Selection as an agency based independently on its own foundations is not to mimimize its importance in the theory of evolution.On the contrary, as soon as we require to form opinions by other means than by comparison and analogy, such an indepen- dent deductive basis becomes a necessity.This necessity is particu- larly to be noted for mankind; since we have some knowledge of the structure of society, of human motives, and of the vital statistics of this species, the use of the deductive method can supply a more intimate knowledge of the evolutionary processes than is elsewhere possible.In addition it will be of importance for our subject to call) attention to several consequences of the principle of Natural Selection!which, since they do not consist in the adaptive modification of specific I forms, have necessarily escaped attention.The genetic phenomena of I dominance and linkage seem to offer examples of this class, the future ' investigation of which may add greatly to the scope of our subject.No efforts of mine could avail to make the book easy reading.I have endeavoured to assist the reader by giving short summaries at the ends of all chapters, except Chapter IV, which is summarized conjointly with Chapter V.Those who prefer to do so may regardChapter IV as a mathematical appendix to the corresponding part of the summary.The deductions respecting Man are strictly in- separable from the more general chapters, but have been placed together in a group commencing with Chapter VIII.I believe no one will be surprised that a large number of the points considered demand a far fuller, more rigorous, and more comprehensive treat- ment.It seems impossible that full justice should be done to the subject in this way, until there is built up a tradition of mathematical work devoted to biological problems, comparable to the researches upon which a mathematical physicist can draw in the resolution of special difficulties.",
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@article{doi101086394551,
    author = "Sturtevant, A. H.",
    title = "Essays on Evolution. II. On the Effects of Selection on Social Insects",
    year = "1938",
    journal = "The Quarterly Review of Biology",
    url = "https://doi.org/10.1086/394551",
    doi = "10.1086/394551",
    openalex = "W2091507780"
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Journal Article NATURAL SELECTION OF INDIVIDUALLY HARMFUL SOCIAL ADAPTATIONS AMONG SIBS WITH SPECIAL REFERENCE TO SOCIAL INSECTS Get access George C. Williams, George C. Williams Department of Natural Science Michigan State University Search for other works by this author on: Oxford Academic Google Scholar Doris C. Williams Doris C. Williams Department of Natural Science Michigan State University Search for other works by this author on: Oxford Academic Google Scholar Evolution, Volume 11, Issue 1, 1 March 1957, Pages 32–39, https://doi.org/10.1111/j.1558-5646.1957.tb02873.x Published: 01 March 1957 Article history Received: 24 May 1956 Published: 01 March 1957

@article{doi101111j155856461957tb02873x,
    author = "Williams, George C. and Williams, Doris C.",
    title = "NATURAL SELECTION OF INDIVIDUALLY HARMFUL SOCIAL ADAPTATIONS AMONG SIBS WITH SPECIAL REFERENCE TO SOCIAL INSECTS",
    year = "1957",
    journal = "Evolution",
    abstract = "Journal Article NATURAL SELECTION OF INDIVIDUALLY HARMFUL SOCIAL ADAPTATIONS AMONG SIBS WITH SPECIAL REFERENCE TO SOCIAL INSECTS Get access George C. Williams, George C. Williams Department of Natural Science Michigan State University Search for other works by this author on: Oxford Academic Google Scholar Doris C. Williams Doris C. Williams Department of Natural Science Michigan State University Search for other works by this author on: Oxford Academic Google Scholar Evolution, Volume 11, Issue 1, 1 March 1957, Pages 32–39, https://doi.org/10.1111/j.1558-5646.1957.tb02873.x Published: 01 March 1957 Article history Received: 24 May 1956 Published: 01 March 1957",
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@article{doi101038200623a0,
    author = "Wynne‐Edwards, V. C.",
    title = "Intergroup Selection in the Evolution of Social Systems",
    year = "1963",
    journal = "Nature",
    url = "https://doi.org/10.1038/200623a0",
    doi = "10.1038/200623a0",
    openalex = "W1975600279"
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@misc{ref1966types1,
    author = "---",
    title = "Types of group selection",
    year = "1966",
    howpublished = "Nature, v. 211, p. 870",
    note = "talkorigins\_source = {true}; raw\_reference = {---, 1966, Types of group selection: Nature, v. 211, p. 870.}"
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@article{doi1010382281218a0,
    author = "Hamilton, W D",
    title = "Selfish and Spiteful Behaviour in an Evolutionary Model",
    year = "1970",
    journal = "Nature",
    url = "https://doi.org/10.1038/2281218a0",
    doi = "10.1038/2281218a0",
    openalex = "W2144228882"
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@inproceedings{darlington1971nonmathematical2,
    author = "Darlington, P. J",
    title = "Nonmathematical models for evolution of altruism and for group selection",
    year = "1971",
    booktitle = "Proceedings of the National Academy of Sciences, v. 69, p. 293-297",
    note = "talkorigins\_source = {true}; raw\_reference = {Darlington, P. J., 1971, Nonmathematical models for evolution of altruism and for group selection: Proceedings of the National Academy of Sciences, v. 69, p. 293-297.}"
}

A model is presented to account for the natural selection of what is termed reciprocally altruistic behavior. The model shows how selection can operate against the cheater (non-reciprocator) in the system. Three instances of altruistic behavior are discussed, the evolution of which the model can explain: (1) behavior involved in cleaning symbioses; (2) warning cries in birds; and (3) human reciprocal altruism. Regarding human reciprocal altruism, it is shown that the details of the psychological system that regulates this altruism can be explained by the model. Specifically, friendship, dislike, moralistic aggression, gratitude, sympathy, trust, suspicion, trustworthiness, aspects of guilt, and some forms of dishonesty and hypocrisy can be explained as important adaptations to regulate the altruistic system. Each individual human is seen as possessing altruistic and cheating tendencies, the expression of which is sensitive to developmental variables that were selected to set the tendencies at a balance appropriate to the local social and ecological environment.

@article{doi101086406755,
    author = "Trivers, Robert",
    title = "The Evolution of Reciprocal Altruism",
    year = "1971",
    journal = "The Quarterly Review of Biology",
    abstract = "A model is presented to account for the natural selection of what is termed reciprocally altruistic behavior. The model shows how selection can operate against the cheater (non-reciprocator) in the system. Three instances of altruistic behavior are discussed, the evolution of which the model can explain: (1) behavior involved in cleaning symbioses; (2) warning cries in birds; and (3) human reciprocal altruism. Regarding human reciprocal altruism, it is shown that the details of the psychological system that regulates this altruism can be explained by the model. Specifically, friendship, dislike, moralistic aggression, gratitude, sympathy, trust, suspicion, trustworthiness, aspects of guilt, and some forms of dishonesty and hypocrisy can be explained as important adaptations to regulate the altruistic system. Each individual human is seen as possessing altruistic and cheating tendencies, the expression of which is sensitive to developmental variables that were selected to set the tendencies at a balance appropriate to the local social and ecological environment.",
    url = "https://doi.org/10.1086/406755",
    doi = "10.1086/406755",
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}

Mathematical biologists have failed to produce a satisfactory general model for evolution of altruism, i.e., of behaviors by which “altruists” benefit other individuals but not themselves; kin selection does not seem to be a sufficient explanation of nonreciprocal altruism. Nonmathematical (but mathematically acceptable) models are now proposed for evolution of negative altruism in dual-determinant and of positive altruism in tri-determinant systems. Peck orders, territorial systems, and an ant society are analyzed as examples. In all models, evolution is primarily by individual selection, probably supplemented by group selection. Group selection is differential extinction of populations. It can act only on populations preformed by selection at the individual level, but can either cancel individual selective trends (effecting evolutionary homeostasis) or supplement them; its supplementary effect is probably increasingly important in the evolution of increasingly organized populations.

@article{darlington1972nonmathematical,
    author = "Darlington, P. J.",
    title = "Nonmathematical Models for Evolution of Altruism, and for Group Selection",
    year = "1972",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = "Mathematical biologists have failed to produce a satisfactory general model for evolution of altruism, i.e., of behaviors by which “altruists” benefit other individuals but not themselves; kin selection does not seem to be a sufficient explanation of nonreciprocal altruism. Nonmathematical (but mathematically acceptable) models are now proposed for evolution of negative altruism in dual-determinant and of positive altruism in tri-determinant systems. Peck orders, territorial systems, and an ant society are analyzed as examples. In all models, evolution is primarily by individual selection, probably supplemented by group selection. Group selection is differential extinction of populations. It can act only on populations preformed by selection at the individual level, but can either cancel individual selective trends (effecting evolutionary homeostasis) or supplement them; its supplementary effect is probably increasingly important in the evolution of increasingly organized populations.",
    url = "https://doi.org/10.1073/pnas.69.2.293",
    doi = "10.1073/pnas.69.2.293",
    number = "2",
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    pages = "293-297",
    volume = "69",
    references = "doi101038218525a0, doi101073pnas6861254, openalexw2414076348"
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The place of mathematics in hypotheticodeductive processes and in biological research is discussed. (Natural) Selection is defined and described as differential elimination of performed sets at any level. Sets and acting sets are groups of units (themselves sets of smaller units) at any level that may or do interact. A pseudomathematical equation describes directional change (evolution) in sets at any level. Selection is the ram of evolution; it cannot generate, but can only direct, evolutionary energy. The energy of evolution is derived from molecular or chemical levels, is transmitted upwards through the increasingly complex sets of sets that form living systems, and is turned in directions determined by the sum of selective processes, at different levels, which may either supplement or oppose each other. All evolutionary processes conform to the pseudomathematical equation referred to above, use energy as described above, and have a P/OE (ratio of programming to open-endedness) that cannot be measured, but can be related to other P/OE values. Phylogeny and ontogeny are compared as processes af directional change with set selection. Stages in the evolution of multi-cellular individuals are suggested, and are essentially the same as stages in the evolution of some multi-individual insect societies. Thinking is considered as a part of ontogeny involving an irreversible, nonrepetitive process of set selection in the brain.

@article{doi101073pnas6951239,
    author = "Darlington, P. J.",
    title = "Nonmathematical Concepts of Selection, Evolutionary Energy, and Levels of Evolution",
    year = "1972",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = "The place of mathematics in hypotheticodeductive processes and in biological research is discussed. (Natural) Selection is defined and described as differential elimination of performed sets at any level. Sets and acting sets are groups of units (themselves sets of smaller units) at any level that may or do interact. A pseudomathematical equation describes directional change (evolution) in sets at any level. Selection is the ram of evolution; it cannot generate, but can only direct, evolutionary energy. The energy of evolution is derived from molecular or chemical levels, is transmitted upwards through the increasingly complex sets of sets that form living systems, and is turned in directions determined by the sum of selective processes, at different levels, which may either supplement or oppose each other. All evolutionary processes conform to the pseudomathematical equation referred to above, use energy as described above, and have a P/OE (ratio of programming to open-endedness) that cannot be measured, but can be related to other P/OE values. Phylogeny and ontogeny are compared as processes af directional change with set selection. Stages in the evolution of multi-cellular individuals are suggested, and are essentially the same as stages in the evolution of some multi-individual insect societies. Thinking is considered as a part of ontogeny involving an irreversible, nonrepetitive process of set selection in the brain.",
    url = "https://doi.org/10.1073/pnas.69.5.1239",
    doi = "10.1073/pnas.69.5.1239",
    openalex = "W2043292425",
    references = "darlington1972nonmathematical, doi101001jama195103670280091044, doi101001jama197103180390056032, doi101016c20130124336, doi101073pnas6861254, doi101073pnas692293, doi101126science1443619718a, doi1023072406953, doi1043249780203706299, openalexw400267313"
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Evolution of altruism by group selection involves sacrifice of some individuals, not to the "group as a whole," but to other individuals in the group. Deme-group selection may establish strictly altruistic genes in a population, but only under limited conditions, and perhaps never among vertebrates, among which apparently altruistic behaviors may always potentially benefit the altruists. Responsive-group selection is a more effective mode of evolution of altruism, conspicuous in man. Evolutionary reinforcement increases the force of selection of advantageous behaviors, including altruistic ones, by making them pleasant or rewarding. It is probably involved also in ecological habitat selection, and may be the source of many human emotions, including esthetic ones. Throwing (of stones and weapons) exemplifies both the possible importance of a difficult-to-measure evolutionary factor and the role of reinforcement; in human evolution throwing may have been decisive in food-getting and fighting, in shifting emphasis from brute force to skill, and in inducing evolution of a brain able to handle three-body geometric problems precisely and thus preadapted for more complex functions.

@article{darlington1975group,
    author = "Darlington, P J",
    title = "Group selection, altruism, reinforcement, and throwing in human evolution.",
    year = "1975",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = {Evolution of altruism by group selection involves sacrifice of some individuals, not to the "group as a whole," but to other individuals in the group. Deme-group selection may establish strictly altruistic genes in a population, but only under limited conditions, and perhaps never among vertebrates, among which apparently altruistic behaviors may always potentially benefit the altruists. Responsive-group selection is a more effective mode of evolution of altruism, conspicuous in man. Evolutionary reinforcement increases the force of selection of advantageous behaviors, including altruistic ones, by making them pleasant or rewarding. It is probably involved also in ecological habitat selection, and may be the source of many human emotions, including esthetic ones. Throwing (of stones and weapons) exemplifies both the possible importance of a difficult-to-measure evolutionary factor and the role of reinforcement; in human evolution throwing may have been decisive in food-getting and fighting, in shifting emphasis from brute force to skill, and in inducing evolution of a brain able to handle three-body geometric problems precisely and thus preadapted for more complex functions.},
    url = "https://doi.org/10.1073/pnas.72.9.3748",
    doi = "10.1073/pnas.72.9.3748",
    number = "9",
    openalex = "W1966138547",
    pages = "3748-3752",
    volume = "72",
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}

@article{doi101007bf02984178,
    author = "Scudo, Francesco M. and Ghiselin, Michael T.",
    title = "Familial selection and the evolution of social behavior",
    year = "1975",
    journal = "Journal of Genetics",
    url = "https://doi.org/10.1007/bf02984178",
    doi = "10.1007/bf02984178",
    openalex = "W2037836748",
    references = "doi101073pnas6861254"
}

(Uploaded by Plazi for the Bat Literature Project) No abstract provided.

@article{doi101126science327542,
    author = "Emlen, Stephen T. and Oring, Lewis W.",
    title = "Ecology, Sexual Selection, and the Evolution of Mating Systems",
    year = "1977",
    journal = "Science",
    abstract = "(Uploaded by Plazi for the Bat Literature Project) No abstract provided.",
    url = "https://doi.org/10.1126/science.327542",
    doi = "10.1126/science.327542",
    openalex = "W1977320179",
    references = "doi1010160003347273900043, doi1010160022519364900384, doi1010160022519364900396, doi101086406755, doi101093icb141249, doi101126science1563774477, doi101146annureves05110174001545, doi101163156853974x00345, doi1015159780691185507, doi101537ase188722495, doi1023072874, doi105962bhltitle27468"
}

@incollection{armstrong1978altruism,
    author = "ARMSTRONG, RICHARD D.",
    title = "Altruism, Group Selection, Human Evolution",
    year = "1978",
    booktitle = "Evolutionary Models and Studies in Human Diversity",
    url = "https://doi.org/10.1515/9783110800043.39",
    doi = "10.1515/9783110800043.39",
    openalex = "W2480302470",
    pages = "39-46"
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Altruism is a group phenomenon in which some genes or individuals, which must be presumed to be selfish, benefit others at cost to themselves. The presumption of selfishness and the fact of altruism are reconciled by kin-group selection and by reciprocal altruism. Kin-group selection is clearly visible only in special cases; its role even among social insects may be overestimated; it is probably usually inhibited by competition. However, reciprocal altruism is ubiquitous. All altruism is: (i) potentially reciprocal; (ii) potentially profitable to altruists as well as to recipients; (iii) environmentally determined, usually by position of individuals in group or environmental situations; and (iv) a net-gain lottery. These generalizations are illustrated by four idealized cases; the difficulty of applying them to real cases is illustrated by alarm-calling in groups of birds. Although altruism is a group phenomenon, it evolves by individual selection, by processes equivalent to co-evolutions. Its evolution is: (i) opposed by competition; (ii) costly, complex, and slow, and tending to produce an imprecise flexible altruism rather than a precisely detailed one; and (iii) supplemented by group selection (differential extinction of groups). That altruism in human beings conforms to these generalizations is a good working hypothesis. However, analysis does not "take the altruism out of (human) altruism." Humans do not calculate it, but behave altruistically because they have human altruistic emotions.

