Exploring Altruism’s Origins: Evolution and Beyond

by Stephanie Yu (PO ‘22)

The Christian viewpoint is that we should be altruistic, moral, and loving — not only to the in-group, but even more so to those who are different from us, to those whom we may perceive as the enemy. This is because God — to whom we owe our lives — did it first for us, and now He’s asking us to do the same for each other.

COVID-19 has brought out the best and the worst in humanity. News outlets broadcast heartwarming stories of essential workers and everyday citizens venturing to unexpected, altruistic lengths to serve their communities — apartment owners waiving rent for struggling tenants, the elderly sewing and donating masks to local hospitals, and survivors donating antibody-rich blood to save patients.[1] Communities of all types have mobilized: teenagers and young adults running errands and picking up groceries for grandmas and grandpas, the employed patronizing local businesses whenever they can, and neighbors delivering precious rolls of toilet paper to each other’s doorsteps. In this time of extreme societal need, scarcity is more palpable than ever, yet intentional acts of altruism are seemingly on the rise.

At the same time, COVID-19 has exposed the nakedly selfish behavior of many who have abandoned morality and compassion in the name of survival. We may know people who are hoarding toilet paper, hand sanitizer, and meat and contributing to mass shortages. And it’s impossible to ignore the social media cacophony demanding that everything open back up, with no concern for the safety of essential workers and families in their own communities.

Perhaps these unprecedented times have us mulling over selfishness and selflessness, asking ourselves why we should or should not be altruistic. I believe that the jarring nature of the pandemic has engendered many reflections on the concept of altruism: where does it come from? How did we develop altruistic behavior, and how should that inform our actions today?

To answer these questions, we must begin by defining altruism. Interestingly enough, there are two ways to do this, depending on whether you take a psychological or biological perspective. Merriam-Webster psychologically defines altruism as an “unselfish regard for or devotion to the welfare of others” — essentially, a motivation from the “goodness of your heart.” In contrast, biology removes the element of intention and focuses exclusively on the outcome — an action is labeled altruistic only if it’s “costly to the actor and beneficial to the recipient,” with cost and benefit understood in the context of “direct fitness consequences,” or an individual’s own survival and reproduction.[2] To summarize, the colloquial understanding of altruism falls predominantly on the psychological or motivational side; however, science acknowledges altruism only from a biological or consequential standpoint.

If we trace altruism back to its inception, the biological definition is of more interest to us, as it necessitates and invites an exploration of evolution — the secular underpinning of all human processes, physical and emotional. Nevertheless, as we move forward in this piece, it’s important to be aware of the difference and avoid conflating biological/consequential altruism with psychological/motivational altruism. Let us turn to evolution.

The Evolutionary Explanation

At first glance, consequential altruism is incompatible with Darwinian evolution, and there doesn’t seem to be any place for motivational altruism either. When my goal in life is to maximize my fitness and procreate, why should I engage in selfless behavior, which only disadvantages or even threatens my livelihood? Altruism seems useless, illogical, and plain foolish in a world that operates according to “survival of the fittest.” Indeed, Darwin himself identified altruistic behavior, notably by worker ants and bees who choose to cultivate the queen’s progeny rather than engage in reproduction, as a “special difficulty” for his proposition.[3]

1963 marked the formal presentation of the inclusive fitness theory, a revolutionary explanation for the origins of altruism.[4] British evolutionary biologist William Donald Hamilton resolved Darwin’s issue by proposing a second metric for evolutionary success, indirect fitness: an organism’s ability to propagate its genes through related individuals. According to this model, opting for altruism in select situations is quite advantageous — you can still “win” by passing your genes down through your kin.

Richard Dawkins popularized this “survival of the fittest gene” theory in bestseller The Selfish Gene (1976), which introduced a thought experiment founded on inclusive fitness. He pioneered the “green-beard effect” to explain altruism. Hypothetically, altruism would ensue if a form of a gene, an allele, were to induce three expressed effects. First, the allele must produce an easily identifiable signal (such as a green beard). Next, the allele must provide the ability to discern whether others carry the same allele. Lastly, the allele must direct selective, altruistic behavior towards signal-bearers, who would also be allele-carriers.

Although the green-beard effect was initially considered an abstract possibility, recent scientific discoveries have reified this phenomenon. In the past decade, scientists have observed that within the unicellular slime mold Dictyostelium discoideum, carriers of the csa gene assemble into multicellular fruiting bodies or stalks, with 20% of the cells sacrificing their lives to ensure that the other 80% disperse and survive. Unfortunately, non-carriers of the csa gene cannot obtain membership into the stalk. Thus, we see that within a society of cells united by a common gene, a utilitarian, altruistic mechanism exists, one that maximizes prospects for survival of the gene.[5]

If we take a step back and evaluate the green-beard effect in technical terms, we can understand it as a form of kin selection, which favors altruism towards genealogical relatives — those with whom you share the same genes. Kin selection is intuitive; we are (for the most part) inclined to exhibit altruistic tendencies towards our family. But while the concentric concepts of the green-beard effect, kin selection, and inclusive fitness seem to offer thought-provoking insights into the evolutionary inception of altruistic behavior, many biologists would argue that it’s not really altruistic at all. Why? Because the benefits of the behavior nullify or outweigh the costs. Under these models, you get to see your genes live on and thus gain significantly from your “sacrifice”. You could be altruistic for the sake of altruism (motivationally altruistic) and (temporarily) harm your direct fitness, but you almost certainly gain it back later through compensations in direct and/or indirect fitness.

