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eBook Evolutionary Restraints: The Contentious History of Group Selection ePub

eBook Evolutionary Restraints: The Contentious History of Group Selection ePub

by Mark E. Borrello

  • ISBN: 0226067033
  • Category: Evolution
  • Subcategory: Math Science
  • Author: Mark E. Borrello
  • Language: English
  • Publisher: University of Chicago Press; Reprint edition (May 15, 2012)
  • Pages: 228
  • ePub book: 1435 kb
  • Fb2 book: 1910 kb
  • Other: doc lrf rtf mobi
  • Rating: 4.6
  • Votes: 739

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PDF On Jan 1, 2013, Eugene Earnshaw and others published Mark Borrello, Evolutionary Restraints: The Contentious . that was my impression. But Borrello's Evolutionary Restraints contextualises and corrects this view

PDF On Jan 1, 2013, Eugene Earnshaw and others published Mark Borrello, Evolutionary Restraints: The Contentious History of Group Selection. But Borrello's Evolutionary Restraints contextualises and corrects this view. Borello's work, despite its generic title, focuses closely on Wynne-Edwards and his struggle to express and defend his. views on group selection. In so doing, it provides an illuminating case study of scientific theory change. as well and motivates a reappraisal of Wynne-Edwards' work in light of current theory. The core of the book chronologically traces Wynne-Edwards development, presentation, and.

As the subtitle of Mark Borrello's Evolutionary Restraints indicates, this book is a detailed genealogy of the & history of group selection' in evolutionary biology, which Borrello explores, in part.

As the subtitle of Mark Borrello's Evolutionary Restraints indicates, this book is a detailed genealogy of the & history of group selection' in evolutionary biology, which Borrello explores, in part, through the biography of Vero Copner Wynne Edwards, the man whose name has become synonymous with the concept following the 1962 publication of Animal Dispersal in relation to Social Behaviour. As Borello shows, though, the history of group selection is deeper and broader than this

Request PDF On Mar 1, 2013, Patricia Princehouse and others published Mark E. Borrello. The aim of this paper is to provide information on the history of MELEO 2, its configuration, actual status, opportunity of flight and the selection criteria

Request PDF On Mar 1, 2013, Patricia Princehouse and others published Mark E. Evolutionary Restraints: The Contentious History of Group Selection. The aim of this paper is to provide information on the history of MELEO 2, its configuration, actual status, opportunity of flight and the selection criteria. General information is provided on the experiments that have been proposed in response to the informal Announcement of Opportunity (AO). A copy of the AO is attached.

Evolutionary Restraints: The Contentious History of Group Selection. Most biologists acknowledge multiple levels of selection-from the gene to the species. The debate about group selection, however, is the focus of Mark E. Borrello’s Evolutionary Restraints. Tracing the history of biological attempts to determine whether selection leads to the evolution of fitter groups, Borrello takes as his focus the British naturalist V. C. Wynne-Edwards, who proposed that animals could regulate their own populations and thus avoid overexploitation of their resources.

The debate about group selection, however, is the focus of Mark E. Borrello’s Evolutionary Restraints

The debate about group selection, however, is the focus of Mark E. Wynne-Edwards, who proposed that animals could regulate their own populations and thus avoid overexploitation of their resources

The publication of Mark Borrello’s Evolutionary Restraints: The Contentious History of Group Selection13 couldn’t have been better timed to bring context to the history of this dispute.

The publication of Mark Borrello’s Evolutionary Restraints: The Contentious History of Group Selection13 couldn’t have been better timed to bring context to the history of this dispute.

Evolutionary Restraints book. However, it is the debate about group selection that Mark E. Borrello focuses on in Evolutionary Restraints. Tracing the history of biological attempts to determine whether selection could lead to the evolution of fitter groups, Borrello takes as his focus the British naturalist V. Wynne-Edwards, who proposed that animals could regulate their own population levels and thereby avoid overexploitation of their food and other resources.

