Hostname: page-component-745bb68f8f-grxwn Total loading time: 0 Render date: 2025-02-12T04:14:19.912Z Has data issue: false hasContentIssue false

The Evolution of Psychological Altruism

Published online by Cambridge University Press:  01 January 2022

Rights & Permissions [Opens in a new window]

Abstract

We argue that there are two different kinds of altruistic motivation: classical psychological altruism, which generates ultimate desires to help other organisms at least partly for those organisms’ sake, and nonclassical psychological altruism, which generates ultimate desires to help other organisms for the sake of the organism providing the help. We then argue that classical psychological altruism is adaptive if the desire to help others is intergenerationally reliable and, thus, need not be learned. Nonclassical psychological altruism is adaptive when the desire to help others is adaptively learnable. This theory opens new avenues for the interdisciplinary study of psychological altruism.

Type
Cognitive Sciences
Copyright
Copyright © The Philosophy of Science Association

1. Introduction

Some organisms behave altruistically—they reduce their own reproductive (direct) fitness (i.e., the expected number of their offspring) while increasing the reproductive (direct) fitness of another organism (West, Griffin, and Gardner Reference West, Griffin and Gardner2007; Okasha Reference Okasha and Zalta2013). Such altruistic behavior can be selected for (Sober and Wilson Reference Sober and Wilson1998; Okasha Reference Okasha2006; Gardner and West Reference Gardner and West2010): its inclusive fitness can be positive (Grafen Reference Grafen2006; Okasha Reference Okasha2016). (An organism’s inclusive fitness is defined as the sum of an organism’s direct fitness—i.e., the expected number of its own offspring—and its relatedness-weighted contribution to the direct fitness of every other organism in the population; see West et al. Reference West, Griffin and Gardner2007; West, El Mouden, and Gardner Reference West, Mouden and Gardner2011; Okasha Reference Okasha2016.)

What is controversial is whether behavior is ever motivated altruistically (i.e., by a motive to benefit others rather than by selfish interests or automatic control mechanisms). Equally controversial is whether psychological altruism—the disposition to behave based on altruistic motives—could be or has been selected for. We propose a new evolutionary framework for investigating psychological altruism and, on that basis, argue that some organisms have evolved by natural selection to be altruistically motivated.

In section 2, we make the questions we seek to answer more precise. In section 3, we articulate different kinds of motivation for altruistic behavior. In section 4, we outline circumstances under which different kinds of motivation for altruistic behavior should be expected to evolve. We bring out the importance of our findings in section 5. We conclude in section 6.

2. Determining When to Help

For obvious reasons, in general it is not adaptive for a given organism to either always help other organisms—or even all organisms from the same species—or never help anyone else. Organisms generally have to determine when to help others. Two points are important about this.

First, determining when to help others does not require that an organism deploy a concept of adaptiveness or fitness. All that is required is that the organism relies on a psychological mechanism for selecting altruistic behaviors that is based on psychological variables that correlate with the (inclusive) fitness of these behaviors. It is plausible that organisms have such a mechanism, as organisms that systematically picked out maladaptive behaviors would eventually go extinct. Furthermore, the assumption that many organisms select altruistic behaviors in ways that are by and large adaptive for them has empirical support (Houston and McNamara Reference Houston and McNamara1999; Jensen Reference Jensen, Hammerstein and Stevens2012; Chudek, Zhao, and Henrich Reference Chudek, Zhao, Henrich, Sterelny, Joyce, Calcott and Fraser2013, 436–37).Footnote 1

Second, there are two types of mechanism by which an organism can determine when to help. On one hand, the organism could rely on automatisms such as reflexes or fixed action patterns (i.e., its helping behaviors might simply by triggered in appropriate situations). On the other hand, the organism could rely on desires (i.e., representational conative states) to produce the behavior.Footnote 2 For current purposes, we restrict ourselves to the latter: the discussion to follow concerns organisms that are cognitively sophisticated—that is, organisms that rely on representational mental states when interacting with their environment (for more on this, see Schulz Reference Schulz2016, Reference Schulz2018). This is not to say that representational decision making is the biologically most widely instantiated (or otherwise most important) way of interacting with the environment. The point is just to focus on those forms of helping behavior for which the question of altruistic motivation arises: there is just no reason to classify automatism-driven behaviors as either psychologically altruistic or psychologically egoistic. That said, we return to some aspects of automatism-driven helping behavior below.