@article{doi101073pnas751385,
    author = "Darlington, P. J.",
    title = "Altruism: Its characteristics and evolution",
    year = "1978",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = {Altruism is a group phenomenon in which some genes or individuals, which must be presumed to be selfish, benefit others at cost to themselves. The presumption of selfishness and the fact of altruism are reconciled by kin-group selection and by reciprocal altruism. Kin-group selection is clearly visible only in special cases; its role even among social insects may be overestimated; it is probably usually inhibited by competition. However, reciprocal altruism is ubiquitous. All altruism is: (i) potentially reciprocal; (ii) potentially profitable to altruists as well as to recipients; (iii) environmentally determined, usually by position of individuals in group or environmental situations; and (iv) a net-gain lottery. These generalizations are illustrated by four idealized cases; the difficulty of applying them to real cases is illustrated by alarm-calling in groups of birds. Although altruism is a group phenomenon, it evolves by individual selection, by processes equivalent to co-evolutions. Its evolution is: (i) opposed by competition; (ii) costly, complex, and slow, and tending to produce an imprecise flexible altruism rather than a precisely detailed one; and (iii) supplemented by group selection (differential extinction of groups). That altruism in human beings conforms to these generalizations is a good working hypothesis. However, analysis does not "take the altruism out of (human) altruism." Humans do not calculate it, but behave altruistically because they have human altruistic emotions.},
    url = "https://doi.org/10.1073/pnas.75.1.385",
    doi = "10.1073/pnas.75.1.385",
    openalex = "W1966263378",
    references = "darlington1972nonmathematical, darlington1975group, doi101002j0022033719774110tb01133x, doi101073pnas69113151, doi101073pnas692293, doi101073pnas6951239, doi101073pnas7441647, doi101126science1954280773, doi101126science19743101246, doi1023072063069, openalexw1585956443"
}

Group selection is defined as that process of genetic change which is caused by the differential extinction or proliferation of groups of organisms. A very large proportion of the literature pertaining to group selection consists of theoretical papers; the genetic problems of group selection have been addressed from many different mathematical viewpoints. The general conclusion has been that, although group selection is possible, it cannot override the effects of individual selection within populations except for a highly restricted set of parameter values. Since it is unlikely that conditions in natural populations would fall within the bounds imposed by the models, group selection, by and large, has been considered an insignificant force for evolutionary change. These theoretical conclusions and the assumptions from which they have been derived are reexamined in the light of recent empirical studies of group selection with laboratory populations of the flour beetle, Tribolium (Wade, 1976, 1977). It is shown that the models have a number of assumptions in common which are inherently unfavorable to the operation of group selection. Alternative assumptions derived from the empirical results are suggested and discussed in the hope that they will stimulate further theoretical and empirical study of this controversial subject.

@article{doi101086410450,
    author = "Wade, Michael J.",
    title = "A Critical Review of the Models of Group Selection",
    year = "1978",
    journal = "The Quarterly Review of Biology",
    abstract = "Group selection is defined as that process of genetic change which is caused by the differential extinction or proliferation of groups of organisms. A very large proportion of the literature pertaining to group selection consists of theoretical papers; the genetic problems of group selection have been addressed from many different mathematical viewpoints. The general conclusion has been that, although group selection is possible, it cannot override the effects of individual selection within populations except for a highly restricted set of parameter values. Since it is unlikely that conditions in natural populations would fall within the bounds imposed by the models, group selection, by and large, has been considered an insignificant force for evolutionary change. These theoretical conclusions and the assumptions from which they have been derived are reexamined in the light of recent empirical studies of group selection with laboratory populations of the flour beetle, Tribolium (Wade, 1976, 1977). It is shown that the models have a number of assumptions in common which are inherently unfavorable to the operation of group selection. Alternative assumptions derived from the empirical results are suggested and discussed in the hope that they will stimulate further theoretical and empirical study of this controversial subject.",
    url = "https://doi.org/10.1086/410450",
    doi = "10.1086/410450",
    openalex = "W2030152342"
}

The joint evolution of female mating preferences and secondary sexual characters of males is modeled for polygamous species in which males provide only genetic material to the next generation and females have many potential mates to choose among. Despite stabilizing natural selection on males, various types of mating preferences may create a runaway process in which the outcome of phenotypic evolution depends critically on the genetic variation parameters and initial conditions of a population. Even in the absence of genetic instability, rapid evolution can result from an interaction of natural and sexual selection with random genetic drift along lines of equilibria. The models elucidate genetic mechanisms that can initiate or contribute to rapid speciation by sexual isolation and divergence of secondary sexual characters.

@article{doi101073pnas7863721,
    author = "Lande, Russell",
    title = "Models of speciation by sexual selection on polygenic traits",
    year = "1981",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = "The joint evolution of female mating preferences and secondary sexual characters of males is modeled for polygamous species in which males provide only genetic material to the next generation and females have many potential mates to choose among. Despite stabilizing natural selection on males, various types of mating preferences may create a runaway process in which the outcome of phenotypic evolution depends critically on the genetic variation parameters and initial conditions of a population. Even in the absence of genetic instability, rapid evolution can result from an interaction of natural and sexual selection with random genetic drift along lines of equilibria. The models elucidate genetic mechanisms that can initiate or contribute to rapid speciation by sexual isolation and divergence of secondary sexual characters.",
    url = "https://doi.org/10.1073/pnas.78.6.3721",
    doi = "10.1073/pnas.78.6.3721",
    openalex = "W2002652493",
    references = "doi101017s0016672300016037, doi1023072341823, doi105281zenodo10742832, doi105962bhltitle2112"
}

On theoretical grounds, we argue that parasite-host systems are ideal candidates for interdemic or "group" selection because of the potential for selection for avirulence based upon reduced host (hence group) survival. Such selection appears to have been an important ingredient in the stabilization of the myxomatosis-rabbit system in Australia, although clearly the evolution of resistance in the rabbit population also played a part. We present a simple mathematical model to demonstrate how easily group selection can (in theory) stabilize a parasite-host system. This model is not meant to be a literal translation of the myxomatosis-rabbit interaction, and in fact intentionally disregards host evolution; its purpose is to isolate the role of group selection in the parasite population. Fenner has established that both interdemic selection and host evolution were important in the stabilization of that system, although our model demonstrates that in theory interdemic selection alone could stabilize the interaction. Our central point is that selection for avirulence in the parasite population is important in the evolution of stability in parasite-host systems (Fenner and Myers 1978). The obvious next step, both from the general theoretical viewpoint and specifically with regard to the myxoma-rabbit system, will be to consider models in which both host and parasite are permitted to coevolve. We feel, as do many others, that modeling of coevolution of parasite-host systems has not received the attention deserved in theoretical ecology, and that because of the interdependence of the elements these associations represent much better candidates for such theory than the more popular predator-prey or competition systems. Moreover, as Donald R. Strong, Jr., has pointed out to us, a broad range of organisms are functionally parasitic and the ideas that we have developed herein apply to these as well as to the more restricted class of species which are the conventional organisms treated in parasitology.

@article{doi101086283708,
    author = "Levin, Simon A. and Pimentel, David",
    title = "Selection of Intermediate Rates of Increase in Parasite-Host Systems",
    year = "1981",
    journal = "The American Naturalist",
    abstract = {On theoretical grounds, we argue that parasite-host systems are ideal candidates for interdemic or "group" selection because of the potential for selection for avirulence based upon reduced host (hence group) survival. Such selection appears to have been an important ingredient in the stabilization of the myxomatosis-rabbit system in Australia, although clearly the evolution of resistance in the rabbit population also played a part. We present a simple mathematical model to demonstrate how easily group selection can (in theory) stabilize a parasite-host system. This model is not meant to be a literal translation of the myxomatosis-rabbit interaction, and in fact intentionally disregards host evolution; its purpose is to isolate the role of group selection in the parasite population. Fenner has established that both interdemic selection and host evolution were important in the stabilization of that system, although our model demonstrates that in theory interdemic selection alone could stabilize the interaction. Our central point is that selection for avirulence in the parasite population is important in the evolution of stability in parasite-host systems (Fenner and Myers 1978). The obvious next step, both from the general theoretical viewpoint and specifically with regard to the myxoma-rabbit system, will be to consider models in which both host and parasite are permitted to coevolve. We feel, as do many others, that modeling of coevolution of parasite-host systems has not received the attention deserved in theoretical ecology, and that because of the interdependence of the elements these associations represent much better candidates for such theory than the more popular predator-prey or competition systems. Moreover, as Donald R. Strong, Jr., has pointed out to us, a broad range of organisms are functionally parasitic and the ideas that we have developed herein apply to these as well as to the more restricted class of species which are the conventional organisms treated in parasitology.},
    url = "https://doi.org/10.1086/283708",
    doi = "10.1086/283708",
    openalex = "W1988240917"
}

@article{doi101111j155856461982tb05003x,
    author = "Kirkpatrick, Mark",
    title = "SEXUAL SELECTION AND THE EVOLUTION OF FEMALE CHOICE",
    year = "1982",
    journal = "Evolution",
    url = "https://doi.org/10.1111/j.1558-5646.1982.tb05003.x",
    doi = "10.1111/j.1558-5646.1982.tb05003.x",
    openalex = "W2019598428",
    references = "doi101111j155856461980tb04817x, doi105962bhltitle2112"
}

Natural selection acts on phenotypes, regardless of their genetic basis, and produces immediate phenotypic effects within a generation that can be measured without recourse to principles of heredity or evolution. In contrast, evolutionary response to selection, the genetic change that occurs from one generation to the next, does depend on genetic variation. Animal and plant breeders routinely distinguish phenotypic selection from evolutionary response to selection (Mayo, 1980; Falconer, 1981). Upon making this critical distinction, emphasized by Haldane (1954), precise methods can be formulated for the measurement of phenotypic natural selection. Correlations between characters seriously complicate the measurement of phenotypic selection, because selection on a particular trait produces not only a direct effect on the distribution of that trait in a population, but also produces indirect effects on the distribution of correlated characters. The problem of character correlations has been largely ignored in current methods for measuring natural selection on quantitative traits. Selection has usually been treated as if it acted only on single characters (e.g., Haldane, 1954; Van Valen, 1965a; O'Donald, 1968, 1970; reviewed by Johnson, 1976 Ch. 7). This is obviously a tremendous oversimplification, since natural selection acts on many characters simultaneously and phenotypic correlations between traits are ubiquitous. In an important but neglected paper, Pearson (1903) showed that multivariate statistics could be used to disentangle the direct and indirect effects of selection to determine which traits in a correlated ensemble are the focus of direct selection. Here we extend and generalize Pearson's major results. The purpose of this paper is to derive measures of directional and stabilizing (or disruptive) selection on each of a set of phenotypically correlated characters. The analysis is retrospective, based on observed changes in the multivariate distribution of characters within a generation, not on the evolutionary response to selection. Nevertheless, the measures we propose have a close connection with equations for evolutionary change. Many other commonly used measures of the intensity of selection (such as selective mortality, change in mean fitness, variance in fitness, or estimates of particular forms of fitness functions) have little predictive value in relation to evolutionary change in quantitative traits. To demonstrate the utility of our approach, we analyze selection on four morphological characters in a population of pentatomid bugs during a brief period of high mortality. We also summarize a multivariate selection analysis on nine morphological characters of house sparrows caught in a severe winter storm, using the classic data of Bumpus (1899). Direct observations and measurements of natural selection serve to clarify one of the major factors of evolution. Critiques of the adaptationist program (Lewontin, 1978; Gould and Lewontin, 1979) stress that adaptation and selection are often invoked without strong supporting evidence. We suggest quantitative measurements of selection as the best alternative to the fabrication of adaptive scenarios. Our optimism that measurement can replace rhetorical claims for adaptation and selection is founded in the growing success of field workers in their efforts to measure major components of fitness in natural populations (e.g., Thornhill, 1976; Howard, 1979; Downhower and Brown, 1980; Boag and Grant, 1981; Clutton-Brock et

@article{doi101111j155856461983tb00236x,
    author = "Lande, Russell and Arnold, Stevan J.",
    title = "THE MEASUREMENT OF SELECTION ON CORRELATED CHARACTERS",
    year = "1983",
    journal = "Evolution",
    abstract = "Natural selection acts on phenotypes, regardless of their genetic basis, and produces immediate phenotypic effects within a generation that can be measured without recourse to principles of heredity or evolution. In contrast, evolutionary response to selection, the genetic change that occurs from one generation to the next, does depend on genetic variation. Animal and plant breeders routinely distinguish phenotypic selection from evolutionary response to selection (Mayo, 1980; Falconer, 1981). Upon making this critical distinction, emphasized by Haldane (1954), precise methods can be formulated for the measurement of phenotypic natural selection. Correlations between characters seriously complicate the measurement of phenotypic selection, because selection on a particular trait produces not only a direct effect on the distribution of that trait in a population, but also produces indirect effects on the distribution of correlated characters. The problem of character correlations has been largely ignored in current methods for measuring natural selection on quantitative traits. Selection has usually been treated as if it acted only on single characters (e.g., Haldane, 1954; Van Valen, 1965a; O'Donald, 1968, 1970; reviewed by Johnson, 1976 Ch. 7). This is obviously a tremendous oversimplification, since natural selection acts on many characters simultaneously and phenotypic correlations between traits are ubiquitous. In an important but neglected paper, Pearson (1903) showed that multivariate statistics could be used to disentangle the direct and indirect effects of selection to determine which traits in a correlated ensemble are the focus of direct selection. Here we extend and generalize Pearson's major results. The purpose of this paper is to derive measures of directional and stabilizing (or disruptive) selection on each of a set of phenotypically correlated characters. The analysis is retrospective, based on observed changes in the multivariate distribution of characters within a generation, not on the evolutionary response to selection. Nevertheless, the measures we propose have a close connection with equations for evolutionary change. Many other commonly used measures of the intensity of selection (such as selective mortality, change in mean fitness, variance in fitness, or estimates of particular forms of fitness functions) have little predictive value in relation to evolutionary change in quantitative traits. To demonstrate the utility of our approach, we analyze selection on four morphological characters in a population of pentatomid bugs during a brief period of high mortality. We also summarize a multivariate selection analysis on nine morphological characters of house sparrows caught in a severe winter storm, using the classic data of Bumpus (1899). Direct observations and measurements of natural selection serve to clarify one of the major factors of evolution. Critiques of the adaptationist program (Lewontin, 1978; Gould and Lewontin, 1979) stress that adaptation and selection are often invoked without strong supporting evidence. We suggest quantitative measurements of selection as the best alternative to the fabrication of adaptive scenarios. Our optimism that measurement can replace rhetorical claims for adaptation and selection is founded in the growing success of field workers in their efforts to measure major components of fitness in natural populations (e.g., Thornhill, 1976; Howard, 1979; Downhower and Brown, 1980; Boag and Grant, 1981; Clutton-Brock et",
    url = "https://doi.org/10.1111/j.1558-5646.1983.tb00236.x",
    doi = "10.1111/j.1558-5646.1983.tb00236.x",
    openalex = "W2322715144",
    references = "doi101038227520a0, doi101086404940, doi101093aesa383396, doi101093aibsbulletin2214b, doi101098rspb19790086, doi101111j146918091957tb01874x, doi101111j155856461979tb04694x, doi101890001296582006871445soefdd20co2, doi1023071435536, doi1023071439305, doi1023072344782, doi1023072529912, doi1023074471, doi105962bhltitle27468, doi107312simp93764"
}

The amounts of inbreeding depression upon selfing and of heterosis upon outcrossing determine the strength of selection on the selfing rate in a population when this evolves polygenically by small steps. Genetic models are constructed which allow inbreeding depression to change with the mean selfing rate in a population by incorporating both mutation to recessive and partially dominant lethal and sublethal alleles at many loci and mutation in quantitative characters under stabilizing selection. The models help to explain observations of high inbreeding depression (> 50%) upon selfing in primarily outcrossing populations, as well as considerable heterosis upon outcrossing in primarily selfing populations. Predominant selfing and predominant outcrossing are found to be alternative stable states of the mating system in most plant populations. Which of these stable states a species approaches depends on the history of its population structure and the magnitude of effect of genes influencing the selfing rate.

@article{doi101111j155856461985tb04077x,
    author = "Lande, Russell and Schemske, Douglas W.",
    title = "THE EVOLUTION OF SELF‐FERTILIZATION AND INBREEDING DEPRESSION IN PLANTS. I. GENETIC MODELS",
    year = "1985",
    journal = "Evolution",
    abstract = "The amounts of inbreeding depression upon selfing and of heterosis upon outcrossing determine the strength of selection on the selfing rate in a population when this evolves polygenically by small steps. Genetic models are constructed which allow inbreeding depression to change with the mean selfing rate in a population by incorporating both mutation to recessive and partially dominant lethal and sublethal alleles at many loci and mutation in quantitative characters under stabilizing selection. The models help to explain observations of high inbreeding depression (> 50\%) upon selfing in primarily outcrossing populations, as well as considerable heterosis upon outcrossing in primarily selfing populations. Predominant selfing and predominant outcrossing are found to be alternative stable states of the mating system in most plant populations. Which of these stable states a species approaches depends on the history of its population structure and the magnitude of effect of genes influencing the selfing rate.",
    url = "https://doi.org/10.1111/j.1558-5646.1985.tb04077.x",
    doi = "10.1111/j.1558-5646.1985.tb04077.x",
    openalex = "W2316973238",
    references = "doi101017s0016672300016037, doi101111j155856461983tb00236x, doi1023072407274"
}

@article{nunney1985group,
    author = "Nunney, Len",
    title = "Group Selection, Altruism, and Structured-Deme Models",
    year = "1985",
    journal = "The American Naturalist",
    url = "https://doi.org/10.1086/284410",
    doi = "10.1086/284410",
    number = "2",
    openalex = "W2018945684",
    pages = "212-230",
    volume = "126",
    references = "doi1010160022519364900384, doi101038227520a0, doi101038269578a0, doi101073pnas721143, doi101086410450, doi101093genetics16297, doi101111j1474919x1962tb08690x, doi1015159781400820108, doi105962bhltitle27468, openalexw1493831303"
}

Nature of presents a powerful analysis of the evolutionary concepts of natural selection, fitness, and adaptation. The book clarifies controversial issues concerning altruism, group selection, and the idea that organisms are survival machines built for the good of the genes that inhabit them. As the book unfolds, it provides a straightforward and self-contained introduction to philosophical and biological problems in evolutionary theory.In the first part, Evolutionary Theory as a Theory of Forces, Sober offers an illuminating characterization of the structure of evolutionary theory. Besides laying to rest the spurious charge that theory is vacuous and unscientific, Sober describes the role of chance in evolution and pinpoints the characteristic structure of evolutionary explanations.The book's second part, Group Above and the Gene Below: The Units of Selection Controversy, thoroughly explores the problem of the units of selection, superseding the author's earlier essays, which are widely regarded as the best available treatment of this problem.