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 Much of the evolutionary debate since Darwin has focused on the level at which natural selection occurs. Most biologists acknowledge multiple levels of selection—from the gene to the species. The debate about group selection, however, is the focus of Mark E. Borrello’s Evolutionary Restraints.            Tracing the history of biological attempts to determine whether selection leads to the evolution of fitter groups, Borrello takes as his focus the British naturalist V. C. Wynne-Edwards, who proposed that animals could regulate their own populations and thus avoid overexploitation of their resources. By the mid-twentieth century, Wynne-Edwards became an advocate for group selection theory and led a debate that engaged the most significant evolutionary biologists of his time, including Ernst Mayr, G. C. Williams, and Richard Dawkins. This important dialogue bled out into broader conversations about population regulation, environmental crises, and the evolution of human social behavior. By examining a single facet in the long debate about evolution, Borrello provides powerful insight into an intellectual quandary that remains relevant and alive to this day.

Comments

Joni_Dep Joni_Dep
As the subtitle of Mark Borrello's Evolutionary Restraints indicates, this book is a detailed genealogy of the `contentious history of group selection' in evolutionary biology, which Borrello explores, in part, through the biography of Vero Copner Wynne Edwards, the man whose name has become synonymous with the concept following the 1962 publication of Animal Dispersal in relation to Social Behaviour. As Borello shows, though, the history of group selection is deeper and broader than this. Not only did the Russian geographer and zoologist Peter Kropotkin explore the idea fully in the 1880s and `90s, in his protracted arguments against the Malthusian individualism of `Darwin's Bulldog' Thomas Huxley, but, (and in vindication of Kropotkin's argument), Borrello shows that Darwin also explicitly invoked group selection not only in relation to the evolution of the social insects, but also and more fully, in relation to the social instincts - both of the higher animals and of man. Thus, in contrast to the partisan opinions of the historian of science Michael Ruse (1980) and more recently of Helena Cronin (1991), Darwin and Wallace were not the fountainhead of individual-selectionist orthodoxy, and were in fact quite comfortable appealing to group selection when it suited them. Further, so too did almost every other nineteenth and early-twentieth century biologist of significance, the neo-Darwinian August Weismann among them. As Borrello points out, this much was hardly surprising given debate about the compound individuality of primitive "communal organisms" such as the marine invertebrate Portugese Man-O'-War and coral polyps.

It is chapter three before we meet Wynne Edwards, and Borrello allows the story he tells of the development of Wynne Edwards' ideas to echo those of Kropotkin quite explicitly. Wynne Edwards own history took him from Oxford to McGill in Canada before later returning to Aberdeen. Like Kropotkin, who had surveyed the expansive Siberian Steppes, the environment in which Wynne Edwards pursued his field studies were similarly harsh and sparsely populated; to both men it was evident that the `struggle for existence' went on between organisms and their environment much more so than between competing organisms. By 1939, in his paper on the intermittent breeding of Fulmars published in the Proceedings of the Zoological Society of London, Wynne Edwards was already framing his observations of bird populations in the context of the group selectionist theory with which his name was to become inextricably linked. Wynne Edwards estimated that only 1/3rd - 2/5ths of the breeding colony appeared to be engaged in rearing offspring and hypothesized that this non-breeding behavior was an evolved strategic behavior to regulate their numbers in order to avoid the threat of over-population.

This explanation of the social behavior of breeding populations brought Wynne Edwards into conflict with fellow ornithologist David Lack. Lack, who had been encouraged to study what we now know as `Darwin's finches' on the Galapagos Islands by Julian Huxley, came to quite different conclusions. Lack was the first to make the case that the variable beak morphology of the finches was adaptive. These adaptations were the result of selection operating not - as Wynne Edwards had argued regarding his Fulmars, to prevent the population running up against limited resources, - but as a result of population running up against limited resources. As Rosemary and Peter Grant would later illustrate, `Darwin's finches' are a perfect illustration of the theory of divergence that Darwin had outlined in Origin. The battle lines between Lack and Wynne Edwards were publically drawn at a session of the British Ornithological Union on population ecology, the proceedings of which were later published in the journal Ibis in 1959. As Borrello tells it, the battle of words was carried on by proxy, with Wynne Edwards stuck in the United States at the time, George Dunnett, one of his students, presented in his stead. It was in "Control of Population Density through Social Behaviour, A Hypothesis", the paper that Dunnett read, that Wynne Edwards suggested that populations self -regulated through social conventions informed by what he called `epideictic' behavior - behavior through which populations actively assessed their own population density. With Lack on home ground, he sealed the advantage, and this was set the tone for the professional reception of group selection theory henceforward. Lack argued that there were alternative explanations to each of the behaviours that Wynne Edwards reported, explanations that supported Lack's own hypothesis. As Borrello's research shows, however, Lack had no better empirical grounding for his own theory than his opponent.