With this in mind, we seek to provide answers to the following concerns. Consider the different ways in which a representationally driven organism can determine whether to help another. Which ones deserve to be called altruistic, which egoistic, and which deserve neither label? We will answer this question in the next section. The subsequent section will examine which of the different mechanisms are likely to be selected for, and under which circumstances.

3. Psychological Altruism: Classical versus Nonclassical

A common way to distinguish altruistic from egoistic motivations to help others is based on the content of the organism’s ultimate desires (i.e., those desires that are not derived from other desires; see Sober and Wilson Reference Sober and Wilson1998; Stich, Doris, and Roedder Reference Stich, Doris, Roedder and Doris2010; Garson Reference Garson2016; Schulz Reference Schulz2016). Organisms motivated by ultimate other-involving desires (i.e., desires directed at increasing others’ well-being) are said to be psychological altruists. Organisms motivated by ultimate self-involving desires (i.e., desires directed at increasing one’s own well-being) are said to be psychological egoists. (Organisms motivated by ultimate neutral desires—i.e., desires directed neither at the self nor at another organism—are thus neither altruists nor egoists.) As we show in this section, however, this characterization is too simplistic: we must also consider how desires are produced. A desire can be produced in ways that deserve to be called either “altruistic” or “egoistic.” To make progress on the empirical and philosophical investigation of psychological altruism, therefore, the ways in which the relevant desires are produced must be considered.

A more adequate way of distinguishing between psychological altruism and egoism considers two dimensions along which motivations to help others can differ. First, as the recent discussion of this topic in the literature makes clear (Batson Reference Batson1991; Sober and Wilson Reference Sober and Wilson1998; Stich et al. Reference Stich, Doris, Roedder and Doris2010), helping behaviors might result from desires having different kinds of contents: organisms might be driven by a genuine concern for other organisms, or they might be driven by a concern for themselves only (or even by a concern that is neither other-involving nor self-involving, such as for equal distributions of resources). Second, helping behaviors might result from desires that are produced in different ways: whatever their content is, they could be produced altruistically, egoistically, or neutrally. This aspect thus concerns differences in the psychological mechanisms that generate and maintain an organism’s desires. To make this clearer, consider these differences in desire production in more detail.

A desire is egoistically produced if and only if it is generated by psychological mechanisms that were selected for increasing their bearer’s own reproductive success (i.e., its direct fitness) only, as opposed to that of other organisms (see also West et al. Reference West, Griffin and Gardner2007).Footnote 3 Two mechanisms of this kind can be distinguished. First, organisms may have an innate disposition to form desires furthering their own self-interest, which is triggered in a specific set of circumstances. That is to say, the organisms’ brains may be structured so that detecting the occurrence of certain states of affairs (e.g., low blood sugar levels) engenders the formation of a desire that leads to an increase in their own well-being (e.g., an urge to eat). Second, organisms can learn to produce desires by conditioning: if a desire is produced and maintained because its previous instances were found to be rewarding (or if the lack of such a desire led to punishment), it is egoistically produced. For example, if past tokens of the desire in question (or reasonably similar ones) improved the agent’s emotional state (or their absence worsened it) to such an extent that this past emotional reward (or punishment) alone is sufficient to cause the desire in question in the appropriate circumstances, the desire is egoistically produced (see also Goldman Reference Goldman1970; Rachlin Reference Rachlin2002; Stich Reference Stich2007; Garson Reference Garson2016).Footnote 4 In this last case, the psychological mechanism that produces the desire after conditioning is the organism’s tendency to pursue rewards and avoid punishments; for our purposes, we conservatively assume that this mechanism has been selected for only because it contributed to the organism’s own (direct) fitness. If there are cases in which the organism’s disposition to pursue rewards and avoid punishments has also been selected because it contributed to other organisms’ fitness, then those are cases of altruistically produced desires.

By contrast, a desire is altruistically produced if and only if it is generated by psychological mechanisms that were selected for because they increase the reproductive success of other organisms (i.e., their direct fitness) as well as, possibly but not necessarily, that of their bearer.Footnote 5 The main such mechanism consists of altruistically evolved innate dispositions to form desires to help others, which are triggered in a specific set of circumstances. That is to say, the organism’s brain may have evolved to be structured so that detecting the occurrence of certain states of affairs (e.g., a crying baby) engenders the formation of a desire that leads to an increase in the other’s well-being (e.g., an urge to soothe the baby). (It is worth noting that the generation of this desire may need to be mediated by an intermediate internal state, such as empathy,Footnote 6 as well as appropriate background conditions, such as bonding between agent and target and sufficient resources. This does not affect the substance of the issue here.)