@article{doi1023072185056,
    author = "Brandon, Robert N. and Sober, Elliott",
    title = "The Nature of Selection: Evolutionary Theory in Philosophical Focus.",
    year = "1986",
    journal = "The Philosophical Review",
    abstract = "Nature of presents a powerful analysis of the evolutionary concepts of natural selection, fitness, and adaptation. The book clarifies controversial issues concerning altruism, group selection, and the idea that organisms are survival machines built for the good of the genes that inhabit them. As the book unfolds, it provides a straightforward and self-contained introduction to philosophical and biological problems in evolutionary theory.In the first part, Evolutionary Theory as a Theory of Forces, Sober offers an illuminating characterization of the structure of evolutionary theory. Besides laying to rest the spurious charge that theory is vacuous and unscientific, Sober describes the role of chance in evolution and pinpoints the characteristic structure of evolutionary explanations.The book's second part, Group Above and the Gene Below: The Units of Selection Controversy, thoroughly explores the problem of the units of selection, superseding the author's earlier essays, which are widely regarded as the best available treatment of this problem.",
    url = "https://doi.org/10.2307/2185056",
    doi = "10.2307/2185056",
    openalex = "W1498210519"
}

Abstract Various systemic aspects of animal and human minds are explored. Formulation of a replicative evolutionary model of the mind is presented which is based upon the recognition of this entity as a component system. It can be demonstrated that interactions of neurons have replicative organization. It was concluded that intelligent activity of the animal brain manifest itself in producing and maintaining a kind of environmental model. The environmental model is a higher organization above the level of neurons, its basic functional units are called concepts. Each single concept consists of three parts: (1(cue; (2) referential‐structure; and (3) behavioral‐instructions. Interactions among the various concepts of the brain create a concept‐superstructure which behaves as a dynamic replicative component system in controlling animal and human actions. It is assumed that selection operating in the replicative process of concept making which acts as the main factor in creating ontogenetic variability of behavior. The essential process of learning is selection of concepts generated by the brain and in that way the construction of the evolutionary dynamic concept‐superstructures of the brain's environmental model.

@article{doi1010800260402719879972045,
    author = "Csånanyi, V.",
    title = "The replicative evolutionary model of animal and human minds",
    year = "1987",
    journal = "World Futures",
    abstract = "Abstract Various systemic aspects of animal and human minds are explored. Formulation of a replicative evolutionary model of the mind is presented which is based upon the recognition of this entity as a component system. It can be demonstrated that interactions of neurons have replicative organization. It was concluded that intelligent activity of the animal brain manifest itself in producing and maintaining a kind of environmental model. The environmental model is a higher organization above the level of neurons, its basic functional units are called concepts. Each single concept consists of three parts: (1(cue; (2) referential‐structure; and (3) behavioral‐instructions. Interactions among the various concepts of the brain create a concept‐superstructure which behaves as a dynamic replicative component system in controlling animal and human actions. It is assumed that selection operating in the replicative process of concept making which acts as the main factor in creating ontogenetic variability of behavior. The essential process of learning is selection of concepts generated by the brain and in that way the construction of the evolutionary dynamic concept‐superstructures of the brain's environmental model.",
    url = "https://doi.org/10.1080/02604027.1987.9972045",
    doi = "10.1080/02604027.1987.9972045",
    openalex = "W1965938887",
    references = "doi101073pnas6951239"
}

Journal Article Individual selection, kin selection, and the shifting balance in the evolution of warning colours: the evidence from butterflies Get access JAMES MALLET, JAMES MALLET 1Gallon Laboratory, Department of Genetics and Biometry, University College London, Wolfson House, 4 Stephenson Way, London NW1 2HE Search for other works by this author on: Oxford Academic Google Scholar MICHAEL C. SINGER MICHAEL C. SINGER 2Department of Zoology, University of Texas at Austin, Austin, Texas 78712, U.S.A. Search for other works by this author on: Oxford Academic Google Scholar Biological Journal of the Linnean Society, Volume 32, Issue 4, December 1987, Pages 337–350, https://doi.org/10.1111/j.1095-8312.1987.tb00435.x Published: 14 January 2008 Article history Received: 05 February 1986 Accepted: 10 July 1987 Published: 14 January 2008

@article{doi101111j109583121987tb00435x,
    author = "Mallet, James and Singer, Michael C.",
    title = "Individual selection, kin selection, and the shifting balance in the evolution of warning colours: the evidence from butterflies",
    year = "1987",
    journal = "Biological Journal of the Linnean Society",
    abstract = "Journal Article Individual selection, kin selection, and the shifting balance in the evolution of warning colours: the evidence from butterflies Get access JAMES MALLET, JAMES MALLET 1Gallon Laboratory, Department of Genetics and Biometry, University College London, Wolfson House, 4 Stephenson Way, London NW1 2HE Search for other works by this author on: Oxford Academic Google Scholar MICHAEL C. SINGER MICHAEL C. SINGER 2Department of Zoology, University of Texas at Austin, Austin, Texas 78712, U.S.A. Search for other works by this author on: Oxford Academic Google Scholar Biological Journal of the Linnean Society, Volume 32, Issue 4, December 1987, Pages 337–350, https://doi.org/10.1111/j.1095-8312.1987.tb00435.x Published: 14 January 2008 Article history Received: 05 February 1986 Accepted: 10 July 1987 Published: 14 January 2008",
    url = "https://doi.org/10.1111/j.1095-8312.1987.tb00435.x",
    doi = "10.1111/j.1095-8312.1987.tb00435.x",
    openalex = "W2103395895",
    references = "brower1964birds, doi10103712293000, doi101073pnas7863721, doi101126science188418319, doi101126science3576198, doi101537ase188722495, doi1023071437762, doi1023072063068, doi1023072576242, doi1023074510368, doi105962bhltitle27468, doi105962p203298, huheey1961studies, openalexw2624262714"
}

@article{doi101016s0022519388802194,
    author = "Boyd, Robert and Richerson, Peter J.",
    title = "The evolution of reciprocity in sizable groups",
    year = "1988",
    journal = "Journal of Theoretical Biology",
    url = "https://doi.org/10.1016/s0022-5193(88)80219-4",
    doi = "10.1016/s0022-5193(88)80219-4",
    openalex = "W2046307793",
    references = "doi101086410450, nunney1985group"
}

@article{doi101007bf00129882,
    author = "Sober, Elliott",
    title = "The evolution of altruism: Correlation, cost, and benefit",
    year = "1992",
    journal = "Biology \& Philosophy",
    url = "https://doi.org/10.1007/bf00129882",
    doi = "10.1007/bf00129882",
    openalex = "W2015293245",
    references = "doi1010160022519364900384, doi1010160169534789900372, doi101016s0022519389801699, doi101073pnas7941331, doi101086406755, doi101111j143903101979tb00682x, doi101126science7466396, doi1023072174952, doi102307257983, openalexw2624262714"
}

@article{doi101016016230959290032y,
    author = "Boyd, Robert and Richerson, Peter J.",
    title = "Punishment allows the evolution of cooperation (or anything else) in sizable groups",
    year = "1992",
    journal = "Ethology and Sociobiology",
    url = "https://doi.org/10.1016/0162-3095(92)90032-y",
    doi = "10.1016/0162-3095(92)90032-y",
    openalex = "W2147570389",
    references = "doi1023072063069, nunney1985group"
}

@article{goodnight1992contextual,
    author = "Goodnight, Charles J. and Schwartz, James M. and Stevens, Lori",
    title = "Contextual Analysis of Models of Group Selection, Soft Selection, Hard Selection, and the Evolution of Altruism",
    year = "1992",
    journal = "The American Naturalist",
    url = "https://doi.org/10.1086/285438",
    doi = "10.1086/285438",
    number = "5",
    openalex = "W2006819253",
    pages = "743-761",
    volume = "140",
    references = "doi1010160022519364900384, doi101016s0022519389801699, doi1010382011145a0, doi101038227520a0, doi101073pnas721143, doi101111j146918091957tb01874x, doi101111j155856461983tb00236x, doi101111j155856461984tb00344x, doi1015159781400820108, doi1023072529912"
}

Abstract In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as “replicators” which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be “vehicles” of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units. The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to “organs” of group-level “organisms.” Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals.

@article{doi101017s0140525x00036104,
    author = "Wilson, David Sloan and Sober, Elliott",
    title = "Reintroducing group selection to the human behavioral sciences",
    year = "1994",
    journal = "Behavioral and Brain Sciences",
    abstract = "Abstract In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as “replicators” which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be “vehicles” of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units. The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to “organs” of group-level “organisms.” Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals.",
    url = "https://doi.org/10.1017/s0140525x00036104",
    doi = "10.1017/s0140525x00036104",
    openalex = "W2091199874",
    references = "doi101007bf00129882, doi1010160022519364900384, doi101016s0022519389801699, doi101016s0065345408603526, doi101017s0140525x00029939, doi10103712293000, doi101073pnas722646, doi101086406755, doi101086410450, doi101093genetics16297, doi101111j143903101978tb01823x, doi101111j155856461977tb00991x, doi101111j155856461984tb00344x, doi101111j204483091987tb00799x, doi101126science16238591243, doi101126science7466396, doi101146annureves01110170000245, doi1015159781400858712, doi1015159781503621534, doi101537ase188722495, doi1023072026633, doi1023072026953, doi1023072529912, doi105962bhltitle2092, doi105962bhltitle27468, doi107208chicago97802261495160010001, goodnight1992contextual, nunney1985group, openalexw2616504082, openalexw2624262714, openalexw645218623"
}

Abstract Sociopaths are “outstanding” members of society in two senses: politically, they draw our attention because of the inordinate amount of crime they commit, and psychologically, they hold our fascination because most ofus cannot fathom the cold, detached way they repeatedly harm and manipulate others. Proximate explanations from behavior genetics, child development, personality theory, learning theory, and social psychology describe a complex interaction of genetic and physiological risk factors with demographic and micro environmental variables that predispose a portion of the population to chronic antisocial behavior. More recent, evolutionary and game theoretic models have tried to present an ultimate explanation of sociopathy as the expression of a frequency-dependent life strategy which is selected, in dynamic equilibrium, in response to certain varying environmental circumstances. This paper tries to integrate the proximate, developmental models with the ultimate, evolutionary ones, suggesting that two developmentally different etiologies of sociopathy emerge from two different evolutionary mechanisms. Social strategies for minimizing the incidence of sociopathic behavior in modern society should consider the two different etiologies and the factors that contribute to them.

@article{doi101017s0140525x00039595,
    author = "Mealey, Linda",
    title = "The sociobiology of sociopathy: An integrated evolutionary model",
    year = "1995",
    journal = "Behavioral and Brain Sciences",
    abstract = "Abstract Sociopaths are “outstanding” members of society in two senses: politically, they draw our attention because of the inordinate amount of crime they commit, and psychologically, they hold our fascination because most ofus cannot fathom the cold, detached way they repeatedly harm and manipulate others. Proximate explanations from behavior genetics, child development, personality theory, learning theory, and social psychology describe a complex interaction of genetic and physiological risk factors with demographic and micro environmental variables that predispose a portion of the population to chronic antisocial behavior. More recent, evolutionary and game theoretic models have tried to present an ultimate explanation of sociopathy as the expression of a frequency-dependent life strategy which is selected, in dynamic equilibrium, in response to certain varying environmental circumstances. This paper tries to integrate the proximate, developmental models with the ultimate, evolutionary ones, suggesting that two developmentally different etiologies of sociopathy emerge from two different evolutionary mechanisms. Social strategies for minimizing the incidence of sociopathic behavior in modern society should consider the two different etiologies and the factors that contribute to them.",
    url = "https://doi.org/10.1017/s0140525x00039595",
    doi = "10.1017/s0140525x00039595",
    openalex = "W2015467510",
    references = "doi1010160169534789900372, doi101017s0140525x00026522, doi101017s0140525x00029939, doi1023074086942, doi105860choice321538"
}

Once upon a time in evolutionary theory, everything happened for the best. Predators killed only the old or the sick. Pecking orders and other dominance hierarchies minimized wasteful conflict within the group. Male displays ensured that only the best and the fittest had mates. In the culmination of this tradition, Wynne-Edwards (1962, 1986) argued that many species have mechanisms that ensure groups do not over-exploit their resource base. The “central function” of territoriality in birds and other higher animals is “of limiting the numbers of occupants per unit area of habitat” (1986, 6). Species with dominance hierarchies, species with lekking breeding systems, and species with communal breeding regulate their populations. These social mechanisms have population regulation as their “underlying primary function” (1986, 9). Wynne-Edwards argued that these mechanisms evolve through group selection. Populations without such mechanisms are apt to go extinct by eroding their own resource base.

@article{doi101086289977,
    author = "Sterelny, Kim",
    title = "The Return of the Group",
    year = "1996",
    journal = "Philosophy of Science",
    abstract = "Once upon a time in evolutionary theory, everything happened for the best. Predators killed only the old or the sick. Pecking orders and other dominance hierarchies minimized wasteful conflict within the group. Male displays ensured that only the best and the fittest had mates. In the culmination of this tradition, Wynne-Edwards (1962, 1986) argued that many species have mechanisms that ensure groups do not over-exploit their resource base. The “central function” of territoriality in birds and other higher animals is “of limiting the numbers of occupants per unit area of habitat” (1986, 6). Species with dominance hierarchies, species with lekking breeding systems, and species with communal breeding regulate their populations. These social mechanisms have population regulation as their “underlying primary function” (1986, 9). Wynne-Edwards argued that these mechanisms evolve through group selection. Populations without such mechanisms are apt to go extinct by eroding their own resource base.",
    url = "https://doi.org/10.1086/289977",
    doi = "10.1086/289977",
    openalex = "W1972346449",
    references = "doi1010029780470996362ch12, doi101017s0140525x00036104, doi1010382011145a0, doi101093oso97801950693270010001, doi1015159781400820108, doi1023072174952, doi1023072828, doi105860choice304384, openalexw2624262714, openalexw2798374369"
}

Several evolutionary processes influence virulence, the amount of damage a parasite causes to its host. For example, parasites are favored to exploit their hosts prudently to prolong infection and avoid killing the host. Parasites also need to use some host resources to reproduce and transmit infections to new hosts. Thus parasites face a tradeoff between prudent exploitation and rapid reproduction-a life history tradeoff between longevity and fecundity. Other tradeoffs among components of parasite fitness also influence virulence. For example, competition among parasite genotypes favors rapid growth to achieve greater relative success within the host. Rapid growth may, however, lower the total productivity of the local group by overexploiting the host, which is a potentially renewable food supply. This is a problem of kin selection and group selection. I summarize models of parasite virulence with the theoretical tools of life history analysis, kin selection, and epidemiology. I then apply the theory to recent empirical studies and models of virulence. These applications, to nematodes, to the extreme virulence of hospital epidemics, and to bacterial meningitis, show the power of simple life history theory to highlight interesting questions and to provide a rich array of hypotheses. These examples also show the kinds of conceptual mistakes that commonly arise when only a few components of parasite fitness are analysed in isolation. The last part of the article connects standard models of parasite virulence to diverse topics, such as the virulence of bacterial plasmids, the evolution of genomes, and the processes that influenced conflict and cooperation among the earliest replicators near the origin of life.