It is important to recognize, and this is one of the key contributions that Borrello makes, that Wynne Edwards was not the lone group-selectionist voice in the wilderness that received wisdom would suggest. Indeed, he was encouraged not only by the population-centred theoretical studies of the synthesis to see his own work as a part of the intellectual vanguard of modern biology, but by the fact that almost all the main players in the synthesis entertained some level of group selectionist thought. An analysis of first edition synthesis literature shows that E.B. Ford, Theodosius Dobzhansky, Sewewll Wright, Ernst Mayr, G.G. Simpson and Julian Huxley, among others, each acknowledged that selection occurred across a hierarchy of levels. As Wynne Edwards himself noted in a 1948 paper to the Oxford Ornithological Society, "The fundamental new idea is that populations, rather than independent individuals, are the basic units upon which evolutionary processes act" (54).

The context of the prevailing state of the discipline was important, and despite the loudly proclaimed unity in biology that accompanied the 1859 centenary of Origin, the reaction to the publication of Wynne Edwards Animal Dispersal in Relation to Social Behaviour in 1962 reveals that there was still much that remained unsettled on what might reasonably seen to be some fairly basic issues. It was a monumental work of some 653 pages, and although it provided little in the way of statistical analysis or sophisticated population modeling, it was packed with decades of detailed field observations in support of Wynne Edwards' hypothesis. As Borello notes, the book was consciously modeled on the concilient structure that Darwin had employed in Origin, it was `one long argument' linking behavior and population size. Selection at the level of the group, Wynne Edwards concluded, "was much more important that selection at the individual level" (80).

Public concern over broader environmental issues about sustainability and population growth that would later fuel enthusiasm for Paul Erlich's 1968 The Population Bomb, ensured Wynne Edward's book a ready audience. Non-specialist commentators read his book as corroborating their own beliefs that any number of modern social ills and neuroses from homosexuality to increasing youth crime could be explained in terms of a population stressed beyond its limits. Initial public acclaim, however, gave way to professional ambivalence. Even former mentors like animal ecologist Charles Elton were little more than lukewarm, confessing he found Wynne Edwards' account not only dogmatic, but its author guilty of "woolly thinking" (86). Lack was not far behind, and even less kind in his appraisal, affronted, perhaps, that Wynne Edwards had appropriated the term `dispersal' from his own Natural Regulation of Animal Numbers (1954) only to give the term an entirely different meaning than Lack had employed. To Lack it had signified no more than the non-random distribution of species, Wynne Edwards, however, implied the result of a communally policed strategy. It is notable, though, that although Lack's criticisms were perhaps the loudest, as Borello points out, his argument relied primarily on parsimony and a sympathetic ear from the individualist inclinations of the post-synthesis community.

G.C. Williams made Lack's rejection of Wynne Edwards orthodox in his now canonical Adaptation and Natural Selection (1966). In it Williams clearly stated that identification of adaptations "should be attributed to no higher level of organisation than is demanded by the evidence" (107-8). Further, selection would always act to maximize the mean reproductive level, over and above any role that multiple environments played in determining selection coefficients. Most damning, though, was his rejection outright of group selection which he presented as fundamentally opposed to genic selection. In private correspondence to Lack, Williams confessed he found Wynne Edwards' thesis laughable and had found it difficult "to avoid the appearance of sarcasm or ridicule" (111).
Wynne Edwards faired little better at the hands of ethologists, Niko Tinbergen and Konrad Lorenz, despite the superficial similarities of their research interests also found reason to differ. Tinbergen's focus was on fitness that particular behaviours bestowed upon individuals, Lorenz stressed the invariability of instinctive behaviours. Wynne Edwards perspective kept him always an outsider in the developing field. The fact that Tinbergen and Lorenz shared the 1973 Nobel Prize for Physiology and Medicine with Karl von Frisch, while Wynne Edwards was only further marginalized, only set the tone for future developments.