Finally, a desire could be neutrally produced. This happens if and only if it is neither egoistically nor altruistically produced. A desire may be produced by mechanisms that have not been selected for at all, or it may be a by-product of other mechanisms. For example, desires might sometimes be produced similarly to how some skills are acquired. So, after repeatedly helping B, organism A may come to form a desire to help B, much like the skill of riding a bike comes after an organism has practiced this for a while. It is empirically unclear to what extent this possibility is actually instantiated (Schroeder Reference Schroeder2004; Garson Reference Garson2016); here we leave it open as a possibility.

Taking a step back, we can now put together these two aspects—the content focused and the production focused—of how helping behavior can be motivated. This will lead to at least three different motivational architectures underlying helping behavior:Footnote 7

Psychological egoism selects actions based on desires with egoistic contents.Footnote 8

Classical psychological altruism selects actions based on nonegoistically produced ultimate desires with altruistic contents.Footnote 9

Nonclassical psychological altruism selects actions based on egoistically produced ultimate desires with altruistic contents.

This division among types of helping motivations makes clear that the traditional definitions of psychological altruism and egoism are oversimplified: they make it appear that there are only two motivations for altruistic behavior, when in fact there are at least three.Footnote 10 This matters, as it can lead researchers to overlook important options that need to be considered in our theorizing about the psychological structures that might underlie altruistic behaviors. Making this clearer is the aim of the next section.

4. Psychological Altruism by Natural Selection

Are these different psychological motivations to help others merely theoretical possibilities, or is there reason to think that they have in fact evolved to drive the helping behaviors of different organisms? To answer this question, we now lay out some of the major adaptive pressures on these different ways of making helping decisions. This sort of evolutionary psychological project is associated with many uncertainties (Buller Reference Buller2005; Richardson Reference Richardson2007; Garson Reference Garson2014), so its conclusions should not be overstated. Nevertheless, evolutionary analysis is a useful launch pad for investigating how helping behavior is motivated (Schulz Reference Schulz2018).

4.1. Psychological Egoism

Psychological egoism is the most flexible but also the most cognitively demanding way to generate altruistic desires. In some cases, it may well be the best strategy—such as when helping someone makes it very likely that the organism will reciprocate that help in the future. However, in many cases it is simply too cognitively demanding and hence unfeasible. There are a large number of variables that influence whether helping another organism is adaptive (e.g., Queller Reference Queller1992; Skyrms Reference Skyrms1996, Reference Skyrms2004; Frank Reference Frank1998; Sober and Wilson Reference Sober and Wilson1998; Stevens and Hauser Reference Stevens and Hauser2004; Okasha Reference Okasha2006; West et al. Reference West, Mouden and Gardner2011; Birch and Okasha Reference Birch and Okasha2014). Consider meeting hungry strangers; whether feeding them is adaptive depends, inter alia, on how closely they are biologically related, whether they are likely to reciprocate, whether there are any social mechanisms for rewarding sharers or punishing nonsharers, and whether such mechanisms are likely to be triggered in that circumstance. Given how many unknowns there are, calculating whether to feed a hungry stranger from egoistic first principles is generally unfeasible. In many practical circumstances, organisms simply lack sufficient information to conduct the relevant instrumental reasoning with any hope of reaching reliable conclusions. For this reason, psychological egoism is unlikely to be the most important way of making helping decisions: it may be plausible in a restricted set of cases, but it is unlikely to be the primary explanation of helping behavior.

4.2. Classical Psychological Altruism

Classical psychological altruism should be expected to be an important motivator of helping behavior. The easiest way to see this is by noting that it is widely accepted that evolution can select for (innate) automatisms that produce altruistic behavior. Given this, there is no reason to rule out that it can also select for (innate) ultimate other-involving desires.