@article{doi101086419267,
    author = "Frank, Steven A.",
    title = "Models of Parasite Virulence",
    year = "1996",
    journal = "The Quarterly Review of Biology",
    abstract = "Several evolutionary processes influence virulence, the amount of damage a parasite causes to its host. For example, parasites are favored to exploit their hosts prudently to prolong infection and avoid killing the host. Parasites also need to use some host resources to reproduce and transmit infections to new hosts. Thus parasites face a tradeoff between prudent exploitation and rapid reproduction-a life history tradeoff between longevity and fecundity. Other tradeoffs among components of parasite fitness also influence virulence. For example, competition among parasite genotypes favors rapid growth to achieve greater relative success within the host. Rapid growth may, however, lower the total productivity of the local group by overexploiting the host, which is a potentially renewable food supply. This is a problem of kin selection and group selection. I summarize models of parasite virulence with the theoretical tools of life history analysis, kin selection, and epidemiology. I then apply the theory to recent empirical studies and models of virulence. These applications, to nematodes, to the extreme virulence of hospital epidemics, and to bacterial meningitis, show the power of simple life history theory to highlight interesting questions and to provide a rich array of hypotheses. These examples also show the kinds of conceptual mistakes that commonly arise when only a few components of parasite fitness are analysed in isolation. The last part of the article connects standard models of parasite virulence to diverse topics, such as the virulence of bacterial plasmids, the evolution of genomes, and the processes that influenced conflict and cooperation among the earliest replicators near the origin of life.",
    url = "https://doi.org/10.1086/419267",
    doi = "10.1086/419267",
    openalex = "W2157813453",
    references = "doi101007978146847862422, doi101007bf00623322, doi1010160022519364900384, doi101017s0031182000055360, doi101038280445a0, doi10108019390450903037302, doi101086410450, doi101093oso97801985029440010001, doi101098rstb19810005, doi101126science7466396, doi101146annureves01110170000245, doi1023075403, doi105962bhltitle27468, doi1073260003481911721744, openalexw2040525210"
}

Variability selection (abbreviated as VS) is a process considered to link adaptive change to large degrees of environment variability. Its application to hominid evolution is based, in part, on the pronounced rise in environmental remodeling that took place over the past several million years. The VS hypothesis differs from prior views of hominid evolution, which stress the consistent selective effects associated with specific habitats or directional trends (e.g., woodland, savanna expansion, cooling). According to the VS hypothesis, wide fluctuations over time created a growing disparity in adaptive conditions. Inconsistency in selection eventually caused habitat-specific adaptations to be replaced by structures and behaviors responsive to complex environmental change. Key hominid adaptations, in fact, emerged during times of heightened variability. Early bipedality, encephalized brains, and complex human sociality appear to signify a sequence of VS adaptations—i.e., a ratcheting up of versatility and responsiveness to novel environments experienced over the past 6 million years. The adaptive results of VS cannot be extrapolated from selection within a single environmental shift or relatively stable habitat. If some complex traits indeed require disparities in adaptive setting (and relative fitness) in order to evolve, the VS idea counters the prevailing view that adaptive change necessitates long-term, directional consistency in selection. © 1998 Wiley-Liss, Inc.

@article{doi101002sici1520650519987381aidevan330co2a,
    author = "Potts, Richard",
    title = "Variability selection in hominid evolution",
    year = "1998",
    journal = "Evolutionary Anthropology Issues News and Reviews",
    abstract = "Variability selection (abbreviated as VS) is a process considered to link adaptive change to large degrees of environment variability. Its application to hominid evolution is based, in part, on the pronounced rise in environmental remodeling that took place over the past several million years. The VS hypothesis differs from prior views of hominid evolution, which stress the consistent selective effects associated with specific habitats or directional trends (e.g., woodland, savanna expansion, cooling). According to the VS hypothesis, wide fluctuations over time created a growing disparity in adaptive conditions. Inconsistency in selection eventually caused habitat-specific adaptations to be replaced by structures and behaviors responsive to complex environmental change. Key hominid adaptations, in fact, emerged during times of heightened variability. Early bipedality, encephalized brains, and complex human sociality appear to signify a sequence of VS adaptations—i.e., a ratcheting up of versatility and responsiveness to novel environments experienced over the past 6 million years. The adaptive results of VS cannot be extrapolated from selection within a single environmental shift or relatively stable habitat. If some complex traits indeed require disparities in adaptive setting (and relative fitness) in order to evolve, the VS idea counters the prevailing view that adaptive change necessitates long-term, directional consistency in selection. © 1998 Wiley-Liss, Inc.",
    url = "https://doi.org/10.1002/(sici)1520-6505(1998)7:3<81::aid-evan3>3.0.co;2-a",
    doi = "10.1002/(sici)1520-6505(1998)7:3<81::aid-evan3>3.0.co;2-a",
    openalex = "W2025328236",
    references = "doi101007bf02547562, doi101038371306a0, doi101038scientificamerican096062"
}

Part 1 Introduction: game theory equilibrium evolutionary games evolution evolution and equilibrium the path ahead. Part 2 The evolution of models: evolutionarily stable strategies alternative notions of evolutionary stability synamic systems. Part 3 A model of evolution: the aspiration and imitation model time sample paths stationary distributions limits. Appendix: proofs. Part 4 The dynamics of sample paths: dynamics equilibrium strictly dominated strategies weakly dominated strategies. Appendix: proofs. Part 5 The ultimatum game: the ultimatum game an ultimatum minigame numerical calculations relevance to experimental data? leaving money on the table. Appendix: proofs. Part 6 Drift: introduction the model when can drift be ignored? when drift matters examples discussion. Part 7 Noise: the model limit distributions alternative best replies trembles. Appendix: proofs. Part 8 Backward and forward induction: introduction the model recurrent outcomes backward induction forward induction markets. appendix: proofs. Part 9 Strict Nash equilibria: a muddling model dynamics equilibrium selection risk dominance discussion.

@article{doi1023071061220,
    author = "Mezzetti, Claudio and Samuelson, Larry",
    title = "Evolutionary Games and Equilibrium Selection",
    year = "1998",
    journal = "Southern Economic Journal",
    abstract = "Part 1 Introduction: game theory equilibrium evolutionary games evolution evolution and equilibrium the path ahead. Part 2 The evolution of models: evolutionarily stable strategies alternative notions of evolutionary stability synamic systems. Part 3 A model of evolution: the aspiration and imitation model time sample paths stationary distributions limits. Appendix: proofs. Part 4 The dynamics of sample paths: dynamics equilibrium strictly dominated strategies weakly dominated strategies. Appendix: proofs. Part 5 The ultimatum game: the ultimatum game an ultimatum minigame numerical calculations relevance to experimental data? leaving money on the table. Appendix: proofs. Part 6 Drift: introduction the model when can drift be ignored? when drift matters examples discussion. Part 7 Noise: the model limit distributions alternative best replies trembles. Appendix: proofs. Part 8 Backward and forward induction: introduction the model recurrent outcomes backward induction forward induction markets. appendix: proofs. Part 9 Strict Nash equilibria: a muddling model dynamics equilibrium selection risk dominance discussion.",
    url = "https://doi.org/10.2307/1061220",
    doi = "10.2307/1061220",
    openalex = "W1546006912"
}

Social interactions often affect the fitness of interactants. Because of this, social selection has been described as a process distinct from other forms of natural selection. Social selection has been predicted to result in different evolutionary dynamics for interacting phenotypes, including rapid or extreme evolution and evolution of altruism. Despite the critical role that social selection plays in theories of social evolution, few studies have measured the force of social selection or the conditions under which this force changes. Here we present a model of social selection acting on interacting phenotypes that can be evaluated independently from the genetics of interacting phenotypes. Our model of social selection is analogous to covariance models of other forms of selection. We observe that an opportunity for social selection exists whenever individual fitness varies as a result of interactions with conspecifics. Social selection occurs, therefore, when variation in fitness due to interactions covaries with traits, resulting in a net force of selection acting on the interacting phenotypes. Thus, there must be a covariance between the phenotypes of the interactants for social selection to exist. This interacting phenotype covariance is important because it measures the degree to which a particular trait covaries with the selective environment provided by conspecifics. A variety of factors, including nonrandom interactions, behavioral modification during interactions, relatedness, and indirect genetic effects may contribute to the covariance of interacting phenotypes, which promotes social selection. The independent force of social selection (measured as a social selection gradient) can be partitioned empirically from the force of natural selection (measured by the natural selection gradient) using partial regression. This measure can be combined with genetic models of interacting phenotypes to provide insights into social evolution.

@article{doi101086303168,
    author = "Wolf, Jason B. and Brodie, Edmund D. and Moore, Allen J.",
    title = "Interacting Phenotypes and the Evolutionary Process. II. Selection Resulting from Social Interactions",
    year = "1999",
    journal = "The American Naturalist",
    abstract = "Social interactions often affect the fitness of interactants. Because of this, social selection has been described as a process distinct from other forms of natural selection. Social selection has been predicted to result in different evolutionary dynamics for interacting phenotypes, including rapid or extreme evolution and evolution of altruism. Despite the critical role that social selection plays in theories of social evolution, few studies have measured the force of social selection or the conditions under which this force changes. Here we present a model of social selection acting on interacting phenotypes that can be evaluated independently from the genetics of interacting phenotypes. Our model of social selection is analogous to covariance models of other forms of selection. We observe that an opportunity for social selection exists whenever individual fitness varies as a result of interactions with conspecifics. Social selection occurs, therefore, when variation in fitness due to interactions covaries with traits, resulting in a net force of selection acting on the interacting phenotypes. Thus, there must be a covariance between the phenotypes of the interactants for social selection to exist. This interacting phenotype covariance is important because it measures the degree to which a particular trait covaries with the selective environment provided by conspecifics. A variety of factors, including nonrandom interactions, behavioral modification during interactions, relatedness, and indirect genetic effects may contribute to the covariance of interacting phenotypes, which promotes social selection. The independent force of social selection (measured as a social selection gradient) can be partitioned empirically from the force of natural selection (measured by the natural selection gradient) using partial regression. This measure can be combined with genetic models of interacting phenotypes to provide insights into social evolution.",
    url = "https://doi.org/10.1086/303168",
    doi = "10.1086/303168",
    openalex = "W2071931278",
    references = "doi101007978146847862422, doi1010160022519364900384, doi101016s001600323892229x, doi101017cbo9780511806292, doi101046j14390388200200356x, doi101111j155856461983tb00236x, doi101126science7466396, doi1023072529912, doi104159harvard9780674865327, goodnight1992contextual, openalexw2624262714"
}

@article{silva1999deterministic,
    author = "Silva, A.T.C. and Fontanari, J.F.",
    title = "Deterministic group selection model for the evolution of altruism",
    year = "1999",
    journal = "The European Physical Journal B",
    url = "https://doi.org/10.1007/s100510050626",
    doi = "10.1007/s100510050626",
    number = "3",
    openalex = "W2597472162",
    pages = "385-392",
    volume = "7"
}

@article{silva1999stochastic,
    author = "Silva, Ana T.C. and Fontanari, J.F.",
    title = "Stochastic group selection model for the evolution of altruism",
    year = "1999",
    journal = "Physica A: Statistical Mechanics and its Applications",
    url = "https://doi.org/10.1016/s0378-4371(99)00065-5",
    doi = "10.1016/s0378-4371(99)00065-5",
    number = "1-2",
    openalex = "W2098685263",
    pages = "257-268",
    volume = "268",
    references = "doi101007bf00623322, doi101086283708, doi101086419267, doi101093genetics16297, doi101093genetics282114, doi101111j1474919x1962tb08690x, doi101111j155856461993tb01266x, doi101111j174966321975tb35099x, doi1023072174952, doi1023072410218, silva1999deterministic"
}

The group selection controversy is about whether natural selection ever operates at the level of groups, rather than at the level of individual organisms. Traditionally, group selection has been invoked to explain the existence of altruistic behaviour in nature. However, most contemporary evolutionary biologists are highly sceptical of the hypothesis of group selection, which they regard as biologically implausible and not needed to explain the evolution of altruism anyway. But in their recent book, Elliot Sober and David Sloan Wilson [1998] argue that the widespread opposition to group selection is founded on conceptual confusion. The theories that have been propounded as alternatives to group selection are actually group selection in disguise, they maintain. I examine their arguments for this claim, and John Maynard Smith's arguments against it. I argue that Sober and Wilson arrive at a correct position by faulty reasoning. In the final section, I examine the issue of how to apply the principle of natural selection at different levels of the biological hierarchy, which underlies the dispute between Sober and Wilson and Maynard Smith.

@article{doi101093bjps52125,
    author = "Okasha, Samir",
    title = "Why Won't the Group Selection Controversy Go Away?",
    year = "2001",
    journal = "The British Journal for the Philosophy of Science",
    abstract = "The group selection controversy is about whether natural selection ever operates at the level of groups, rather than at the level of individual organisms. Traditionally, group selection has been invoked to explain the existence of altruistic behaviour in nature. However, most contemporary evolutionary biologists are highly sceptical of the hypothesis of group selection, which they regard as biologically implausible and not needed to explain the evolution of altruism anyway. But in their recent book, Elliot Sober and David Sloan Wilson [1998] argue that the widespread opposition to group selection is founded on conceptual confusion. The theories that have been propounded as alternatives to group selection are actually group selection in disguise, they maintain. I examine their arguments for this claim, and John Maynard Smith's arguments against it. I argue that Sober and Wilson arrive at a correct position by faulty reasoning. In the final section, I examine the issue of how to apply the principle of natural selection at different levels of the biological hierarchy, which underlies the dispute between Sober and Wilson and Maynard Smith.",
    url = "https://doi.org/10.1093/bjps/52.1.25",
    doi = "10.1093/bjps/52.1.25",
    openalex = "W1984721570",
    references = "doi101086289977"
}

@article{barrett2002group,
    author = "BARRETT, MATTHEW and GODFREY‐SMITH, PETER",
    title = "Group Selection, Pluralism, and the Evolution of Altruism",
    year = "2002",
    journal = "Philosophy and Phenomenological Research",
    url = "https://doi.org/10.1111/j.1933-1592.2002.tb00233.x",
    doi = "10.1111/j.1933-1592.2002.tb00233.x",
    number = "3",
    openalex = "W2001924915",
    pages = "685-691",
    volume = "65",
    references = "doi101007bf00129882, doi101016s0065345408603526, doi10103831383, doi101046j14390310199900372x, doi101086289977, doi1023072026633, doi1023072026953"
}

@article{doi101006jtbi20033144,
    author = "Ono, Seiji and Misawa, Kazuharu and Tsuji, Kazuki",
    title = "Effect of Group Selection on the Evolution of Altruistic Behavior",
    year = "2002",
    journal = "Journal of Theoretical Biology",
    url = "https://doi.org/10.1006/jtbi.2003.3144",
    doi = "10.1006/jtbi.2003.3144",
    openalex = "W2031506983",
    references = "silva1999stochastic"
}

@article{cooper2004group,
    author = "Cooper, B.",
    title = "Group selection and the evolution of altruism",
    year = "2004",
    journal = "Oxford Economic Papers",
    url = "https://doi.org/10.1093/oep/gpf043",
    doi = "10.1093/oep/gpf043",
    number = "2",
    openalex = "W2098687748",
    pages = "307-330",
    volume = "56",
    references = "doi1010160022519364900384, doi101017cbo9780511493607, doi1010382011145a0, doi101046j14390310199900372x, doi101073pnas721143, doi101111j251761611951tb00088x, doi101537ase188722495, doi105962bhltitle2112, openalexw1494315736, openalexw2624262714"
}

Group selection is one acknowledged mechanism for the evolution of altruism. It is well known that for altruism to spread by natural selection, interactions must be correlated; that is, altruists must tend to associate with one another. But does group selection itself require correlated interactions? Two possible arguments for answering this question affirmatively are explored. The first is a bad argument, for it rests on a product/process confusion. The second is a more subtle argument, whose validity (or otherwise) turns on issues concerning the meaning of multi-level selection and how it should be modelled. A cautious defence of the second argument is offered. 1. Introduction 2. Multi-level selection and the evolution of altruism 3. Price's equation and multi-level selection 4. Contextual analysis and multi-level selection 5. The neighbour approach 6. Recapitulation and conclusion

@article{doi101093bjpsaxi143,
    author = "Okasha, Samir",
    title = "Altruism, Group Selection and Correlated Interaction",
    year = "2005",
    journal = "The British Journal for the Philosophy of Science",
    abstract = "Group selection is one acknowledged mechanism for the evolution of altruism. It is well known that for altruism to spread by natural selection, interactions must be correlated; that is, altruists must tend to associate with one another. But does group selection itself require correlated interactions? Two possible arguments for answering this question affirmatively are explored. The first is a bad argument, for it rests on a product/process confusion. The second is a more subtle argument, whose validity (or otherwise) turns on issues concerning the meaning of multi-level selection and how it should be modelled. A cautious defence of the second argument is offered. 1. Introduction 2. Multi-level selection and the evolution of altruism 3. Price's equation and multi-level selection 4. Contextual analysis and multi-level selection 5. The neighbour approach 6. Recapitulation and conclusion",
    url = "https://doi.org/10.1093/bjps/axi143",
    doi = "10.1093/bjps/axi143",
    openalex = "W2134905101",
    references = "doi101007bf00129882"
}

The Trust Game is regarded as an important model of corporate culture in that social learning to promote trust creates organizational efficiencies consistent with the existence of firms. We examine the Trust Game from an evolutionary perspective in which player types are defined in terms of their commitment to trustworthy behavior. In so doing, a condition for multilevel selection (group effects) is identified. When group effects exist—implying that the matching process is not independent of players' commitment—a corporate culture emerges that promotes trust. This results in an evolutionary theory of business ethics.