The increasing emphasis upon mathematical modeling at the molecular level cannot be under-estimated in keeping Wynne Edwards work under fire. G.C. Williams, W.D. Hamilton and John Maynard Smith proffered gene-level explanations for all the observed phenomena that Wynne Edwards described. Even though Maynard Smith's Haystack modeling showed that group selection was a theoretical possibility, the parameters of the model showed it not only to be extremely unlikely, but that any altruistic community that came about by such mechanisms were fragile, being vulnerable to invasion by selfish individuals. Indeed, it quickly became the go-to weapon in the arsenal of critics of group selection. E.O. Wilson's Sociobiology and Richard Dawkins' Selfish Gene took popular biology to a new level and in doing so sealed the fate of group selection in what Borrello calls "the dark ages of group selection" (136). Dawkins made explicit the contempt for group selection theory that Williams had kept private.
Wynne Edwards was not to be silenced, however, finally publishing his second magnum opus, Evolution through Group Selection in 1986, in which he attempted to explicitly align his own work with Sewell Wright's much earlier simple quantitative model showing that individually disadvantageous, but socially advantageous traits could evolve through group selection. "My task" he wrote, "is obviously to show that the group selection I advocate is not essentially different from Wright's in raising average individual fitness, and is not fragile either", his intention, Borrello notes, was clearly to "salvage his theory from oblivion" (145).

It was not to be. Wynne Edwards was either unable or unwilling to adapt his position on group selection to adequately take account of the many advances that had been made in the field since 1962. Being seen to be talking past rather than to his colleagues undermined his professional reputation and, in effect, Borrello argues, "he essentially wrote himself out of the debate" (147). In the intervening years, philosophers and biologists had taken the debate to new and theoretically more sophisticated territory. David Sloan Wilson and Michael Wade had embraced kin selection as one form of group selection, negating the either-or scenario that Williams had painted, and D.S. Wilson and Elliott Sober challenged the limited presumptions of Maynard Smith's Haystack model of group selection. Selection at the level of the group, they demonstrated, was not only mathematically plausible, but a highly likely, and a highly stable possibility.

As Borello concludes, Wynne Edwards' attempts to remain a significant player in the debate over group selection as it became reframed around the levels of selection were frustrated in part by the mathematical turn in theoretical biology, but, perhaps more seriously, were dogged by his continued refusal to engage with developments in the field, and this despite the pleas of editors and referees. He was not helped by the fact that few journal editors were willing to entertain what was by now deemed out-dated heterodoxy.
But what should we make of this story? What was the significance of Wynne Edwards' career in biology? His work certainly remains well-cited, although one suspects that this is as much in reference to how not to proceed as out of endorsement. Borrello contends that his work and this story is notable for a number of reasons. Certainly Wynne Edwards challenged others to think deeply about how one might account for the observed phenomena as a result of either genic or organismic selection, however, in light of more recent, if grudging, acceptance of hierarchical selection, Wynne Edwards also served as an acknowledged influence for present day advocates of group selection. It might be said that his persistence paid off, perhaps, even if it was this same blinkered insistence that ultimately undermined his own credibility.

Further, though, this story is also about the nature of science and about the peculiar relationship between the empirical world and our best perceptions of it. Borello asks, when Kropotkin and Wynne Edwards look at the world and see what they see, and when Huxley and, say, Williams look at the world and see what they see, what is it that leads them to such different conclusions? In this instance, both Kropotkin and Wynne Edwards approached nature with a full appreciation of Darwinian natural selection. With Kropotkin and Huxley we might appeal to their deeply held political commitments. As an anarchist, Kropotkin just could not see the competition that to Huxley's liberal mind was only too evident. To what extent is biology still deeply imbued with politics? As Borrello points out, there remains a lot at stake about the nature of man in the question over whether altruism can ever be truly genuine, as well as about whether the group dynamic is the faciltator or the oppressor of personal liberties. It would be strange if the politics of biology were not at work in the post World War Two to cold war years just as they were in the industrial revolution. As Borrello notes, it is here that historians not only have an insight into what occurred in the past in this debate, but have an opportunity to contribute to ongoing discussions of the hierarchical nature of evolutionary theory. In addition, of course, the history of a science is relevant to its practitioners too, not only in terms of thinking about the processes and practices of science, but in the sense that, as Borrello points out, many of the common misunderstandings of evolution to which students in biology frequently fall victim, are a direct result of the exclusive application of natural selection at the level of the individual.