One way to see that classical psychological altruism is likely to evolve is by noting that, by definition, an organism’s inclusive fitness is a positive function of the (direct) fitness of organisms that are sufficiently closely (genetically) related to the focal organism (Queller Reference Queller1992; Taylor and Frank Reference Taylor and Frank1996; Frank Reference Frank1998; van Veelen Reference van Veelen2009; Gardner, West, and Wild Reference Gardner, West and Wild2011; Birch and Okasha Reference Birch and Okasha2014). Therefore, other things being equal, it is adaptive for many organisms to help sufficiently close kin when they are in need. This is not true for all organisms—it depends on the exact nature of the benefits and costs involved, but it is true for a number of organisms, including many mammals (Taylor and Frank Reference Taylor and Frank1996; Frank Reference Frank1998; Gardner et al. Reference Gardner, West and Wild2011).Footnote 11 Of course, for some organisms, helping their kin can be done automatically, without involving desires at all (see, e.g., Kuzdzal-Fick et al. Reference Kuzdzal-Fick, Fox, Strassmann and Queller2011; Strassman, Gilbert, and Queller Reference Strassmann, Gilbert and Queller2011). In cognitively sophisticated, representation-driven organisms, however, this circumstance creates adaptive pressures to have an innate disposition to form a desire to help kin when they are in need. As noted above, this is a form of classical psychological altruism: it is an altruistically produced desire with the content to help another organism. Moreover, there are probably other examples of classical psychological altruism as well: in some populations of organisms, similar reasoning might well support helping injured in-group members, at least in some cases (Sober and Wilson Reference Sober and Wilson1998). For present purposes, the key point to note is just that there are important sets of circumstances in which classical psychological altruism can be expected to be adaptive.

4.3. Nonclassical Psychological Altruism

Nonclassical psychological altruism is also likely to play a large role in animal psychology. Like classical psychological altruism, nonclassical psychological altruism is less cognitively demanding than egoism: it need not derive all helping behaviors from egoistic first principles. Unlike classical psychological altruism, however, nonclassical psychological altruism does not depend on the fact that the desire to help certain other organisms is cross-generationally adaptive. Instead, it can allow for the adaptiveness of this desire to be dependent on environmental factors, such as the likelihood of reciprocation. Put more positively: nonclassical altruism is adaptively favored whenever learning whether to want to help others is adaptive, even though calculating whether to do so from egoistic first principles is unfeasible.

For example, nonclassical altruism is adaptive if it is intergenerationally variable whether helping is adaptive but intragenerationally stable. For example, many social animals form alliances with select members of their group to share resources and maintain social status. In cases like this, it is not adaptive for (representationally driven) organisms to be born with an innate disposition to want to help certain other organisms: after all, whether helping is adaptive depends on the precise conditions the organism faces. However, if the conditions are such that helping is adaptive, it is also adaptive for the organism not to derive the helping behavior, every time, from egoistic ultimate desires. The existence of circumstances that fit this general pattern is well known: much of gene-culture coevolutionary theory is dedicated to investigating these circumstances (see, e.g., Fehr and Fischbacher Reference Fehr and Fischbacher2003; Boyd and Richerson Reference Boyd and Richerson2005; Henrich Reference Henrich2015). In a nutshell, given that it is sometimes adaptive for organisms to learn when to behave altruistically, there will likely be circumstances in which it is adaptive for organisms to learn to be motivated to behave altruistically (i.e., to be nonclassical altruists).

5. The Importance of Psychological Altruism for Other Disciplines

These conclusions have several important consequences for the many disciplines studying altruism. For starters, the moral value of psychological altruism vis-à-vis egoism must be reassessed. This is not only due to the fact that altruism and egoism can equally reliably lead to the same kinds of helping behaviors (as has been argued by Stich et al. Reference Stich, Doris, Roedder and Doris2010) but also because even behaviors caused by desires with other-regarding contents might be egoistically produced. This matters, as it is not obvious that there is anything morally problematic about nonclassical psychological altruism. Therefore, to see “the basic goal of morality as ‘selflessness’” is too strong (see Rachels Reference Rachels and LaFollette2000, 81; Schroder Reference Schroder and LaFollette2000, 396): moral action can countenance some egoistic influences on helping behavior.

Further, a better understanding of the differences between egoism, classical altruism, and nonclassical altruism matters, as it may help us determine how to create a more cooperative global culture. For example, knowing more about how we can learn to be altruists (e.g., through training our empathy mechanisms to recognize all human beings as in-group members) might help lessen the prevalence and severity of the kinds of racist and discriminatory behaviors that are still so common (Greene Reference Greene2013; Klimecki Reference Klimecki2015).