@article{doi101002j232580122006tb00755x,
    author = "Arce, Daniel G.",
    title = "Taking Corporate Culture Seriously: Group Effects in the Trust Game",
    year = "2006",
    journal = "Southern Economic Journal",
    abstract = "The Trust Game is regarded as an important model of corporate culture in that social learning to promote trust creates organizational efficiencies consistent with the existence of firms. We examine the Trust Game from an evolutionary perspective in which player types are defined in terms of their commitment to trustworthy behavior. In so doing, a condition for multilevel selection (group effects) is identified. When group effects exist—implying that the matching process is not independent of players' commitment—a corporate culture emerges that promotes trust. This results in an evolutionary theory of business ethics.",
    url = "https://doi.org/10.1002/j.2325-8012.2006.tb00755.x",
    doi = "10.1002/j.2325-8012.2006.tb00755.x",
    openalex = "W2026185580",
    references = "cooper2004group, doi1010160160932782900369, doi1010160167268185900174, doi101016s0167268103000945, doi101017cbo9780511571657006, doi101017s0140525x00036104, doi101038227520a0, doi101093oso97801987743580030005, doi101111j251761611951tb00088x, doi1023071061220, openalexw1512416168"
}

We propose a minimalist stochastic model of multilevel (or group) selection. A population is subdivided into groups. Individuals interact with other members of the group in an evolutionary game that determines their fitness. Individuals reproduce, and offspring are added to the same group. If a group reaches a certain size, it can split into two. Faster reproducing individuals lead to larger groups that split more often. In our model, higher-level selection emerges as a byproduct of individual reproduction and population structure. We derive a fundamental condition for the evolution of cooperation by group selection: if b/c > 1 + n/m, then group selection favors cooperation. The parameters b and c denote the benefit and cost of the altruistic act, whereas n and m denote the maximum group size and the number of groups. The model can be extended to more than two levels of selection and to include migration.

@article{doi101073pnas0602530103,
    author = "Traulsen, Arne and Nowak, Martin A.",
    title = "Evolution of cooperation by multilevel selection",
    year = "2006",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = "We propose a minimalist stochastic model of multilevel (or group) selection. A population is subdivided into groups. Individuals interact with other members of the group in an evolutionary game that determines their fitness. Individuals reproduce, and offspring are added to the same group. If a group reaches a certain size, it can split into two. Faster reproducing individuals lead to larger groups that split more often. In our model, higher-level selection emerges as a byproduct of individual reproduction and population structure. We derive a fundamental condition for the evolution of cooperation by group selection: if b/c > 1 + n/m, then group selection favors cooperation. The parameters b and c denote the benefit and cost of the altruistic act, whereas n and m denote the maximum group size and the number of groups. The model can be extended to more than two levels of selection and to include migration.",
    url = "https://doi.org/10.1073/pnas.0602530103",
    doi = "10.1073/pnas.0602530103",
    openalex = "W2009481751",
    references = "doi101023a1020504900646, doi101038nature01906, doi101111j155856461977tb00991x, nunney1985group, openalexw645218623"
}

Does natural selection act primarily on individual organisms, on groups, on genes, or on whole species? This book provides a comprehensive analysis of the debate in evolutionary biology over the levels of selection, focusing on conceptual philosophical and foundational questions.

@book{doi101093acprofoso97801992679720010001,
    author = "Okasha, Samir",
    title = "Evolution and the Levels of Selection",
    year = "2006",
    booktitle = "Oxford University Press eBooks",
    abstract = "Does natural selection act primarily on individual organisms, on groups, on genes, or on whole species? This book provides a comprehensive analysis of the debate in evolutionary biology over the levels of selection, focusing on conceptual philosophical and foundational questions.",
    url = "https://doi.org/10.1093/acprof:oso/9780199267972.001.0001",
    doi = "10.1093/acprof:oso/9780199267972.001.0001",
    openalex = "W2098069022",
    references = "doi1010020470099704, doi101525bio20106059, doi105860choice396411, doi105962bhltitle3163"
}

One of the enduring puzzles in biology and the social sciences is the origin and persistence of intraspecific cooperation and altruism in humans and other species. Hundreds of theoretical models have been proposed and there is much confusion about the relationship between these models. To clarify the situation, we developed a synthetic conceptual framework that delineates the conditions necessary for the evolution of altruism and cooperation. We show that at least one of the four following conditions needs to be fulfilled: direct benefits to the focal individual performing a cooperative act; direct or indirect information allowing a better than random guess about whether a given individual will behave cooperatively in repeated reciprocal interactions; preferential interactions between related individuals; and genetic correlation between genes coding for altruism and phenotypic traits that can be identified. When one or more of these conditions are met, altruism or cooperation can evolve if the cost-to-benefit ratio of altruistic and cooperative acts is greater than a threshold value. The cost-to-benefit ratio can be altered by coercion, punishment and policing which therefore act as mechanisms facilitating the evolution of altruism and cooperation. All the models proposed so far are explicitly or implicitly built on these general principles, allowing us to classify them into four general categories.

@article{doi101111j14209101200601119x,
    author = "Lehmann, Laurent and Keller, Laurent",
    title = "The evolution of cooperation and altruism – a general framework and a classification of models",
    year = "2006",
    journal = "Journal of Evolutionary Biology",
    abstract = "One of the enduring puzzles in biology and the social sciences is the origin and persistence of intraspecific cooperation and altruism in humans and other species. Hundreds of theoretical models have been proposed and there is much confusion about the relationship between these models. To clarify the situation, we developed a synthetic conceptual framework that delineates the conditions necessary for the evolution of altruism and cooperation. We show that at least one of the four following conditions needs to be fulfilled: direct benefits to the focal individual performing a cooperative act; direct or indirect information allowing a better than random guess about whether a given individual will behave cooperatively in repeated reciprocal interactions; preferential interactions between related individuals; and genetic correlation between genes coding for altruism and phenotypic traits that can be identified. When one or more of these conditions are met, altruism or cooperation can evolve if the cost-to-benefit ratio of altruistic and cooperative acts is greater than a threshold value. The cost-to-benefit ratio can be altered by coercion, punishment and policing which therefore act as mechanisms facilitating the evolution of altruism and cooperation. All the models proposed so far are explicitly or implicitly built on these general principles, allowing us to classify them into four general categories.",
    url = "https://doi.org/10.1111/j.1420-9101.2006.01119.x",
    doi = "10.1111/j.1420-9101.2006.01119.x",
    openalex = "W2112573298",
    references = "doi101016s0065345408603526, doi101086383541, nunney1985group"
}

From an evolutionary perspective, social behaviours are those which have fitness consequences for both the individual that performs the behaviour, and another individual. Over the last 43 years, a huge theoretical and empirical literature has developed on this topic. However, progress is often hindered by poor communication between scientists, with different people using the same term to mean different things, or different terms to mean the same thing. This can obscure what is biologically important, and what is not. The potential for such semantic confusion is greatest with interdisciplinary research. Our aim here is to address issues of semantic confusion that have arisen with research on the problem of cooperation. In particular, we: (i) discuss confusion over the terms kin selection, mutualism, mutual benefit, cooperation, altruism, reciprocal altruism, weak altruism, altruistic punishment, strong reciprocity, group selection and direct fitness; (ii) emphasize the need to distinguish between proximate (mechanism) and ultimate (survival value) explanations of behaviours. We draw examples from all areas, but especially recent work on humans and microbes.

@article{doi101111j14209101200601258x,
    author = "West, Stuart A. and Griffin, Ashleigh S. and Gardner, Andy",
    title = "Social semantics: altruism, cooperation, mutualism, strong reciprocity and group selection",
    year = "2006",
    journal = "Journal of Evolutionary Biology",
    abstract = "From an evolutionary perspective, social behaviours are those which have fitness consequences for both the individual that performs the behaviour, and another individual. Over the last 43 years, a huge theoretical and empirical literature has developed on this topic. However, progress is often hindered by poor communication between scientists, with different people using the same term to mean different things, or different terms to mean the same thing. This can obscure what is biologically important, and what is not. The potential for such semantic confusion is greatest with interdisciplinary research. Our aim here is to address issues of semantic confusion that have arisen with research on the problem of cooperation. In particular, we: (i) discuss confusion over the terms kin selection, mutualism, mutual benefit, cooperation, altruism, reciprocal altruism, weak altruism, altruistic punishment, strong reciprocity, group selection and direct fitness; (ii) emphasize the need to distinguish between proximate (mechanism) and ultimate (survival value) explanations of behaviours. We draw examples from all areas, but especially recent work on humans and microbes.",
    url = "https://doi.org/10.1111/j.1420-9101.2006.01258.x",
    doi = "10.1111/j.1420-9101.2006.01258.x",
    openalex = "W2124337033",
    references = "doi101006jtbi20002111, doi101016s0065345408603526, doi101016s1090513804000054, doi1010382011145a0, doi101046j14390310199900372x, doi101073pnas721143, doi101086383541, doi101093oso97801985029440010001, doi101111j143903101963tb01161x, doi101111j155856461995tb04464x, doi101126science1563774477, doi1015159780691206820, doi1023072828, openalexw2616504082"
}

Interaction among individuals is universal, both in animals and in plants, and substantially affects evolution of natural populations and responses to artificial selection in agriculture. Although quantitative genetics has successfully been applied to many traits, it does not provide a general theory accounting for interaction among individuals and selection acting on multiple levels. Consequently, current quantitative genetic theory fails to explain why some traits do not respond to selection among individuals, but respond greatly to selection among groups. Understanding the full impacts of heritable interactions on the outcomes of selection requires a quantitative genetic framework including all levels of selection and relatedness. Here we present such a framework and provide expressions for the response to selection. Results show that interaction among individuals may create substantial heritable variation, which is hidden to classical analyses. Selection acting on higher levels of organization captures this hidden variation and therefore always yields positive response, whereas individual selection may yield response in the opposite direction. Our work provides testable predictions of response to multilevel selection and reduces to classical theory in the absence of interaction. Statistical methodology provided elsewhere enables empirical application of our work to both natural and domestic populations.

@article{doi101534genetics106062711,
    author = "Bijma, Piter and Muir, William M. and van Arendonk, J.A.M.",
    title = "Multilevel Selection 1: Quantitative Genetics of Inheritance and Response to Selection",
    year = "2006",
    journal = "Genetics",
    abstract = "Interaction among individuals is universal, both in animals and in plants, and substantially affects evolution of natural populations and responses to artificial selection in agriculture. Although quantitative genetics has successfully been applied to many traits, it does not provide a general theory accounting for interaction among individuals and selection acting on multiple levels. Consequently, current quantitative genetic theory fails to explain why some traits do not respond to selection among individuals, but respond greatly to selection among groups. Understanding the full impacts of heritable interactions on the outcomes of selection requires a quantitative genetic framework including all levels of selection and relatedness. Here we present such a framework and provide expressions for the response to selection. Results show that interaction among individuals may create substantial heritable variation, which is hidden to classical analyses. Selection acting on higher levels of organization captures this hidden variation and therefore always yields positive response, whereas individual selection may yield response in the opposite direction. Our work provides testable predictions of response to multilevel selection and reduces to classical theory in the absence of interaction. Statistical methodology provided elsewhere enables empirical application of our work to both natural and domestic populations.",
    url = "https://doi.org/10.1534/genetics.106.062711",
    doi = "10.1534/genetics.106.062711",
    openalex = "W2132362070",
    references = "cooper2004group, doi1010160022519364900384, doi101016s0065345408603526, doi101038238413a0, doi10103835012234, doi101046j14390388200200356x, doi101111j146918091949tb02451x, doi101111j155856461965tb01731x, doi101111j155856461977tb00991x, doi101111j155856461983tb00236x, doi1023072408842, doi1023072529912, doi105962bhltitle27468, goodnight1992contextual"
}

Evolutionary theory postulates that altruistic behavior evolved for the return-benefits it bears the performer. For return-benefits to play a motivational role, however, they need to be experienced by the organism. Motivational analyses should restrict themselves, therefore, to the altruistic impulse and its knowable consequences. Empathy is an ideal candidate mechanism to underlie so-called directed altruism, i.e., altruism in response to anothers's pain, need, or distress. Evidence is accumulating that this mechanism is phylogenetically ancient, probably as old as mammals and birds. Perception of the emotional state of another automatically activates shared representations causing a matching emotional state in the observer. With increasing cognition, state-matching evolved into more complex forms, including concern for the other and perspective-taking. Empathy-induced altruism derives its strength from the emotional stake it offers the self in the other's welfare. The dynamics of the empathy mechanism agree with predictions from kin selection and reciprocal altruism theory.

@article{doi101146annurevpsych59103006093625,
    author = "de Waal, Frans Β. Μ.",
    title = "Putting the Altruism Back into Altruism: The Evolution of Empathy",
    year = "2007",
    journal = "Annual Review of Psychology",
    abstract = "Evolutionary theory postulates that altruistic behavior evolved for the return-benefits it bears the performer. For return-benefits to play a motivational role, however, they need to be experienced by the organism. Motivational analyses should restrict themselves, therefore, to the altruistic impulse and its knowable consequences. Empathy is an ideal candidate mechanism to underlie so-called directed altruism, i.e., altruism in response to anothers's pain, need, or distress. Evidence is accumulating that this mechanism is phylogenetically ancient, probably as old as mammals and birds. Perception of the emotional state of another automatically activates shared representations causing a matching emotional state in the observer. With increasing cognition, state-matching evolved into more complex forms, including concern for the other and perspective-taking. Empathy-induced altruism derives its strength from the emotional stake it offers the self in the other's welfare. The dynamics of the empathy mechanism agree with predictions from kin selection and reciprocal altruism theory.",
    url = "https://doi.org/10.1146/annurev.psych.59.103006.093625",
    doi = "10.1146/annurev.psych.59.103006.093625",
    openalex = "W2134084792",
    references = "doi101002ajp1350020302, doi101002j153892351995tb03988x, doi1010160022519364900384, doi101017s0140525x02000018, doi10103700121649281126, doi101038nature04271, doi101046j14390310199900372x, doi101073pnas0608062103, doi101073pnas101086398, doi101086406755, doi101093oso97801950967360010001, doi101111j143903101963tb01161x, doi101126science1093535, doi105860choice294797, doi105860choice351500, openalexw1581387623, openalexw1849553904, openalexw2001431842, openalexw2126474339, openalexw2624262714"
}

Group-structured populations, of the kind prominent in discussions of multilevel selection, are contrasted with ‘neighbor-structured’ populations. I argue that it is a necessary condition on multilevel description of a selection process that there should be a nonarbitrary division of the population into equivalence classes (or an approximation to this situation). The discussion is focused via comparisons between two famous problem cases involving group structure (altruism and heterozygote advantage) and two neighbor-structured cases that resemble them. Conclusions are also drawn about the role of correlated interaction in the evolution of altruism. 1. Introduction2. Two Kinds of Population Structure3. Objections and Replies4. Particles on a Line5. ConclusionAppendix: Neighborhoods and Selection

@article{doi101093bjpsaxm044,
    author = "Godfrey‐Smith, Peter",
    title = "Varieties of Population Structure and the Levels of Selection",
    year = "2008",
    journal = "The British Journal for the Philosophy of Science",
    abstract = "Group-structured populations, of the kind prominent in discussions of multilevel selection, are contrasted with ‘neighbor-structured’ populations. I argue that it is a necessary condition on multilevel description of a selection process that there should be a nonarbitrary division of the population into equivalence classes (or an approximation to this situation). The discussion is focused via comparisons between two famous problem cases involving group structure (altruism and heterozygote advantage) and two neighbor-structured cases that resemble them. Conclusions are also drawn about the role of correlated interaction in the evolution of altruism. 1. Introduction2. Two Kinds of Population Structure3. Objections and Replies4. Particles on a Line5. ConclusionAppendix: Neighborhoods and Selection",
    url = "https://doi.org/10.1093/bjps/axm044",
    doi = "10.1093/bjps/axm044",
    openalex = "W1995109037",
    references = "doi101007bf00129882, doi101086289977"
}

We present a simple framework that highlights the most fundamental requirement for the evolution of altruism: assortment between individuals carrying the cooperative genotype and the helping behaviours of others with which these individuals interact. We partition the fitness effects on individuals into those due to self and those due to the 'interaction environment', and show that it is the latter that is most fundamental to understanding the evolution of altruism. We illustrate that while kinship or genetic similarity among those interacting may generate a favourable structure of interaction environments, it is not a fundamental requirement for the evolution of altruism, and even suicidal aid can theoretically evolve without help ever being exchanged among genetically similar individuals. Using our simple framework, we also clarify a common confusion made in the literature between alternative fitness accounting methods (which may equally apply to the same biological circumstances) and unique causal mechanisms for creating the assortment necessary for altruism to be favoured by natural selection.