In sum, Evolutionary Restraints is a compelling book and a challenging one. It provides us with a new perspective on the history of group selection that really does make us have to rethink its most vociferous condemnation in preference to selection at the level of the gene. The prevalence of group selectionist ideas throughout the synthesis years cannot be ignored in the same way that a solitary figure, however prominent, can be marginalized and cast as eccentric and imprecise. Sewell Wright, and Dobzhnsky, as well as Mayr were only three notable figures who argued that selection occurred at multiple levels prior to what has been referred to as the `hardening' or even the `constriction' of the synthesis. Mayr, indeed, insisted that the individual organism, as the entity that interacted with the environment, must be the entity exposed to selection, and not merely the genes it carried. Of course, he acknowledged, counting genes was important, but this much was really just so much book-keeping without an awareness of the interactions that led to their survival. This, of course, demands attention not only to the gene, but to the individual in the context of the population and the environment.
Teonyo Teonyo
George Santayana one famously said "Those who cannot remember the past are condemned to repeat it." Perhaps even those who remember the past may be condemned to repeat it, unless they learn sufficiently from it. Borrello adds a bit to the well-known history of the debate over levels of selection, but he does not help us understand what the real issues are, and how they might be resolved. Perhaps it is not his fault: the whole issue seems to be a hopeless muddle even in the hands of the most adept contemporary thinkers--present company excepted, of course. ;)

Biologists have debated the nature and status of group selection since the early twentieth century. The debate continues. Curiously, there have always been both extremely eminent supporters and equally eminent detractors of group selection, such as J. B. S. Haldane vs. Ernst Mayr (early years), John Maynard Smith vs. Stephen Jay Gould (recent past), and most recently Edward O. Wilson vs. Richard Dawkins. So pitched has this debate been that in reviewing Wilson's book The Social Conquest of Earth (which eloquently defends and extends group selection arguments), Richard Dawkins (who treats such arguments with characteristic distain) wrote "To borrow from Dorothy Parker, this is not a book to be tossed lightly aside. It should be thrown with great force." My more temperate and supportive review of Wilson's book, by the way, is published in BioScience and entitled The Clash of the Titans The paper is available from my web site.

To add fuel to the fire, in a response to a paper in Nature in 2010 by Wilson and coauthors Martin Nowak and Corina Tarnita that trashes inclusive fitness and extols group selection, some 150 well-known biologists wrote a vitriolic reply in a letter to Nature in 2010, resoundingly rejecting Wilson and coauthors' arguments. More heat, in my estimation, but less light.

I am an outsider to this debate. My training and most of my research until a couple of decades ago was in mathematics and economics. But I have come to work in biology as an adult, a situation that may have allowed me to avoid the emotional baggage attached to the dispute.

The group selection dispute shows the immaturity of contemporary population biology. No mature scientific dispute can sustain itself at so a high level of emotion and so low a level of progress for over a century without resolution. Indeed, most participants to the dispute agree that one can describe sociobiological behavior equally validly at the level of the gene, at the level of the individual (in a sexually reproducing species with a sequestered germ plasm), or at the group level (in species that live in groups engaged in social interaction). This admission, surprisingly, does not stop the disputants from intemperate diatribes at each other's expense.

Borrello's book does a competent if breezy and non-technical overview of this controversy, with some historical emphasis on particular papers presented by the disputants at different times. He does, however, seriously underestimate the importance of R. A. Fisher in laying the groundwork for the position of the Oxford school in this research area, and he has a long, unoriginal, and irrelevant analysis of Lamarckism. However, Borrello makes no attempt to cut through the verbiage and miscommunication to put the issue into insightful theoretical perspective. History, for Borrello, is not one damn thing after another---it is the same damn thing over and over!

So let me here explain what is really going on in the controversy over group selection. Population biology is based on careful accounting at the level of the gene (this is why Ernst Mayr criticized J. B. S. Haldane's work as "beanbag genetics"). Indeed, all of Darwinian evolution can be analyzed, at least in principle, in terms of changes in gene frequencies and correlations among genes in the genome. The two sides to the debate over group selection differ as to how correlations among genetic loci in the genome are to be treated. In the tradition of Fisher and Haldane, inspired by the mathematics of heredity developed to deal with animal husbandry, only the additive component of a gene's contribution to the fitness of its carriers need be analyzed because "evolution only sees the additive component." Those in the tradition of Ernst Mayr and Theodosius Dobzhansky, by contrast, consider the genome itself as a unit that is subject to natural selection, and it is the cooperative interaction of genes in the genome that governs its evolutionary success.