In cognitive neuroscience, recent work has emphasized that understanding helping behavior requires investigating its underlying neurocognitive structures (Kurzban, Burton-Chellew, and West Reference Kurzban, Burton-Chellew and West2015; Gluth and Fontanesi Reference Gluth and Fontanesi2016; Greene, Morrison, and Seligman Reference Greene, Morrison and Seligman2016; Hein et al. Reference Hein, Morishima, Leiberg, Sul and Fehr2016). We wholeheartedly agree with this. Our conclusions show that doing so requires a diachronic approach: we need to take into account not only how the organism is constituted at time t 1 (by assessing the contents of its desires) but also how it was constituted at time t 0 (by assessing how it produced the relevant desires).

Finally, our argument also complements recent work in economics. It is a still typical (although not theoretically required) assumption in much economic modeling that people are egoistically motivated. This assumption has come under much empirical and theoretical criticism, especially from work in behavioral economics and neuroeconomics (Falk, Fehr, and Fischbacher Reference Falk, Fehr and Fischbacher2003; Fehr and Camerer Reference Fehr and Camerer2007; Clavien and Chapuisat Reference Clavien and Chapuisat2016; Rand Reference Rand2016). Our current argument adds evolutionary biological considerations to the body of evidence speaking against egoistic models of human economic behavior.

6. Conclusion

There are two importantly different kinds of altruistic motivation: classical psychological altruism, which generates ultimate desires to help other organisms at least partly for those organisms’ sake, and nonclassical psychological altruism, which generates ultimate desires to help other organisms for the sake of the organism providing the help. Classical psychological altruism is adaptive if the desire to help others is intergenerationally reliable, and thus need not be learned. Nonclassical psychological altruism is adaptive when the desire to help others is adaptively learnable. Thus, both kinds of psychological altruism are likely to be selected for. This theory of the motivational structures underlying helping behaviors opens up new avenues for the productive, interdisciplinary study of psychological altruism.

Footnotes

We thank the referees and our audience at PSA 2016 for useful comments on prior drafts of this article. This material is partially based on work supported by the National Science Foundation under grant SES-1654982 to Gualtiero Piccinini.

1. In cases in which the relevant organisms are cultural learners, what determines whether a behavior will spread through the population can depend on more than its biological adaptiveness (Boyd and Richerson Reference Boyd and Richerson2005; Stich Reference Stich2016).

2. Desires, in the sense relevant here, need not be propositional representations of a state of affairs and need not be explicitly deliberated with: an immediate urge to help another organism—i.e., a representation of the kind “must help so and so”—without representing the precise state of affairs aimed at and without deliberating is sufficient (cf. Clavien Reference Clavien2011). We will remain neutral on the vexed issues of what counts as a representation and how it gets its content. Any reasonable account will do; for an opinionated defense of representational explanation within cognitive neuroscience, see Boone and Piccinini (Reference Boone and Piccinini2016).

3. Put differently, egoistically produced desires are desires produced by mechanisms whose evolution is driven just by the fact that they increase the direct reproductive fitness (i.e., the number of offspring) of the organism in question, while not increasing its indirect fitness (so that the second component of the inclusive fitness calculation is zero).

4. Note, though, that in cases in which conditioning operates by rewarding a previously occurring desire, the chain of rewards must end in a desire that is produced by some other means.

5. We may distinguish further between two kinds of altruistically produced desires. Strictly altruistically produced desires are produced by mechanisms selected for increasing solely others’ direct fitness—they increase the expected reproductive success of that other organism and do not affect or even decrease the bearer’s expected reproductive success. Broadly altruistically produced desires are produced by mechanisms selected for increasing others’ direct fitness along with their bearer’s (what West et al. [Reference West, Griffin and Gardner2007] call “mutual benefit”). This distinction will not play a role in this article.

6. De Waal (Reference de Waal2008) argues that empathy can play such a role; Klimecki et al. (Reference Klimecki, Mayer, Jusyte, Scheeff and Schönenberg2016) provide additional supporting evidence. Someone might object that empathy-driven altruistic behavior is selfishly motivated because it improves the agent’s emotional state. This objection is confused. It is well established that empathy can lead to either altruistic or selfish desires and, consequently, to either altruistic or selfish behaviors (Schulz Reference Schulz and Maibom2017). Here we are considering cases in which empathy leads to ultimate altruistic desires. In such cases, empathy deserves to be considered a component in an altruistic source of desires. Any improvement in the agent’s emotional state, which may or may not follow the desire’s satisfaction, is not the agent’s motive—it is just a by-product.

7. Clavien and Chapuisat (Reference Clavien and Chapuisat2013), Böckler, Tusche, and Singer (Reference Böckler, Tusche and Singer2016), Garson (Reference Garson2016), and Ramsey (Reference Ramsey2016) also distinguish among different types of altruism, but their theoretical frameworks are very different from ours.

