The theoretical heuristic of assuming distinct alleles (or genotypes) for alternative phenotypes is the foundation of the paradigm of evolutionary explanation we call the Modern Synthesis. In modeling the evolution of sociality, the heuristic has been to set altruism and selfishness as alternative phenotypes under distinct genotypes, which has been dubbed the “phenotypic gambit.” The prevalence of the altruistic genotype that is of lower evolutionary fitness relative to the alternative genotype for non-altruistic behavior in populations is the basis of the (...) “paradox of altruism.” I show in this article that the assumption of contrasting genotypes for altruism and selfishness in our “phenotypic gambit” is inconsistent with the empirical data when viewed in the light of today’s post-Mendelian understanding of gene expression. I demonstrate that however nuanced and sophisticated the models may have become today, they are still rooted in that fundamentally problematic assumption. I then offer a genetic conception of altruism that best fits the field data. (shrink)

The puzzle of how altruism can evolve has been at the center of recent debates over Hamilton’s Rule, inclusive fitness, and kin-selection. In this paper, I use recent debates over altruism and Hamilton’s legacy as an example to illustrate a more general problem in evolutionary theory that has philosophical significance; I attempt to explain this significance and to draw a variety of conclusions about it. The problem is that specific behaviours and general concepts of organism agency and intentionality are defined (...) in terms of concepts of evolutionary “costs” and “benefits,” and these terms have determined the role that agency should play in evolutionary explanation. However, costs, benefits, and agency are not only or even best conceived through evolutionary effects in a biological context. The paper proceeds as follows: first, I explain how the issue of agency relates to the evolutionary puzzle of altruism. Next, I discuss how questions about agency have figured in recent debates over Hamilton’s legacy. In the final section, I argue that Denis Walsh’s “situated Darwinism,” which attempts to return the organism to central status in biological explanation, offers a more productive route for thinking about different forms of costs, benefits, and agency. Finally, I argue that the upshot of all this is that there may be many different, and equally valid, ways to express what organisms are doing and how they are behaving based on different currencies of cost and benefit—even if these may stand in some tension. I illustrate this through returning to the case of altruism and using examples to show that even in non-humans there can be many forms of altruism, even if they are not all biological altruism as defined in the conventional evolutionary terms. (shrink)

A hallmark of Confucian morality is its emphasis on duties to family and kin as weighty features of moral life. The virtue of ‘filiality’ or ‘filial piety’ (xiao 孝), for example, is one of the most important in the Confucian canon. This aspect of Confucianism has been of renewed interest recently. On the one hand, some have claimed that, precisely because it acknowledges the importance of kin duties, Confucianism should be seen as an ethics rooted in human nature that remains (...) a viable system of morality today. On the other hand, some have argued that the extreme emphasis on filial duties is precisely the aspect of Confucian moral philosophy that ought to be jettisoned in favor of greater impartialism; without mitigating its emphasis on filial piety, Confucianism risks irrelevance to modern concerns. In this paper, I will outline the nature of filial morality in the Confucian tradition and discuss these recent contributions to the literature. (shrink)

Kin selection and multilevel selection are alternative approaches for studying the evolution of social behaviour, the relation between which has long been a source of controversy. Many recent theorists regard the two approaches as ultimately equivalent, on the grounds that gene frequency change can be correctly expressed using either. However, this shows only that the two are formally equivalent, not that they offer equally good causal representations of the evolutionary process. This article articulates the notion of an ‘adequate causal representation’ (...) using causal graphs, and then seeks to identify circumstances under which kin and multilevel selection do and do not satisfy the test of causal adequacy. 1 Introduction2 The KS and MLS Approaches2.1 The MLS decomposition2.2 The KS decomposition3 Equivalence and Causality4 Two Problem Cases4.1 The non-social trait case4.2 Genotypic selection with meiotic drive5 Casual Adequacy: A Graphical Approach5.1 The basic idea5.2 Graphs with individual and group variables5.3 Cases where KS is causally adequate5.4 Cases where MLS is causally adequate6 Discussion6.1 Relation to previous work. (shrink)

My comments will focus on the second and third chapters of Sober’s book , which explore Darwin’s ideas about altruism, group selection and kin selection , and sex-ratio evolution . Sober makes a persuasive argument for his main claim: that Darwin was a subtler thinker on these topics than he is often taken to be. While there is much that I admire in Sober’s lucid discussion, I will focus on points of disagreement. Readers should note that this is not the (...) first time that Sober and I have disagreed on these issues .Sober begins his chapter on group selection with a brief history of the modern debate on the topic, which began in earnest in the 1960s. One of the key figures in this debate was George Williams, whose book Adaptation and Natural Selection helped convince many biologists that group selection was unlikely to be an important factor in evolution. Sober is critical of many of Williams’ arguments, though .. (shrink)