This distinction is then carried over from interactions among genes in the genome to interactions among individuals in a social species: inclusive fitness theorists, the heirs to the Fisher-Haldane gene's-eye view, reduce complex interactions among individuals in a social species to simple influences at single genetic locus (inclusive fitness), whereas the heirs to the Mayr-Dobzhansky tradition take social cooperation to involve complex dynamical and nonlinear interactions (multi-level selection). The inclusive fitness adherents argue that the genome cannot be an object of selection because it is broken up in every generation through meiosis and crossover, while the gene is conserved intact over many generations. The multilevel selection adherents claim that inclusive fitness analysis ignores all the substantive complexities of social interaction that in fact account for the evolutionary success of a social species.

My own studies have provided me with a third perspective. First, inclusive fitness analysis is extremely important and valid under almost all circumstances in dealing with sexually reproducing populations. Its strength comes from the fact that its reasoning is on the level of a single genetic locus. Popular expositions obfuscate this point by implicitly, without argument, treating a complex social behavior as governed by a single genetic locus---the so-called "phenotypic gambit." The correct analysis, that proposed by William Hamilton in 1964, shows that genes are utterly selfish in the sense popularized in Dawkins' great book "The Selfish Gene." What this means is that the conditions for the evolutionary success of a gene are distinct from the conditions under which it promotes the fitness of the reproduction population. Thus individual genes have absolutely no regard for the evolutionary success of the species in which they reside.

For instance, take the basic inclusive fitness equation, Hamilton's Rule. This says that if an allele at a genetic locus causes a behavior in the individual (the donor) that costs the individual a fitness loss c, and provides a fitness benefit b to another individual (the recipient), then the allele will increase in frequency provided b r > c, where r is the degree of relatedness between the donor and the recipient. The total contribution of the allele to the population is then b-c, which must be strictly positive provided that the degree of relatedness is a probability (the usual case), and b,c > 0. This is because b - c = b(1-r) + br - c > b(1-r), which is non-negative. However, there is no reason to limit the analysis to b, c, and r having particular signs. Hamilton's rule holds no matter what the sign of the cost c (which becomes a benefit when less than zero) and the benefit b (which become a cost when b is less than zero). Thus Hamilton's rule says that a selfish gene that helps its carrier by hurting other members of the population is just as capable of evolving as an altruistic gene that induces its carrier to take a fitness hit in order to confer a fitness benefit on others, despite that the former reduces mean population fitness while the latter increases mean population fitness.

For instance, suppose b and c are both negative, which occurs when the allele helps its carriers by hurting other members of the population. In that case b - c, the fitness contribution to the population, will be negative unless individuals interact almost exclusively with their genetic relatives. For instance, suppose b = -0.4, c = -0.3, and r = 1/2. Then b-c = -0.1, which is negative, but b r - c = 0.1, which is positive, so Hamilton's rule is satisfied.

The validity of inclusive fitness theory allows us to formulate the central problem of sociobiology as follows: Individual genes are utterly selfish, yet evolutionarily successful species consist of genes that predominantly cooperate with one another (this is the reason there is an "appearance of design" in multicellular organisms), and evolutionary successful social species consist predominantly in individuals who cooperate in a manner furthering their joint biological interests. By the very fact that inclusive fitness theory abstracts from interactions among loci, this theory cannot help solve the central problem of sociobiology.

The irony in the debate over group selection is that the theorists involved in the debate fight endlessly over it even though they know that there is nothing to fight about, while the onlookers, such as those tempted to read this book, believe the debate must be over something real and of momentous importance. Usually, onlookers think the debate is about whether "biological altruism" can exist, and more specifically, whether human society is based on self-interest or truly moral behavior. That is a very important question, to which Samuel Bowles and I devoted a whole book: A Cooperative Species (Princeton University Press, 2011). We conclude that our success as a species is based on the essential morality embedded in human nature. But this has nothing to do with the group selection debate as it is properly formulated. We argue in this book that the fact that humans evolved in small groups (hunter-gatherer societies) is central to the evolution of human morality, but our analytical models are equally well described in terms of inclusive fitness (applying the phenotypic gambit) and multi-level selection. Indeed, the models themselves use neither concept, but are simply sets of equations that represent plausible dynamics underlying the current constitution of the human genetic structure.
Anarus Anarus
I chased this rating because the book is really good. I use it in my research project in my University.