8. While psychological egoism often results in egoistic behavior, it can also result in altruistic behavior.

9. Classical psychological altruism can, in turn, be divided into two kinds: a pure kind, where the ultimate desires with altruistic content are all produced altruistically (whether strictly or broadly), and an impure kind, where the ultimate desires with altruistic content are produced neutrally. We will not consider this subdivision further here.

10. Note also that one type of helping motivation that is not on the list is impersonal agency, which selects actions based on desires with neutral (i.e., nonegoistic and nonaltruistic) contents. Since this kind of impersonal helping is most relevant to highly social and cognitively sophisticated creatures subject to cultural pressures, such as human beings, and presumably it piggybacks on other forms of altruism, we set it aside here.

11. This may require complex decisions as to which kin to help (also taking into account potential future kin). See also Trivers (Reference Trivers1974) and Hausfater and Hrdy (Reference Hausfater and Hrdy1984).

References

Batson, D. 1991. The Altruism Question: Toward a Social-Psychological Answer. Hillsdale, NJ: Erlbaum.Google Scholar
Birch, J., and Okasha, S.. 2014. “Kin Selection and Its Critics.” BioScience 65 (1): 2232..10.1093/biosci/biu196CrossRefGoogle Scholar
Böckler, A., Tusche, A., and Singer, T.. 2016. “The Structure of Human Prosociality: Differentiating Altruistically Motivated, Norm Motivated, Strategically Motivated, and Self-Reported Prosocial Behavior.” Social Psychological and Personality Science 7 (6): 530–41..CrossRefGoogle Scholar
Boone, W., and Piccinini, G.. 2016. “The Cognitive Neuroscience Revolution.” Synthese 193 (5): 1509–34..CrossRefGoogle Scholar
Boyd, R., and Richerson, P.. 2005. The Origin and Evolution of Cultures. Oxford: Oxford University Press.Google Scholar
Buller, D. 2005. Adapting Minds. Cambridge, MA: MIT Press.Google Scholar
Chudek, M., Zhao, W., and Henrich, J.. 2013. “Culture-Gene Coevolution, Large Scale Cooperation, and the Shaping of Human Social Psychology.” In Cooperation and Its Evolution, ed. Sterelny, K., Joyce, R., Calcott, B., and Fraser, B., 425–57. Cambridge, MA: MIT Press.Google Scholar
Clavien, C. 2011. “Altruistic Emotional Motivation: An Argument in Favour of Psychological Altruism.” In Philosophy of Behavioral Biology: Boston Studies in Philosophy of Science, Vol. 282, ed. K. Plaisance and T. Reydon, 275–96. Dordrecht: Springer.Google Scholar
Clavien, C., and Chapuisat, M.. 2013. “Altruism across Disciplines: One Word, Multiple Meanings.” Biology and Philosophy 28 (1): 125–40..CrossRefGoogle Scholar
Clavien, C., and Chapuisat, M. 2016. “The Evolution of Utility Functions and Psychological Altruism.” Studies in History and Philosophy of Biological and Biomedical Sciences 56:2431.CrossRefGoogle ScholarPubMed
de Waal, F. B. M. 2008. “Putting the Altruism Back into Altruism: The Evolution of Empathy.” Annual Review of Psychology 59:279300.CrossRefGoogle ScholarPubMed
Falk, A., Fehr, E., and Fischbacher, U.. 2003. “On the Nature of Fair Behavior.” Economic Inquiry 41 (1): 2026..CrossRefGoogle Scholar
Fehr, E., and Camerer, C. F.. 2007. “Social Neuroeconomics: The Neural Circuitry of Social Preferences.” Trends in Cognitive Sciences 11 (10): 419–27..CrossRefGoogle ScholarPubMed
Fehr, E., and Fischbacher, U.. 2003. “The Nature of Human Altruism.” Nature 425:785–91.CrossRefGoogle ScholarPubMed
Frank, S. A. 1998. Foundations of Social Evolution. Princeton, NJ: Princeton University Press.10.1515/9780691206820CrossRefGoogle Scholar