Evolutionary debunking arguments appeal to selective etiologies of human morality in an attempt to undermine moral realism. But is morality actually the product of evolution by natural selection? Although debunking arguments have attracted considerable attention in recent years, little of it has been devoted to whether the underlying evolutionary assumptions are credible. In this paper, we take a closer look at the evolutionary hypotheses put forward by two leading debunkers, namely Sharon Street and Richard Joyce. We raise a battery of (...) considerations, both empirical and theoretical, that combine to cast doubt on the plausibility of both hypotheses. We also suggest that it is unlikely that there is in the vicinity a plausible alternative hypothesis suitable for the debunker’s cause. (shrink)

Social evolution theory provides a wide array of successful evolutionary explanations for cooperative traits. However and surprisingly, a number of cases of unexplained cooperative behaviour remain. Shouldn’t they cast doubt on the relevance of the theory, or even disconfirm it? This depends on whether the theory is akin to a research programme such as adaptationism, or closer to a theory – a set of compatible, confirmable hypotheses. In order to find out, we focus on the two main tenets of social (...) evolution theory, namely reciprocity explanations and kin selection. Reciprocity-based explanations are extremely hard to confirm. This is due, first to the multiple realisability of explanatory processes, factors and strategies, despite apparent reasons to the contrary; second, to the high quantity, and limited availability of data needed to eliminate or back up such explanations. One of our target cases vividly illustrates these limitations. Moreover, kin selection, while relatively easy to disconfirm in particular cases, seems to enjoy a more limited explanatory scope than previously thought. Overall, social evolution theory turns out to be neither a research programme nor a theory, but a heterogeneous scientific entity, composed of parts that are amenable to confirmation and others barely so. (shrink)

There is a conspicuous tendency of humans to experience empathy and sympathy preferentially towards members of their own group, whereas empathetic feelings towards outgroup members or strangers are often reduced or even missing. This may culminate in a “dissociation of empathy”: a historical example are the cases of Nazi perpetrators who behaved as compassionate family men on the one hand, yet committed crimes of utter cruelty against Jews on the other. The paper aims at explaining such phenomena and at determining (...) the limits of empathy. To this purpose, it first distinguishes between two levels of empathy, namely primary or intercorporeal and extended or higher-level empathy. It then investigates the mutual interconnection of empathy and recognition, which may be regarded as a principle of extending empathy to others regardless of whether they belong to one’s own group or not. However, this principle is in conflict with ingroup conformism and outgroup biases that hamper the universal extension of empathy. Thus, a denial of recognition and exclusion of others from one’s ingroup usually results in a withdrawal or lack of extended empathy which then influences primary empathy as well. On this basis, and using the historical example of mass executions during the Holocaust, the paper investigates the mechanisms of exclusion which may lead to a withdrawal of recognition and finally to a dissociation of empathy. (shrink)

In an incendiary 2010 Nature article, M. A. Nowak, C. E. Tarnita, and E. O. Wilson present a savage critique of the best-known and most widely used framework for the study of social evolution, W. D. Hamilton’s theory of kin selection. More than a hundred biologists have since rallied to the theory’s defence, but Nowak et al. maintain that their arguments ‘stand unrefuted’. Here I consider the most contentious claim Nowak et al. defend: that Hamilton’s rule, the core explanatory principle (...) of kin selection theory, ‘almost never holds’. I first distinguish two versions of Hamilton’s rule in contemporary theory: a special version (HRS) that requires restrictive assumptions, and a general version (HRG) that does not. I then show that Nowak et al. are most charitably construed as arguing that HRS almost never holds, while HRG buys its generality at the expense of explanatory power. While their arguments against HRS are fairly uncontroversial, their arguments against HRG are more contentious, yet these have been largely overlooked in the ensuing furore. I consider the arguments for and against the explanatory value of HRG, with a view to assessing what exactly is at stake in the debate. I suggest that the debate hinges on issues concerning the causal interpretability of regression coefficients, and concerning the explanatory function Hamilton’s rule is intended to serve. (shrink)

Cooperation is rife in the microbial world, yet our best current theories of the evolution of cooperation were developed with multicellular animals in mind. Hamilton’s theory of inclusive fitness is an important case in point: applying the theory in a microbial setting is far from straightforward, as social evolution in microbes has a number of distinctive features that the theory was never intended to capture. In this article, I focus on the conceptual challenges posed by the project of extending Hamilton’s (...) theory to accommodate the effects of gene mobility. I begin by outlining the basics of the theory of inclusive fitness, emphasizing the role that the concept of relatedness is intended to play. I then provide a brief history of this concept, showing how, over the past fifty years, it has departed from the intuitive notion of genealogical kinship to encompass a range of generalized measures of genetic similarity. I proceed to argue that gene mobility forces a further revision of the concept. The reason in short is that, when the genes implicated in producing social behaviour are mobile, we cannot talk of an organism’s genotype simpliciter; we can talk only of an organism’s genotype at a particular stage in its life cycle. We must therefore ask: with respect to which stage(s) in the life cycle should relatedness be evaluated? For instance: is it genetic similarity at the time of social interaction that matters to the evolution of social behaviour, or is it genetic similarity at the time of reproduction? I argue that, strictly speaking, it is neither of these: what really matters to the evolution of social behaviour is diachronic genetic similarity between the producers of fitness benefits at the time they produce them and the recipients of those benefits at the end of their life-cycle. I close by discussing the implications of this result. The main payoff is that it makes room for a possible new mechanism for the evolution of altruism in microbes that does not require correlated interaction among bearers of the genes for altruism. The importance of this mechanism in nature remains an open empirical question. (shrink)