@article{doi101098rspb20080829,
    author = "Fletcher, Jeffrey and Doebeli, Michael",
    title = "A simple and general explanation for the evolution of altruism",
    year = "2008",
    journal = "Proceedings of the Royal Society B Biological Sciences",
    abstract = "We present a simple framework that highlights the most fundamental requirement for the evolution of altruism: assortment between individuals carrying the cooperative genotype and the helping behaviours of others with which these individuals interact. We partition the fitness effects on individuals into those due to self and those due to the 'interaction environment', and show that it is the latter that is most fundamental to understanding the evolution of altruism. We illustrate that while kinship or genetic similarity among those interacting may generate a favourable structure of interaction environments, it is not a fundamental requirement for the evolution of altruism, and even suicidal aid can theoretically evolve without help ever being exchanged among genetically similar individuals. Using our simple framework, we also clarify a common confusion made in the literature between alternative fitness accounting methods (which may equally apply to the same biological circumstances) and unique causal mechanisms for creating the assortment necessary for altruism to be favoured by natural selection.",
    url = "https://doi.org/10.1098/rspb.2008.0829",
    doi = "10.1098/rspb.2008.0829",
    openalex = "W2019988542",
    references = "doi10103831383, nunney1985group"
}

Kin and levels-of-selection models are common approaches for modelling social evolution. Indirect genetic effect (IGE) models represent a different approach, specifying social effects on trait values rather than fitness. We investigate the joint effect of relatedness, multilevel selection and IGEs on response to selection. We present a measure for the degree of multilevel selection, which is the natural partner of relatedness in expressions for response. Response depends on both relatedness and the degree of multilevel selection, rather than only one or the other factor. Moreover, response is symmetric in relatedness and the degree of multilevel selection, indicating that both factors have exactly the same effect. Without IGEs, the key parameter is the product of relatedness and the degree of multilevel selection. With IGEs, however, multilevel selection without relatedness can explain evolution of social traits. Thus, next to relatedness and multilevel selection, IGEs are a key element in the genetical theory of social evolution.

@article{doi101111j14209101200801550x,
    author = "Bijma, Piter and Wade, Michael J.",
    title = "The joint effects of kin, multilevel selection and indirect genetic effects on response to genetic selection",
    year = "2008",
    journal = "Journal of Evolutionary Biology",
    abstract = "Kin and levels-of-selection models are common approaches for modelling social evolution. Indirect genetic effect (IGE) models represent a different approach, specifying social effects on trait values rather than fitness. We investigate the joint effect of relatedness, multilevel selection and IGEs on response to selection. We present a measure for the degree of multilevel selection, which is the natural partner of relatedness in expressions for response. Response depends on both relatedness and the degree of multilevel selection, rather than only one or the other factor. Moreover, response is symmetric in relatedness and the degree of multilevel selection, indicating that both factors have exactly the same effect. Without IGEs, the key parameter is the product of relatedness and the degree of multilevel selection. With IGEs, however, multilevel selection without relatedness can explain evolution of social traits. Thus, next to relatedness and multilevel selection, IGEs are a key element in the genetical theory of social evolution.",
    url = "https://doi.org/10.1111/j.1420-9101.2008.01550.x",
    doi = "10.1111/j.1420-9101.2008.01550.x",
    openalex = "W1985894030",
    references = "doi101534genetics106062711"
}

Biological evolution is a fact--but the many conflicting theories of evolution remain controversial even today. In 1966, simple Darwinism, which holds that evolution functions primarily at the level of the individual organism, was threatened by opposing concepts such as group selection, a popular idea stating that evolution acts to select entire species rather than individuals. George Williams's famous argument in favor of the Darwinists struck a powerful blow to those in opposing camps. His Adaptation and Natural Selection, now a classic of science literature, is a thorough and convincing essay in defense of Darwinism; its suggestions for developing effective principles for dealing with the evolution debate and its relevance to many fields outside biology ensure the timelessness of this critical work.

@book{doi1015159781400820108,
    author = "Williams, George C.",
    title = "Adaptation and Natural Selection",
    year = "2008",
    booktitle = "Princeton University Press eBooks",
    abstract = "Biological evolution is a fact--but the many conflicting theories of evolution remain controversial even today. In 1966, simple Darwinism, which holds that evolution functions primarily at the level of the individual organism, was threatened by opposing concepts such as group selection, a popular idea stating that evolution acts to select entire species rather than individuals. George Williams's famous argument in favor of the Darwinists struck a powerful blow to those in opposing camps. His Adaptation and Natural Selection, now a classic of science literature, is a thorough and convincing essay in defense of Darwinism; its suggestions for developing effective principles for dealing with the evolution debate and its relevance to many fields outside biology ensure the timelessness of this critical work.",
    url = "https://doi.org/10.1515/9781400820108",
    doi = "10.1515/9781400820108",
    openalex = "W2020289104"
}

@article{doi101016jjpolmod200807002,
    author = "Arce, Daniel G. and Sandler, Todd",
    title = "Fitting in: Group effects and the evolution of fundamentalism",
    year = "2009",
    journal = "Journal of Policy Modeling",
    url = "https://doi.org/10.1016/j.jpolmod.2008.07.002",
    doi = "10.1016/j.jpolmod.2008.07.002",
    openalex = "W2100437646",
    references = "cooper2004group, doi1010160160932782900369, doi101016s0003347276801108, doi101016s0024630197809661, doi101038227520a0, doi101038246015a0, doi101046j14390310199900372x, doi101093oepgpf064, doi101111j251761611951tb00088x, doi1023071906951, doi1023072069338"
}

The origin of altruistic behavior, i.e. the behavior that is useful for a population or a species but goes at the expense of an altruistic individual, has long been a challenge for students of evolutionary biology. The populations with altruistic individuals thrive better than those without altruists; however, the altruists within a population thrive worse than the non-altruists and their prevalence in the population decreases due to individual selection. Under certain conditions, the strength of group selection, i.e. the competition between populations, can surpass the strength of individual selection; however, such conditions are rarely achieved in practice. It was suggested recently that chances for altruistic behavior to spread highly increase when it is controlled not by a single gene but by multiple independent genes substitutable in their effects on the phenotype of the individual. Here we confirm the original verbal model published as part of the frozen plasticity theory by numerical modeling of the spread of altruistic/selfish alleles in a metapopulation consisting of partly isolated groups of organisms (demes) interconnected by migration. We have shown that altruistic behavior coded by multiple substitutable genes can stably coexist with selfish behavior, even under relatively high mutation and migration rates, i.e. under such conditions where altruistic behavior coded by a single gene is quickly outcompeted in a metapopulation.

@misc{doi1048550arxiv10024204,
    author = "Kulich, Tomáš and Flegr, Jaroslav",
    title = "Effects of multiple gene control on the spread of altruism by group selection",
    year = "2010",
    booktitle = "arXiv (Cornell University)",
    abstract = "The origin of altruistic behavior, i.e. the behavior that is useful for a population or a species but goes at the expense of an altruistic individual, has long been a challenge for students of evolutionary biology. The populations with altruistic individuals thrive better than those without altruists; however, the altruists within a population thrive worse than the non-altruists and their prevalence in the population decreases due to individual selection. Under certain conditions, the strength of group selection, i.e. the competition between populations, can surpass the strength of individual selection; however, such conditions are rarely achieved in practice. It was suggested recently that chances for altruistic behavior to spread highly increase when it is controlled not by a single gene but by multiple independent genes substitutable in their effects on the phenotype of the individual. Here we confirm the original verbal model published as part of the frozen plasticity theory by numerical modeling of the spread of altruistic/selfish alleles in a metapopulation consisting of partly isolated groups of organisms (demes) interconnected by migration. We have shown that altruistic behavior coded by multiple substitutable genes can stably coexist with selfish behavior, even under relatively high mutation and migration rates, i.e. under such conditions where altruistic behavior coded by a single gene is quickly outcompeted in a metapopulation.",
    url = "https://doi.org/10.48550/arxiv.1002.4204",
    doi = "10.48550/arxiv.1002.4204",
    openalex = "W1884319009",
    references = "silva1999stochastic"
}

In spite of recent interest in sexual selection in females, debate exists over whether traits that influence female-female competition are sexually selected. This review uses female-female aggressive behavior as a model behavioral trait for understanding the evolutionary mechanisms promoting intrasexual competition, focusing especially on sexual selection. I employ a broad definition of sexual selection, whereby traits that influence competition for mates are sexually selected, whereas those that directly influence fecundity or offspring survival are naturally selected. Drawing examples from across animal taxa, including humans, I examine 4 predictions about female intrasexual competition based on the abundance of resources, the availability of males, and the direct or indirect benefits those males provide. These patterns reveal a key sex difference in sexual selection: Although females may compete for the number of mates, they appear to compete more so for access to high-quality mates that provide direct and indirect (genetic) benefits. As is the case in males, intrasexual selection in females also includes competition for essential resources required for access to mates. If mate quality affects the magnitude of mating success, then restricting sexual selection to competition for quantity of mates may ignore important components of fitness in females and underestimate the role of sexual selection in shaping female phenotype. In the future, understanding sex differences in sexual selection will require further exploration of the extent of mutual intrasexual competition and the incorporation of quality of mating success into the study of sexual selection in both sexes.

@article{doi101093behecoarr106,
    author = "Rosvall, Kimberly A.",
    title = "Intrasexual competition in females: evidence for sexual selection?",
    year = "2011",
    journal = "Behavioral Ecology",
    abstract = "In spite of recent interest in sexual selection in females, debate exists over whether traits that influence female-female competition are sexually selected. This review uses female-female aggressive behavior as a model behavioral trait for understanding the evolutionary mechanisms promoting intrasexual competition, focusing especially on sexual selection. I employ a broad definition of sexual selection, whereby traits that influence competition for mates are sexually selected, whereas those that directly influence fecundity or offspring survival are naturally selected. Drawing examples from across animal taxa, including humans, I examine 4 predictions about female intrasexual competition based on the abundance of resources, the availability of males, and the direct or indirect benefits those males provide. These patterns reveal a key sex difference in sexual selection: Although females may compete for the number of mates, they appear to compete more so for access to high-quality mates that provide direct and indirect (genetic) benefits. As is the case in males, intrasexual selection in females also includes competition for essential resources required for access to mates. If mate quality affects the magnitude of mating success, then restricting sexual selection to competition for quantity of mates may ignore important components of fitness in females and underestimate the role of sexual selection in shaping female phenotype. In the future, understanding sex differences in sexual selection will require further exploration of the extent of mutual intrasexual competition and the incorporation of quality of mating success into the study of sexual selection in both sexes.",
    url = "https://doi.org/10.1093/beheco/arr106",
    doi = "10.1093/beheco/arr106",
    openalex = "W2154986968",
    references = "doi101016jtree200603015, doi101111j15585646201001012x, doi105962bhltitle19780"
}

Natural selection operates both directly, via the impact of a trait upon the individual's own fitness, and indirectly, via the impact of the trait upon the fitness of the individual's genetically related social partners. These effects are often framed in terms of Hamilton's rule, rb - c > 0, which provides the central result of social-evolution theory. However, a number of studies have questioned the generality of Hamilton's rule, suggesting that it requires restrictive assumptions. Here, we use Fisher's genetical paradigm to demonstrate the generality of Hamilton's rule and to clarify links between different studies. We show that confusion has arisen owing to researchers misidentifying model parameters with the b and c terms in Hamilton's rule, and misidentifying measures of genotypic similarity or genealogical relationship with the coefficient of genetic relatedness, r. More generally, we emphasize the need to distinguish between general kin-selection theory that forms the foundations of social evolution, and streamlined kin-selection methodology that is used to solve specific problems.

@article{doi101111j14209101201102236x,
    author = "Gardner, Andy and West, Stuart A. and Wild, Geoff",
    title = "The genetical theory of kin selection",
    year = "2011",
    journal = "Journal of Evolutionary Biology",
    abstract = "Natural selection operates both directly, via the impact of a trait upon the individual's own fitness, and indirectly, via the impact of the trait upon the fitness of the individual's genetically related social partners. These effects are often framed in terms of Hamilton's rule, rb - c > 0, which provides the central result of social-evolution theory. However, a number of studies have questioned the generality of Hamilton's rule, suggesting that it requires restrictive assumptions. Here, we use Fisher's genetical paradigm to demonstrate the generality of Hamilton's rule and to clarify links between different studies. We show that confusion has arisen owing to researchers misidentifying model parameters with the b and c terms in Hamilton's rule, and misidentifying measures of genotypic similarity or genealogical relationship with the coefficient of genetic relatedness, r. More generally, we emphasize the need to distinguish between general kin-selection theory that forms the foundations of social evolution, and streamlined kin-selection methodology that is used to solve specific problems.",
    url = "https://doi.org/10.1111/j.1420-9101.2011.02236.x",
    doi = "10.1111/j.1420-9101.2011.02236.x",
    openalex = "W1539514422",
    references = "doi101016jevolhumbehav201008001, doi101017cbo9780511542053, doi101086303168, doi101093acprofoso97801992679720010001, doi101111j14209101200801681x, doi101111j15585646201001012x, doi1015159781400832019"
}

Ornaments, weapons and aggressive behaviours may evolve in female animals by mate choice and intrasexual competition for mating opportunities-the standard forms of sexual selection in males. However, a growing body of evidence suggests that selection tends to operate in different ways in males and females, with female traits more often mediating competition for ecological resources, rather than mate acquisition. Two main solutions have been proposed to accommodate this disparity. One is to expand the concept of sexual selection to include all mechanisms related to fecundity; another is to adopt an alternative conceptual framework-the theory of social selection-in which sexual selection is one component of a more general form of selection resulting from all social interactions. In this study, we summarize the history of the debate about female ornaments and weapons, and discuss potential resolutions. We review the components of fitness driving ornamentation in a wide range of systems, and show that selection often falls outside the limits of traditional sexual selection theory, particularly in females. We conclude that the evolution of these traits in both sexes is best understood within the unifying framework of social selection.

@article{doi101098rstb20110280,
    author = "Tobias, Joseph A. and Montgomerie, Robert and Lyon, Bruce E.",
    title = "The evolution of female ornaments and weaponry: social selection, sexual selection and ecological competition",
    year = "2012",
    journal = "Philosophical Transactions of the Royal Society B Biological Sciences",
    abstract = "Ornaments, weapons and aggressive behaviours may evolve in female animals by mate choice and intrasexual competition for mating opportunities-the standard forms of sexual selection in males. However, a growing body of evidence suggests that selection tends to operate in different ways in males and females, with female traits more often mediating competition for ecological resources, rather than mate acquisition. Two main solutions have been proposed to accommodate this disparity. One is to expand the concept of sexual selection to include all mechanisms related to fecundity; another is to adopt an alternative conceptual framework-the theory of social selection-in which sexual selection is one component of a more general form of selection resulting from all social interactions. In this study, we summarize the history of the debate about female ornaments and weapons, and discuss potential resolutions. We review the components of fitness driving ornamentation in a wide range of systems, and show that selection often falls outside the limits of traditional sexual selection theory, particularly in females. We conclude that the evolution of these traits in both sexes is best understood within the unifying framework of social selection.",
    url = "https://doi.org/10.1098/rstb.2011.0280",
    doi = "10.1098/rstb.2011.0280",
    openalex = "W2109893252",
    references = "doi101007978140206287210, doi101086303168, doi101111j15585646201001012x, doi105962bhltitle17416"
}

Cooperation in nature is a complex topic and its study has left scientists with many open questions. Over the past two decades research has been undertaken into how cooperation works in an evolutionary context and how we can emulate it for social analysis. Numerous computer models have been developed and analyzed, with many models formulated as spatial or network games. These games use various update rules to evolve cooperative strategies. Despite two decades of effort, arguably little progress has been made. This paper exposes some of the problems with these spatial and network games and shows why they are ill-suited to get any real answers. Recommendations on future research directions that might provide some insight are presented.

@article{doi101109cig20126374132,
    author = "Greenwood, Garrison W. and Avery, Phillipa",
    title = "Update rules, reciprocity and weak selection in evolutionary spatial games",
    year = "2012",
    abstract = "Cooperation in nature is a complex topic and its study has left scientists with many open questions. Over the past two decades research has been undertaken into how cooperation works in an evolutionary context and how we can emulate it for social analysis. Numerous computer models have been developed and analyzed, with many models formulated as spatial or network games. These games use various update rules to evolve cooperative strategies. Despite two decades of effort, arguably little progress has been made. This paper exposes some of the problems with these spatial and network games and shows why they are ill-suited to get any real answers. Recommendations on future research directions that might provide some insight are presented.",
    url = "https://doi.org/10.1109/cig.2012.6374132",
    doi = "10.1109/cig.2012.6374132",
    openalex = "W2028130218",
    references = "doi101109cec20126256426"
}

We study the evolution of both characteristics of reciprocity: the willingness to reward and the willingness to punish. First, both preferences for rewarding and preferences for punishing can survive provided that individuals interact within separate groups. Second, rewarders survive only in coexistence with self-interested preferences, but punishers either vanish or dominate the population entirely. Third, the evolution of preferences for rewarding and the evolution of preferences for punishing influence each other decisively. Rewarders can invade a population of self-interested players. The existence of rewarders enhances the evolutionary success of punishers, who then crowd out all other preferences. (JEL C71, C72, C73, D64, K42)

@article{doi101257aer1022914,
    author = "Herold, Florian",
    title = "Carrot or Stick? The Evolution of Reciprocal Preferences in a Haystack Model",
    year = "2012",
    journal = "American Economic Review",
    abstract = "We study the evolution of both characteristics of reciprocity: the willingness to reward and the willingness to punish. First, both preferences for rewarding and preferences for punishing can survive provided that individuals interact within separate groups. Second, rewarders survive only in coexistence with self-interested preferences, but punishers either vanish or dominate the population entirely. Third, the evolution of preferences for rewarding and the evolution of preferences for punishing influence each other decisively. Rewarders can invade a population of self-interested players. The existence of rewarders enhances the evolutionary success of punishers, who then crowd out all other preferences. (JEL C71, C72, C73, D64, K42)",
    url = "https://doi.org/10.1257/aer.102.2.914",
    doi = "10.1257/aer.102.2.914",
    openalex = "W3125391255",
    references = "cooper2004group, doi1010382011145a0, doi101038227520a0, doi101046j14390310199900372x, doi101126science1133755, doi101257aer1022i, doi101257aer901166, doi101257aer91273, doi101257jep143159, doi1023071367778, doi1023072951777"
}

This paper aims at detecting the presence of group structures in complex artificial societies by solely observing and analysing the interactions occurring among the artificial agents. Our approach combines: (1) an unsupervised method for clustering interactions into two possible classes, namely in-group and out-group, (2) reinforcement learning for deriving the existing levels of collaboration within the society, and (3) an evolutionary algorithm for the detection of group structures and the assignment of group identities to the agents. Under a case study of static societies - i.e. the agents do not evolve their social preferences - where agents interact with each other by means of the Ultimatum Game, our approach proves to be successful for small-sized social networks independently on the underlying social structure of the society; promising results are also registered for mid-size societies.