Gardner, A., and West, S. A.. 2010. “Greenbeards.” Evolution 64 (1): 2538..CrossRefGoogle ScholarPubMed
Gardner, A., West, S. A., and Wild, G.. 2011. “The Genetical Theory of Kin Selection.” Journal of Evolutionary Biology 24 (5): 1020–43..10.1111/j.1420-9101.2011.02236.xCrossRefGoogle ScholarPubMed
Garson, J. 2014. The Biological Mind: A Philosophical Introduction. London: Routledge.CrossRefGoogle Scholar
Garson, J. 2016. “Two Types of Psychological Hedonism.” Studies in History and Philosophy of Biological and Biomedical Sciences 56:714.10.1016/j.shpsc.2015.10.011CrossRefGoogle ScholarPubMed
Gluth, S., and Fontanesi, L.. 2016. “Wiring the Altruistic Brain.” Science 351 (6277): 1028–29..CrossRefGoogle ScholarPubMed
Goldman, A. 1970. A Theory of Human Action. Princeton, NJ: Princeton University Press.Google Scholar
Grafen, A. 2006. “Optimization of Inclusive Fitness.” Journal of Theoretical Biology 238:541–63.10.1016/j.jtbi.2005.06.009CrossRefGoogle ScholarPubMed
Greene, J. D. 2013. Moral Tribes. Penguin.Google Scholar
Greene, J. D., Morrison, I., and Seligman, M. E. P.. 2016. Positive Neuroscience. Oxford: Oxford University Press.CrossRefGoogle Scholar
Hausfater, G., and Hrdy, S. B., eds. 1984. Infanticide: Comparative and Evolutionary Perspectives. Chicago: Aldine.Google Scholar
Hein, G., Morishima, Y., Leiberg, S., Sul, S., and Fehr, E.. 2016. “The Brain’s Functional Network Architecture Reveals Human Motives.” Science 351 (6277): 1074–78..10.1126/science.aac7992CrossRefGoogle ScholarPubMed
Henrich, J. 2015. The Secret of Our Success: How Culture Is Driving Human Evolution, Domesticating Our Species, and Making Us Smarter. Princeton, NJ: Princeton University Press.10.2307/j.ctvc77f0dCrossRefGoogle Scholar
Houston, A. I., and McNamara, J. M.. 1999. Models of Adaptive Behaviour: An Approach Based on State. Cambridge: Cambridge University Press.Google Scholar
Jensen, K. 2012. “Who Cares? Other-Regarding Concerns: Decisions with Feeling.” In Evolution and the Mechanisms of Decision Making, ed. Hammerstein, P. and Stevens, J. R., 299317. Cambridge, MA: MIT Press.Google Scholar
Klimecki, O. M. 2015. “The Plasticity of Social Emotions.” Social Neuroscience 10 (5): 466–73..10.1080/17470919.2015.1087427CrossRefGoogle ScholarPubMed
Klimecki, O. M., Mayer, S. V., Jusyte, A., Scheeff, J., and Schönenberg, M.. 2016. “Empathy Promotes Altruistic Behavior in Economic Interactions.” Scientific Reports 6:31961.10.1038/srep31961CrossRefGoogle ScholarPubMed
Kurzban, R., Burton-Chellew, M. N., and West, S. A.. 2015. “The Evolution of Altruism in Humans.” Annual Review of Psychology 66:575–99.CrossRefGoogle ScholarPubMed
Kuzdzal-Fick, J. A., Fox, S. A., Strassmann, J. E., and Queller, D. C.. 2011. “High Relatedness Is Necessary and Sufficient to Maintain Multicellularity in Dictyostelium.” Science 334:1548–51.CrossRefGoogle ScholarPubMed
Okasha, Samir. 2006. Evolution and the Levels of Selection. Oxford: Oxford University Press.CrossRefGoogle Scholar
Okasha, Samir 2013. “Biological Altruism.” In The Stanford Encyclopedia of Philosophy, ed. Zalta, Edward N.. Stanford, CA: Stanford University. https://plato.stanford.edu/archives/fall2013/entries/altruism-biological/.Google Scholar
Okasha, Samir 2016. “On Hamilton’s Rule and Inclusive Fitness Theory with Nonadditive Payoffs.” Philosophy of Science 83 (5): 873–83..10.1086/687871CrossRefGoogle Scholar
Queller, D. C. 1992. “Quantitative Genetics, Inclusive Fitness and Group Selection.” American Naturalist 139:540–58.10.1086/285343CrossRefGoogle Scholar
Rachels, J. 2000. “Naturalism.” In The Blackwell Guide to Ethical Theory, ed. LaFollette, H., 7491. Oxford: Blackwell.Google Scholar