The ideological underpinning that guides our interaction with non-human animals needs revision. The traditional outlook, according to which humans have a higher moral status vis-à-vis non-human animals, is now otiose. If these claims are to be justified, what ideological framework would serve this end? What are the moral implications of endorsing the view that humans possess no higher moral status than non-human animals? This work takes as foundation Charles Darwins theory of evolution, which affirms that humans emerged from the long (...) chain of evolutionary history, where non-human animals have been the carriers of the genes that shaped humans. A revisit to the discourse on the moral implication of humans cruelty to their ancestors and neighbours becomes pertinent. This essay goes against the mainstream and dominant perspective that non-human animals exist to serve human ends and as such can be treated with disdain. The thesis of this paper goes beyond Peter Singers submission that sentience is the basis for conferring moral worth on non-human animals. It affirms that in addition to sentience, good neighbourliness is a factor in determining the moral worth of non-human animals. It submits that cruelty to reared and domesticated animals may produce violent and wild species of these animals kind in a future evolution, thereby endangering the lives of future human generation, through negative alteration of genes. In the end, this paper proposes the principle of biological altruism as a suitable norm for determining the moral worth of non-human animals. (shrink)

Altruisme pada makhluk sosial, selain manusia, selalu menjadi topik yang menarik. Satu dari contoh tipikalnya adalah semut pekerja – semut mandul yang bekerja tanpa kenal lelah demi ratu mereka. Bisa dibilang, evaluasi menurut asas biaya-manfaat dalam sistem koloni semut sedang berlangsung di tingkat kolektif dan bukan tingkat individu. Perlu dicatat bahwa hal ini secara biologis lebih merupakan batasan yang “keras” daripada pilihan yang bersifat “pribadi”. Perilaku altruistik yang dipilih secara alami pada hewan bermanfaat bagi kelangsungan hidup dan reproduksi spesies yang (...) membawa biaya individu (pengorbanan). Secara keseluruhan, perilaku seperti ini disebut altruisme biologis [1]. Niat dan perilaku manusia sering kali merupakan gabungan yang jauh lebih rumit dari naluri biologis dan kognisi tingkat lanjut. Perilaku altruistik pada manusia yang banyak ditemukan telah dikaitkan secara positif dengan kebahagiaan, kesehatan mental yang lebih baik, dan kesehatan fisik yang lebih baik, yang disarankan selaras dengan prinsip biologis dan evolusioner alam [2]. Dalam sebuah studi di tahun 2019 yang diterbitkan dalam Proceedings of the National Academy of Sciences (PNAS), peneliti menemukan bahwa perilaku altruistik dapat menghilangkan rasa sakit fisik dengan mengurangi aktivitas otak di dorsal anterior cingulate cortex dan bilateral insula dalam merespon rangsangan yang menyakitkan [3]. (shrink)

In this paper I first offer a systematic outline of a series of conceptual novelties in the life-sciences that have favoured, over the last three decades, the emergence of a more social view of biology. I focus in particular on three areas of investigation: (1) technical changes in evolutionary literature that have provoked a rethinking of the possibility of altruism, morality and prosocial behaviours in evolution; (2) changes in neuroscience, from an understanding of the brain as an isolated data processor (...) to the ultrasocial and multiply connected social brain of contemporary neuroscience; and (3) changes in molecular biology, from the view of the gene as an autonomous master of development to the ‘reactive genome’ of the new emerging field of molecular epigenetics. In the second section I reflect on the possible implications for the social sciences of this novel biosocial terrain and argue that the postgenomic language of extended epigenetic inheritance and blurring of the nature/nurture boundaries will be as provocative for neo-Darwinism as it is for the social sciences as we have known them. Signs of a new biosocial language are emerging in several social-science disciplines and this may represent an exciting theoretical novelty for twenty-first social theory. (shrink)

This is the introductory essay to the Italian translation of Matt Ridley's "The origins of virtue", surveying the game-theoretic and evolutionary approaches to the emergence and evolution of cooperation and altruism.

We sometimes joke about the world conspiring against us. For instance, I might suggest that it will rain today because I’m without my umbrella or that my football team will play poorly this weekend because I’m attending the game. But of course the weather is not influenced by my umbrella, and football teams do not perform any differently when I’m present. Any perceived correlations here are most likely illusory or, if real, mere accidents. In other words, the probability of a (...) fine day and the probability of my team playing well are independent of my actions. (shrink)