@article{doi101109alife20136602441,
    author = "Grappiolo, Corrado and Togelius, Julian and Yannakakis, Georgios N.",
    title = "Artificial evolution for the detection of group identities in complex artificial societies",
    year = "2013",
    abstract = "This paper aims at detecting the presence of group structures in complex artificial societies by solely observing and analysing the interactions occurring among the artificial agents. Our approach combines: (1) an unsupervised method for clustering interactions into two possible classes, namely in-group and out-group, (2) reinforcement learning for deriving the existing levels of collaboration within the society, and (3) an evolutionary algorithm for the detection of group structures and the assignment of group identities to the agents. Under a case study of static societies - i.e. the agents do not evolve their social preferences - where agents interact with each other by means of the Ultimatum Game, our approach proves to be successful for small-sized social networks independently on the underlying social structure of the society; promising results are also registered for mid-size societies.",
    url = "https://doi.org/10.1109/alife.2013.6602441",
    doi = "10.1109/alife.2013.6602441",
    openalex = "W2093582041",
    references = "doi101109cec20126256426"
}

We present a computational framework capable of inferring the existence of group identities, built upon social networks of reciprocal friendship, in Complex Adaptive Artificial Societies (CAAS) by solely observing the flow of interactions occurring among the agents. Our modelling framework infers the group identities by following two steps: first, it aims to learn the ongoing levels of cooperation among the agents and, second, it applies evolutionary computation, based on the learned cooperation values, to partition the agents into groups and assign group identities to the agents.

@article{doi10114524645762482722,
    author = "Grappiolo, Corrado and Togelius, Julian and Yannakakis, Georgios N.",
    title = "Interaction-based group identity detection via reinforcement learning and artificial evolution",
    year = "2013",
    abstract = "We present a computational framework capable of inferring the existence of group identities, built upon social networks of reciprocal friendship, in Complex Adaptive Artificial Societies (CAAS) by solely observing the flow of interactions occurring among the agents. Our modelling framework infers the group identities by following two steps: first, it aims to learn the ongoing levels of cooperation among the agents and, second, it applies evolutionary computation, based on the learned cooperation values, to partition the agents into groups and assign group identities to the agents.",
    url = "https://doi.org/10.1145/2464576.2482722",
    doi = "10.1145/2464576.2482722",
    openalex = "W1964745510",
    references = "doi101109cec20126256426"
}

@article{doi101016jmbs201401003,
    author = "Fontanari, José F. and Serva, Maurizio",
    title = "Nonlinear group survival in Kimura’s model for the evolution of altruism",
    year = "2014",
    journal = "Mathematical Biosciences",
    url = "https://doi.org/10.1016/j.mbs.2014.01.003",
    doi = "10.1016/j.mbs.2014.01.003",
    openalex = "W2163795031",
    references = "silva1999deterministic"
}

Many of the most important properties of human groups - including properties that may give one group an evolutionary advantage over another - are properly defined only at the level of group organization. Yet at present, most work on the evolution of culture has focused solely on the transmission of individual-level traits. I propose a conceptual extension of the theory of cultural evolution, particularly related to the evolutionary competition between cultural groups. The key concept in this extension is the emergent group-level trait. This type of trait is characterized by the structured organization of differentiated individuals and constitutes a unit of selection that is qualitatively different from selection on groups as defined by traditional multilevel selection (MLS) theory. As a corollary, I argue that the traditional focus on cooperation as the defining feature of human societies has missed an essential feature of cooperative groups. Traditional models of cooperation assume that interacting with one cooperator is equivalent to interacting with any other. However, human groups involve differential roles, meaning that receiving aid from one individual is often preferred to receiving aid from another. In this target article, I discuss the emergence and evolution of group-level traits and the implications for the theory of cultural evolution, including ramifications for the evolution of human cooperation, technology, and cultural institutions, and for the equivalency of multilevel selection and inclusive fitness approaches.

@article{doi101017s0140525x13001544,
    author = "Smaldino, Paul E.",
    title = "The cultural evolution of emergent group-level traits",
    year = "2014",
    journal = "Behavioral and Brain Sciences",
    abstract = "Many of the most important properties of human groups - including properties that may give one group an evolutionary advantage over another - are properly defined only at the level of group organization. Yet at present, most work on the evolution of culture has focused solely on the transmission of individual-level traits. I propose a conceptual extension of the theory of cultural evolution, particularly related to the evolutionary competition between cultural groups. The key concept in this extension is the emergent group-level trait. This type of trait is characterized by the structured organization of differentiated individuals and constitutes a unit of selection that is qualitatively different from selection on groups as defined by traditional multilevel selection (MLS) theory. As a corollary, I argue that the traditional focus on cooperation as the defining feature of human societies has missed an essential feature of cooperative groups. Traditional models of cooperation assume that interacting with one cooperator is equivalent to interacting with any other. However, human groups involve differential roles, meaning that receiving aid from one individual is often preferred to receiving aid from another. In this target article, I discuss the emergence and evolution of group-level traits and the implications for the theory of cultural evolution, including ramifications for the evolution of human cooperation, technology, and cultural institutions, and for the equivalency of multilevel selection and inclusive fitness approaches.",
    url = "https://doi.org/10.1017/s0140525x13001544",
    doi = "10.1017/s0140525x13001544",
    openalex = "W2108786427",
    references = "doi10103831383, doi101086289977, doi101371journalpbio0020439"
}

Human cooperation is highly unusual. We live in large groups composed mostly of non-relatives. Evolutionists have proposed a number of explanations for this pattern, including cultural group selection and extensions of more general processes such as reciprocity, kin selection, and multi-level selection acting on genes. Evolutionary processes are consilient; they affect several different empirical domains, such as patterns of behavior and the proximal drivers of that behavior. In this target article, we sketch the evidence from five domains that bear on the explanatory adequacy of cultural group selection and competing hypotheses to explain human cooperation. Does cultural transmission constitute an inheritance system that can evolve in a Darwinian fashion? Are the norms that underpin institutions among the cultural traits so transmitted? Do we observe sufficient variation at the level of groups of considerable size for group selection to be a plausible process? Do human groups compete, and do success and failure in competition depend upon cultural variation? Do we observe adaptations for cooperation in humans that most plausibly arose by cultural group selection? If the answer to one of these questions is "no," then we must look to other hypotheses. We present evidence, including quantitative evidence, that the answer to all of the questions is "yes" and argue that we must take the cultural group selection hypothesis seriously. If culturally transmitted systems of rules (institutions) that limit individual deviance organize cooperation in human societies, then it is not clear that any extant alternative to cultural group selection can be a complete explanation.

@article{doi101017s0140525x1400106x,
    author = "Richerson, Peter J. and Baldini, Ryan and Bell, Adrian V. and Demps, Kathryn and Frost, Karl and Hillis, Vicken and Mathew, Sarah and Newton, Emily K. and Naar, Nicole and Newson, Lesley and Ross, Cody T. and Smaldino, Paul E. and Waring, Timothy M. and Zefferman, Matthew",
    title = "Cultural group selection plays an essential role in explaining human cooperation: A sketch of the evidence",
    year = "2014",
    journal = "Behavioral and Brain Sciences",
    abstract = {Human cooperation is highly unusual. We live in large groups composed mostly of non-relatives. Evolutionists have proposed a number of explanations for this pattern, including cultural group selection and extensions of more general processes such as reciprocity, kin selection, and multi-level selection acting on genes. Evolutionary processes are consilient; they affect several different empirical domains, such as patterns of behavior and the proximal drivers of that behavior. In this target article, we sketch the evidence from five domains that bear on the explanatory adequacy of cultural group selection and competing hypotheses to explain human cooperation. Does cultural transmission constitute an inheritance system that can evolve in a Darwinian fashion? Are the norms that underpin institutions among the cultural traits so transmitted? Do we observe sufficient variation at the level of groups of considerable size for group selection to be a plausible process? Do human groups compete, and do success and failure in competition depend upon cultural variation? Do we observe adaptations for cooperation in humans that most plausibly arose by cultural group selection? If the answer to one of these questions is "no," then we must look to other hypotheses. We present evidence, including quantitative evidence, that the answer to all of the questions is "yes" and argue that we must take the cultural group selection hypothesis seriously. If culturally transmitted systems of rules (institutions) that limit individual deviance organize cooperation in human societies, then it is not clear that any extant alternative to cultural group selection can be a complete explanation.},
    url = "https://doi.org/10.1017/s0140525x1400106x",
    doi = "10.1017/s0140525x1400106x",
    openalex = "W2155058526",
    references = "doi101007978140206287210, doi101016jevolhumbehav201008001, doi101016jjhevol201007011, doi101017cbo9780511807763, doi101017s0140525x0999094x, doi101017s0140525x14000090, doi101017s0140525x14001356, doi1010370033295x982224, doi101073pnas0904312106, doi101086303168, doi101086668207, doi101093oxfordjournalscjea013725, doi101098rspb20091650, doi101098rspb20151019, doi101098rstb20090052, doi101111j14677687200700573x, doi101257aer10052060, doi101257aer991431, doi101371journalpone0045150, doi101537ase188722495, doi1023071367778, doi1023072232409, doi1023073146384, doi105860choice416654, doi105860choice485062, openalexw1581387623, openalexw2034328075, openalexw2624262714, openalexw3124182182"
}

Biological evolution is a fact-but the many conflicting theories of evolution remain controversial even today. When Adaptation and Natural Selection was first published in 1966, it struck a powerful blow against those who argued for the concept of group selection-the idea that evolution acts to select entire species rather than individuals. Williams's famous work in favor of simple Darwinism over group selection has become a classic of science literature, valued for its thorough and convincing argument and its relevance to many fields outside of biology. Now with a new foreword by Richard Dawkins, Adaptation and Natural Selection is an essential text for understanding the nature of scientific debate

@book{doi1015159780691185507,
    author = "Williams, George C.",
    title = "Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought",
    year = "2018",
    abstract = "Biological evolution is a fact-but the many conflicting theories of evolution remain controversial even today. When Adaptation and Natural Selection was first published in 1966, it struck a powerful blow against those who argued for the concept of group selection-the idea that evolution acts to select entire species rather than individuals. Williams's famous work in favor of simple Darwinism over group selection has become a classic of science literature, valued for its thorough and convincing argument and its relevance to many fields outside of biology. Now with a new foreword by Richard Dawkins, Adaptation and Natural Selection is an essential text for understanding the nature of scientific debate",
    url = "https://doi.org/10.1515/9780691185507",
    doi = "10.1515/9780691185507",
    openalex = "W1480083809"
}

@article{doi101007s00285019013525,
    author = "Cooney, Daniel B.",
    title = "The replicator dynamics for multilevel selection in evolutionary games",
    year = "2019",
    journal = "Journal of Mathematical Biology",
    url = "https://doi.org/10.1007/s00285-019-01352-5",
    doi = "10.1007/s00285-019-01352-5",
    openalex = "W2898978090",
    references = "doi101007s0018201806301"
}

The literature on the evolution of preferences of individuals in strategic interactions is vast and diverse. We organize the discussion around the following question: Supposing that material outcomes drive evolutionary success, under what circumstances does evolution promote Homo economicus, defined as material self-interest, and when does it instead lead to other preferences? The literature suggests that Homo economicus is favored by evolution only when individuals’ preferences are their private information and the population is large and well-mixed, so that individuals with rare mutant preferences almost never get to interact with each other. If rare mutants instead interact more often (say, due to local dispersion), then evolution instead favors a certain generalization of Homo economicus including a Kantian concern. If individuals interact under complete information about preferences, then evolution destabilizes Homo economicus in virtually all games.

@article{doi101146annureveconomics080218030255,
    author = "Alger, Ingela and Weibull, Jörgen W.",
    title = "Evolutionary Models of Preference Formation",
    year = "2019",
    journal = "Annual Review of Economics",
    abstract = "The literature on the evolution of preferences of individuals in strategic interactions is vast and diverse. We organize the discussion around the following question: Supposing that material outcomes drive evolutionary success, under what circumstances does evolution promote Homo economicus, defined as material self-interest, and when does it instead lead to other preferences? The literature suggests that Homo economicus is favored by evolution only when individuals’ preferences are their private information and the population is large and well-mixed, so that individuals with rare mutant preferences almost never get to interact with each other. If rare mutants instead interact more often (say, due to local dispersion), then evolution instead favors a certain generalization of Homo economicus including a Kantian concern. If individuals interact under complete information about preferences, then evolution destabilizes Homo economicus in virtually all games.",
    url = "https://doi.org/10.1146/annurev-economics-080218-030255",
    doi = "10.1146/annurev-economics-080218-030255",
    openalex = "W2912848664",
    references = "doi101007s0018201806301"
}

Innovation subsidy is of great significance to promoting enterprise innovation development. However, in recent years, the frequent occurrence of R&D subsidy deception in China has greatly reduced effectiveness of innovation. From the perspective of the strategic choice motivation of the innovation subject (including the enterprises, research institutions, and local governments), this paper constructs a multiplayer stochastic evolutionary game model. The influence of each variable on the subject strategy adoption is analyzed by simulation. There are two important findings in this paper. First, the paper confirms that there is an optimal boundary for the high subsidies received by enterprises and academic institutions, and the “subsidy boundary” is solved through the model. Second, this paper analyzes the effectiveness of the regulation of each variable through simulation and provides management and policy implications.

@article{doi10115520209640412,
    author = "Li, Junqiang and Ren, Hao and Zhang, Changcheng and Qing-xia, LI and Duan, Kaifeng",
    title = "Substantive Innovation or Strategic Innovation? Research on Multiplayer Stochastic Evolutionary Game Model and Simulation",
    year = "2020",
    journal = "Complexity",
    abstract = "Innovation subsidy is of great significance to promoting enterprise innovation development. However, in recent years, the frequent occurrence of R\&D subsidy deception in China has greatly reduced effectiveness of innovation. From the perspective of the strategic choice motivation of the innovation subject (including the enterprises, research institutions, and local governments), this paper constructs a multiplayer stochastic evolutionary game model. The influence of each variable on the subject strategy adoption is analyzed by simulation. There are two important findings in this paper. First, the paper confirms that there is an optimal boundary for the high subsidies received by enterprises and academic institutions, and the “subsidy boundary” is solved through the model. Second, this paper analyzes the effectiveness of the regulation of each variable through simulation and provides management and policy implications.",
    url = "https://doi.org/10.1155/2020/9640412",
    doi = "10.1155/2020/9640412",
    openalex = "W3019162526",
    references = "doi101007s0018201806301"
}

Sex-Ratio (SR) chromosomes are selfish X -chromosomes that distort Mendelian segregation and are commonly associated with inversions. These chromosomal rearrangements suppress recombination with Standard (ST) X -chromosomes and are hypothesized to maintain multiple alleles important for distortion in a single large haplotype. Here, we conduct a multifaceted study of the multiply inverted Drosophila p se udoobscura SR chromosome to understand the evolutionary history, genetic architecture, and present-day dynamics that shape this enigmatic selfish chromosome. The D. p se udoobscura SR chromosome has three nonoverlapping inversions of the right arm of the metacentric X -chromosome: basal, medial, and terminal. We find that 23 of 29 Mb of the D. p se udoobscura X -chromosome right arm is highly differentiated between the Standard and SR arrangements, including a 6.6 Mb collinear region between the medial and terminal inversions. Although crossing-over is heavily suppressed on this chromosome arm, we discover it is not completely eliminated, with measured rates indicating recombination suppression alone cannot explain patterns of differentiation or the near-perfect association of the three SR chromosome inversions in nature. We then demonstrate the ancient basal and medial inversions of the SR chromosome contain genes sufficient to cause weak distortion. In contrast, the younger terminal inversion cannot distort by itself, but contains at least one modifier gene necessary for full manifestation of strong sex chromosome distortion. By parameterizing population genetic models for chromosome-wide linkage disequilibrium with our experimental results, we infer that strong selection acts to maintain the near-perfect association of SR chromosome inversions in present-day populations. Based on comparative genomic analyses, direct recombination experiments, segregation distortion assays, and population genetic modeling, we conclude the combined action of suppressed recombination and strong, ongoing, epistatic selection shape the D. p se udoobscura SR arrangement into a highly differentiated chromosome.