Rachlin, H. 2002. “Altruism and Selfishness.” Behavioral and Brain Sciences 25:239–96.CrossRefGoogle ScholarPubMed
Ramsey, G. 2016. “Can Altruism Be Unified?Studies in History and Philosophy of Science C 56:3238.CrossRefGoogle ScholarPubMed
Rand, D. 2016. “Cooperation, Fast and Slow: Meta-analytic Evidence for a Theory of Social Heuristics and Self-Interested Deliberation.” Psychological Science 27 (9): 1192–206..CrossRefGoogle ScholarPubMed
Richardson, R. 2007. Evolutionary Psychology as Maladapted Psychology. Cambridge, MA: MIT Press.CrossRefGoogle Scholar
Schroder, W. 2000. “Continental Ethics.” In The Blackwell Guide to Ethical Theory, ed. LaFollette, H., 375–99. Oxford: Blackwell.Google Scholar
Schroeder, T. 2004. Three Faces of Desire. Oxford: Oxford University Press.CrossRefGoogle Scholar
Schulz, A. 2016. “Altruism, Egoism, or Neither: A Cognitive-Efficiency-Based Evolutionary Biological Perspective on Helping Behavior.” Studies in History and Philosophy of Biological and Biomedical Sciences 56:1523.10.1016/j.shpsc.2015.10.006CrossRefGoogle ScholarPubMed
Schulz, A. 2017. “The Evolution of Empathy.” In Routledge Handbook of the Philosophy of Empathy, ed. Maibom, H.. London: Routledge.Google Scholar
Schulz, A. 2018. Efficient Cognition: The Evolution of Representational Decision Making. Cambridge, MA: MIT Press.Google Scholar
Skyrms, B. 1996. Evolution and the Social Contract. Cambridge: Cambridge University Press.CrossRefGoogle Scholar
Skyrms, B. 2004. The Stag Hunt and the Evolution of Social Structure. Cambridge: Cambridge University Press.Google Scholar
Sober, E., and Wilson, D. S.. 1998. Unto Others: The Evolution and Psychology of Unselfish Behavior. Cambridge, MA: Harvard University Press.Google Scholar
Stevens, J. R., and Hauser, M. D.. 2004. “Why Be Nice? Psychological Constraints on the Evolution of Cooperation.” Trends in Cognitive Sciences 8 (2): 6065..CrossRefGoogle ScholarPubMed
Stich, S. 2007. “Evolution, Altruism and Cognitive Architecture: A Critique of Sober and Wilson’s Argument for Psychological Altruism.” Biology and Philosophy 22:267–81.CrossRefGoogle Scholar
Stich, S. 2016. “Why There Might Not Be an Evolutionary Explanation for Psychological Altruism.” Studies in History and Philosophy of Biological and Biomedical Sciences 56:36.CrossRefGoogle Scholar
Stich, S., Doris, J., and Roedder, E.. 2010. “Altruism.” In The Moral Psychology Handbook, ed. Doris, J. and the Moral Psychology Research Group, 147205. Oxford: Oxford University Press.CrossRefGoogle Scholar
Strassmann, J. E., Gilbert, O. M., and Queller, D. C.. 2011. “Kin Discrimination and Cooperation in Microbes.” Annual Review of Microbiology 65:349–67.10.1146/annurev.micro.112408.134109CrossRefGoogle ScholarPubMed
Taylor, P. D., and Frank, S. A.. 1996. “How to Make a Kin Selection Model.” Journal of Theoretical Biology 180:2737.10.1006/jtbi.1996.0075CrossRefGoogle ScholarPubMed
Trivers, R. 1974. “Parent-Offspring Conflict.” American Zoologist 14:247–62.10.1093/icb/14.1.249CrossRefGoogle Scholar
van Veelen, M. 2009. “Group Selection, Kin Selection, Altruism, and Cooperation: When Inclusive Fitness Is Right and When It Can Be Wrong.” Journal of Theoretical Biology 259:589600.CrossRefGoogle ScholarPubMed
West, S. A., Mouden, C. El, and Gardner, A.. 2011. “Sixteen Common Misconceptions about the Evolution of Cooperation in Humans.” Evolution and Human Behavior 32 (4): 231–62..CrossRefGoogle Scholar
West, S. A., Griffin, A. S., and Gardner, A.. 2007. “Social Semantics: Altruism, Cooperation, Mutualism, Strong Reciprocity and Group Selection.” Journal of Evolutionary Biology 20 (2): 415–32..CrossRefGoogle ScholarPubMed