@article{doi101534genetics120303460,
    author = "Fuller, Zachary L. and Koury, Spencer and Leonard, Christopher J. and Young, Randee E. and Ikegami, Kobe and Westlake, Jonathan and Richards, Stephen and Schaeffer, Stephen W. and Phadnis, Nitin",
    title = "Extensive Recombination Suppression and Epistatic Selection Causes Chromosome-Wide Differentiation of a Selfish Sex Chromosome in Drosophila pseudoobscura",
    year = "2020",
    journal = "Genetics",
    abstract = "Sex-Ratio (SR) chromosomes are selfish X -chromosomes that distort Mendelian segregation and are commonly associated with inversions. These chromosomal rearrangements suppress recombination with Standard (ST) X -chromosomes and are hypothesized to maintain multiple alleles important for distortion in a single large haplotype. Here, we conduct a multifaceted study of the multiply inverted Drosophila p se udoobscura SR chromosome to understand the evolutionary history, genetic architecture, and present-day dynamics that shape this enigmatic selfish chromosome. The D. p se udoobscura SR chromosome has three nonoverlapping inversions of the right arm of the metacentric X -chromosome: basal, medial, and terminal. We find that 23 of 29 Mb of the D. p se udoobscura X -chromosome right arm is highly differentiated between the Standard and SR arrangements, including a 6.6 Mb collinear region between the medial and terminal inversions. Although crossing-over is heavily suppressed on this chromosome arm, we discover it is not completely eliminated, with measured rates indicating recombination suppression alone cannot explain patterns of differentiation or the near-perfect association of the three SR chromosome inversions in nature. We then demonstrate the ancient basal and medial inversions of the SR chromosome contain genes sufficient to cause weak distortion. In contrast, the younger terminal inversion cannot distort by itself, but contains at least one modifier gene necessary for full manifestation of strong sex chromosome distortion. By parameterizing population genetic models for chromosome-wide linkage disequilibrium with our experimental results, we infer that strong selection acts to maintain the near-perfect association of SR chromosome inversions in present-day populations. Based on comparative genomic analyses, direct recombination experiments, segregation distortion assays, and population genetic modeling, we conclude the combined action of suppressed recombination and strong, ongoing, epistatic selection shape the D. p se udoobscura SR arrangement into a highly differentiated chromosome.",
    url = "https://doi.org/10.1534/genetics.120.303460",
    doi = "10.1534/genetics.120.303460",
    openalex = "W3046327068",
    references = "doi101086282886"
}

Abstract In recent years, the study of the evolution of non-compliant behaviour in public procurement has been widely developed due to the growing economic relevance of this phenomenon. When such a question is formalized in terms of a dynamical model, new insights can be pursued, related to the possible evolution from a situation with low dishonesty level to high dishonesty level or vice versa. The present model considers an evolutionary adaptation process explaining whether honest or dishonest behaviour prevails in society at any given time by assuming endogenous monitoring by the State. We will distinguish between a scenario in which firms converge to monomorphic configurations (all honest or all dishonest) and a scenario in which firms converge to polymorphic compositions (that is with coexistence of both groups), depending on the relevant parameters. By making use of both analytical tools and numerical simulations, the present work aims at explaining the effectiveness of economic policies to reduce or eliminate non-compliant behaviour. Social stigma is found to play a key role: if the “inner attitude toward honesty” of a country is not strong enough, then dishonesty cannot be ruled out. However, increasing both the fine level attached to dishonest behaviour and the monitoring effort by the State can reduce asymptotic dishonesty levels and escape form the dishonesty trap.

@article{doi101007s1020302100317y,
    author = "Coppier, Raffaella and Grassetti, Francesca and Michetti, Elisabetta",
    title = "Non-compliant behaviour in public procurement: an evolutionary model with endogenous monitoring",
    year = "2021",
    journal = "Decisions in Economics and Finance",
    abstract = "Abstract In recent years, the study of the evolution of non-compliant behaviour in public procurement has been widely developed due to the growing economic relevance of this phenomenon. When such a question is formalized in terms of a dynamical model, new insights can be pursued, related to the possible evolution from a situation with low dishonesty level to high dishonesty level or vice versa. The present model considers an evolutionary adaptation process explaining whether honest or dishonest behaviour prevails in society at any given time by assuming endogenous monitoring by the State. We will distinguish between a scenario in which firms converge to monomorphic configurations (all honest or all dishonest) and a scenario in which firms converge to polymorphic compositions (that is with coexistence of both groups), depending on the relevant parameters. By making use of both analytical tools and numerical simulations, the present work aims at explaining the effectiveness of economic policies to reduce or eliminate non-compliant behaviour. Social stigma is found to play a key role: if the “inner attitude toward honesty” of a country is not strong enough, then dishonesty cannot be ruled out. However, increasing both the fine level attached to dishonest behaviour and the monitoring effort by the State can reduce asymptotic dishonesty levels and escape form the dishonesty trap.",
    url = "https://doi.org/10.1007/s10203-021-00317-y",
    doi = "10.1007/s10203-021-00317-y",
    openalex = "W3123095917",
    references = "doi101007s0019101403753"
}

In this minireview, we address the trade-off between biological altruism (group adaptation result-ing from the ability of an organism to improve the fitness of an associate at the expense of its own fitness) and symbiogenesis — the evolutionary pathway based on genetic integration of non-related species. We address symbiogenesis as a multi-stage process, which involves for-mation of superspecific hereditary systems — functionally integral symbiogenomes (under the facultative partners’ interactions) reorganized into the structurally integral hologenomes (in the obligatory symbioses). The best studied case of symbiogenesis is represented by the evolution of the eukaryotic cell based on transformation of symbiotic bacteria into cellular organelles. This evolution is associated with the deep reduction of microsymbionts’ genomes and with allocation of their genes into the hosts. As a result, microsymbionts lost their Genetic INdividuality (GIN), characterized by an ability to implement DNA- and RNA-based template syntheses required for genome maintenance and expression. Under facultative symbiotic dependence on hosts, the par-tial loss of GIN is due to a “symbiont → host” altruism which in the N2-fixing microbe–plant symbioses results in formation of non-reproducible bacterial forms (e.g., intracellular bacteroids in rhizobia or multiple heterocysts in Nostoc). If micro-symbionts lose their ability of autonomous existence (e.g., in the vertically transmitted intracellular symbionts), they are switched to the “forced altruism” in which the GIN reduction is required for the stable persistence of symbionts in hosts. Therefore, organellogenesis involves the sequential increase of the symbionts’ de-pendency on hosts: conditional → facultative → obligatory → absolute. It is associated with the reorganization of microbes into semi-autonomous cellular components, which may be completely devoid of their own genomes.

@article{doi1021638spbu032021108,
    author = "Проворов, Н. А.",
    title = "Genetic individuality and interspecies altruism: modelling symbiogenesis using different types of symbiotic bacteria",
    year = "2021",
    journal = "Biological Communications",
    abstract = "In this minireview, we address the trade-off between biological altruism (group adaptation result-ing from the ability of an organism to improve the fitness of an associate at the expense of its own fitness) and symbiogenesis — the evolutionary pathway based on genetic integration of non-related species. We address symbiogenesis as a multi-stage process, which involves for-mation of superspecific hereditary systems — functionally integral symbiogenomes (under the facultative partners’ interactions) reorganized into the structurally integral hologenomes (in the obligatory symbioses). The best studied case of symbiogenesis is represented by the evolution of the eukaryotic cell based on transformation of symbiotic bacteria into cellular organelles. This evolution is associated with the deep reduction of microsymbionts’ genomes and with allocation of their genes into the hosts. As a result, microsymbionts lost their Genetic INdividuality (GIN), characterized by an ability to implement DNA- and RNA-based template syntheses required for genome maintenance and expression. Under facultative symbiotic dependence on hosts, the par-tial loss of GIN is due to a “symbiont → host” altruism which in the N2-fixing microbe–plant symbioses results in formation of non-reproducible bacterial forms (e.g., intracellular bacteroids in rhizobia or multiple heterocysts in Nostoc). If micro-symbionts lose their ability of autonomous existence (e.g., in the vertically transmitted intracellular symbionts), they are switched to the “forced altruism” in which the GIN reduction is required for the stable persistence of symbionts in hosts. Therefore, organellogenesis involves the sequential increase of the symbionts’ de-pendency on hosts: conditional → facultative → obligatory → absolute. It is associated with the reorganization of microbes into semi-autonomous cellular components, which may be completely devoid of their own genomes.",
    url = "https://doi.org/10.21638/spbu03.2021.108",
    doi = "10.21638/spbu03.2021.108",
    openalex = "W3144068874",
    references = "doi101073pnas751385"
}

Abstract This paper defends global realism about the units of selection, the view that there is always (or nearly always) an objective fact of the matter concerning the level at which natural selection acts. The argument proceeds in two stages. First, it is argued that global conventionalist-pluralism is false. This is established by identifying plausible sufficient conditions for irreducible selection at a particular level, and showing that these conditions are sometimes satisfied in nature. Second, it is argued that local pluralism – the view that while realism is true of some selection regimes, pluralist conventionalism holds for others – should also be rejected. I show that the main arguments for local pluralism are consistent with global realism. I also suggest that local pluralism offers an unacceptably disunified view of the metaphysics of selection. It follows that we should accept global realism. But this leaves open the question of how to classify so called ‘multi-level selection type 1’ (MLS1) processes, such as Wilson’s classic trait-group model for the evolution of altruism: should they be interpreted as particle selection or collective selection? On the assumption of global realism, at most one of these is correct. I argue, against global realists such as Sober, that MLS1 processes should be understood as particle, not collective, selection, due to three features of MLS1: the reducibility of collective fitness, the absence of collective reproduction, and the dispensable role of collectives.

@article{doi101007s1053902309931z,
    author = "Boucher, Sandy C.",
    title = "An argument for global realism about the units of selection",
    year = "2023",
    journal = "Biology \& Philosophy",
    abstract = "Abstract This paper defends global realism about the units of selection, the view that there is always (or nearly always) an objective fact of the matter concerning the level at which natural selection acts. The argument proceeds in two stages. First, it is argued that global conventionalist-pluralism is false. This is established by identifying plausible sufficient conditions for irreducible selection at a particular level, and showing that these conditions are sometimes satisfied in nature. Second, it is argued that local pluralism – the view that while realism is true of some selection regimes, pluralist conventionalism holds for others – should also be rejected. I show that the main arguments for local pluralism are consistent with global realism. I also suggest that local pluralism offers an unacceptably disunified view of the metaphysics of selection. It follows that we should accept global realism. But this leaves open the question of how to classify so called ‘multi-level selection type 1’ (MLS1) processes, such as Wilson’s classic trait-group model for the evolution of altruism: should they be interpreted as particle selection or collective selection? On the assumption of global realism, at most one of these is correct. I argue, against global realists such as Sober, that MLS1 processes should be understood as particle, not collective, selection, due to three features of MLS1: the reducibility of collective fitness, the absence of collective reproduction, and the dispensable role of collectives.",
    url = "https://doi.org/10.1007/s10539-023-09931-z",
    doi = "10.1007/s10539-023-09931-z",
    openalex = "W4387014143",
    references = "barrett2002group, doi1010800258013620191706384, doi101093acprofoso97801992679720010001, doi101093acprofosobl97801995520470010001, doi101146annureves01110170000245, doi1015159780691185507, doi1023072185056, doi1023072186103, doi1023072410462, doi105860choice366578, doi107208chicago97802261786530010001, openalexw1541831804"
}

Abstract The formalism used to describe evolutionary change in a multilevel setting can be used equally to re-describe the situation as one where all the selection occurs at the individual level. Thus, whether multilevel or individual-level selection occurs seems to be a matter of convention rather than fact. Yet, group selection is regarded by some as an important concept with factual rather than conventional elements. I flesh out an alternative position that regards groups as a target of selection in a way that is not merely definitional fiat and provide a theoretical basis for this position.

@article{doi101007s10670023007495,
    author = "Bourrat, Pierrick",
    title = "Moving Past Conventionalism About Multilevel Selection",
    year = "2023",
    journal = "Erkenntnis",
    abstract = "Abstract The formalism used to describe evolutionary change in a multilevel setting can be used equally to re-describe the situation as one where all the selection occurs at the individual level. Thus, whether multilevel or individual-level selection occurs seems to be a matter of convention rather than fact. Yet, group selection is regarded by some as an important concept with factual rather than conventional elements. I flesh out an alternative position that regards groups as a target of selection in a way that is not merely definitional fiat and provide a theoretical basis for this position.",
    url = "https://doi.org/10.1007/s10670-023-00749-5",
    doi = "10.1007/s10670-023-00749-5",
    openalex = "W4389388826",
    references = "doi1023072026633"
}

This narrative review aims to contribute to the scientific literature on awe by reviewing seven perspectives on the evolutionary function of awe. Each is presented with accompanying empirical evidence and suggestions for research investigating unanswered questions. Based on the existing perspectives, this review proposes an integrated evolutionary model of awe, postulating the evolutionary selection of awe through three adaptive domains: (1) social cooperation, (2) reflective processing, and (3) signaling suitability as a potential mate.

@article{doi10117717540739231197199,
    author = "Lucht, Antonia and van Schie, Hein T.",
    title = "The Evolutionary Function of Awe: A Review and Integrated Model of Seven Theoretical Perspectives",
    year = "2023",
    journal = "Emotion Review",
    abstract = "This narrative review aims to contribute to the scientific literature on awe by reviewing seven perspectives on the evolutionary function of awe. Each is presented with accompanying empirical evidence and suggestions for research investigating unanswered questions. Based on the existing perspectives, this review proposes an integrated evolutionary model of awe, postulating the evolutionary selection of awe through three adaptive domains: (1) social cooperation, (2) reflective processing, and (3) signaling suitability as a potential mate.",
    url = "https://doi.org/10.1177/17540739231197199",
    doi = "10.1177/17540739231197199",
    openalex = "W4386294224",
    references = "cooper2004group, doi101017cbo9780511620539, doi101037002235146761049, doi101037h0054346, doi10108002699930302297, doi101126science164387586, doi101146annurevpsych59103006093629, openalexw1555328317, openalexw1575922411, openalexw2600536710, openalexw2908473373"
}

Humans often cooperate in groups with friends and family members with varying degrees of genetic relatedness. Past kin selection can also be relevant to interactions between strangers, explaining how the cooperation first arose in the ancestral population. However, modelling the effects of relatedness is difficult when the benefits of cooperation scale nonlinearly with the number of cooperators (e.g., economies of scale). Here, we present a direct fitness method for rigorously accounting for kin selection in n-player interactions with m discrete strategies, where a genetically homophilic group-formation model is used to calculate the necessary higher-order relatedness coefficients. Our approach allows us to properly account for non-additive fitness effects between relatives (synergy). Analytical expressions for dynamics are obtained, and they can be solved numerically for modestly sized groups and numbers of strategies. We illustrate with an example where group members can verbally agree (cheap talk) to contribute to a public good with a sigmoidal benefit function, and we find that such coordinated cooperation is favoured by kin selection. As interactions switched from family to strangers, in order for coordinated cooperation to persist and for the population to resist invasion by liars, either some level of homophily must be maintained or following through on the agreement must be in the self-interests of contributors. Our approach is useful for scenarios where fitness effects are non-additive and the strategies are best modelled in a discrete way, such as behaviours that require a cognitive 'leap' of insight into the situation (e.g., shared intentionality, punishment).

@article{doi101016jjtbi2025112089,
    author = "Kristensen, Nadiah P. and Chisholm, Ryan A. and Ohtsuki, Hisashi",
    title = "Many-strategy games in groups with relatives and the evolution of coordinated cooperation",
    year = "2025",
    journal = "Journal of Theoretical Biology",
    abstract = "Humans often cooperate in groups with friends and family members with varying degrees of genetic relatedness. Past kin selection can also be relevant to interactions between strangers, explaining how the cooperation first arose in the ancestral population. However, modelling the effects of relatedness is difficult when the benefits of cooperation scale nonlinearly with the number of cooperators (e.g., economies of scale). Here, we present a direct fitness method for rigorously accounting for kin selection in n-player interactions with m discrete strategies, where a genetically homophilic group-formation model is used to calculate the necessary higher-order relatedness coefficients. Our approach allows us to properly account for non-additive fitness effects between relatives (synergy). Analytical expressions for dynamics are obtained, and they can be solved numerically for modestly sized groups and numbers of strategies. We illustrate with an example where group members can verbally agree (cheap talk) to contribute to a public good with a sigmoidal benefit function, and we find that such coordinated cooperation is favoured by kin selection. As interactions switched from family to strangers, in order for coordinated cooperation to persist and for the population to resist invasion by liars, either some level of homophily must be maintained or following through on the agreement must be in the self-interests of contributors. Our approach is useful for scenarios where fitness effects are non-additive and the strategies are best modelled in a discrete way, such as behaviours that require a cognitive 'leap' of insight into the situation (e.g., shared intentionality, punishment).",
    url = "https://doi.org/10.1016/j.jtbi.2025.112089",
    doi = "10.1016/j.jtbi.2025.112089",
    openalex = "W4408979304",
    references = "doi101007s0018201806301"
}