If we step out of this model, there are biologists who would actually go a step further and say that biological altruism, if modified to include cost to indirect fitness, does not exist at all — that we are incapable of “genetically self-destructive behavior,” as E. O. Wilson, the founder of sociobiology, would contend. In other words, altruistic behavior paves the way for compensation, ultimately promoting an average net increase in the sacrificer’s fitness. Any selflessness you pursue will more likely than not benefit you; any sacrifice you make can be explained by biological processes — processes that culminate in consequential selfishness. We rest on the “sociobiological maxim that ‘if natural selection is both sufficient and true, it is impossible for a genuinely disinterested or ‘altruistic’ behavior pattern to evolve.[6] ’”

To clarify, allow me to present evolutionary explanations for a few altruistic scenarios. Up to this point, we’ve only considered situations in which altruism is directed towards biological relatives, in which its “genuineness” can be written off by inclusive fitness theory. But what if altruism is pursued in the context of unrelated individuals? Sociobiology (the discipline that studies social behavior in terms of evolution) has quite a few answers to that, a couple of which I’ll mention here.

First, sociobiology offers a “reciprocal altruism.” Evolutionary biologist Robert Trivers (1971) developed this concept,2 which generally plays out in relational settings. Essentially, you and I engage in back-and-forth sacrifices, a quid pro quo — you can expect to receive a future boost in fitness whenever you help me and vice versa even if we’re not genetically related. This pattern of behavior is biologically selfish — we have removed motivational altruism from the equation, so it doesn’t matter whether your sacrificial actions are from disinterested kindness or ulterior motives. We only examine the outcome, which is that (on average), neither your fitness nor mine suffers from our mutually altruistic behavior.

Next, sociobiology extends the concept of “indirect reciprocity,” which explains that any sacrifice you make elevates your reputation, thus positioning you to be on the receiving end of others’ sacrifices.[7] In effect, we are more likely to assist those with “good” reputations (regardless of whether they are our kin), and in the long run, our assistance will likely result in compensation by third parties. Once again, this fails to represent consequential altruism — although our sacrifices may be costly at the moment, they may not ultimately harm our biological fitness.

But what about altruistic acts of service towards strangers? For example, let’s say that you donate anonymously to a stranger’s GoFundMe. Can that be considered biologically altruistic? Theoretically, it could be, but there’s also costly signalling theory (CST) research to suggest that we can derive biological benefits from seemingly “wasteful” sacrifices, albeit in a roundabout way. Research suggests that acts such as “stranger-directed altruism” can be advantageously assimilated into our personalities and body language, causing other strangers to be more inclined to trust us and allowing us to receive indirect recompensation. The idea is that a biological explanation can explain any instance of altruism, not that it must be the one infallible explanation.

Even if we maintain that “stranger-directed altruism” is consequentially altruistic, or consequential altruism must exist, something doesn’t quite match up. Evolutionarily speaking, in a world of every man for himself, altruism-promoting genes should be extinct or heading towards extinction. They would cripple their carriers, who would be inevitably trampled by free-loaders and unable to pass on their genes — unless, that is, natural selection acted on groups as well as individuals, as proposed by the multi-selection theory (MLS). Since we cannot ignore scarcity of resources, we must preserve a framework involving competition of some sort, and the every-group-for-itself worldview is a viable candidate.

Adding group selection to the conversation would give us a plausible explanation for biologically altruistic actions. Within such a model, groups containing only selfish individuals would overexploit resources and thus die out2. Groups with a mix of altruistic and selfish individuals, on the other hand, would win out — it would pay to have members sacrifice themselves for the greater good and fitness of the group; it would pay to preserve altruistic genes in a group’s gene pool. A phenomenon that supports group selection is cooperative breeding, the practice of fostering someone else’s offspring, which extends to those who are genetically unrelated. Biologists believe that cooperative breeding arose because of improved net reproductive success, especially in adverse conditions in which independent rearing of young would have been very difficult or unfeasible. They have recorded observations of cooperative breeding in up to 10% of bird species, meerkats and New World monkeys, including tamarins and marmosets.[8]

It is worth mentioning, however, that biologists are far from universally accepting MLS and group selection — critics of these hypotheses include Maynard Smith and Dawkins, who believe that kin selection at the individual level is a sufficient explanation. For example, Dawkins asserts that occurrences such as grey squirrels outcompeting red squirrels are simply the “side-effect of individual level selection.[9] ” A popular counterexample from the past decade is the case of “facultatively social bees,” Ceratina australensis.[10] These bees can choose to nest by themselves or in colonies; however, compared to their solitary counterparts, social nesters cut their lifetime reproductive success in half.[11] We cannot explain the phenomenon of social nesting using inclusive fitness, so perhaps group selection or MLS holds an answer. Regardless, I believe we must at least entertain the group selection hypothesis — we have not disproved it, and it bears relevance to our discussion of whether biological altruism exists.

So yes — if group selection is true, we have found credible grounds for biological altruism. If natural selection indeed divides organisms up into teams, it can make sense for me to sacrifice for my team and receive no fitness compensation, since my altruism can help my team gain the upper hand.

Does this conclude our search for consequential altruism? Not quite — for there is one more form of altruism we have not yet explored. Recall that group selection provides the rationale for biologically altruistic within-group altruism. Uncontroversially, this process relies entirely upon between-group competition and exclusion6; therefore, it is here that we finally confront the limitations of evolutionary theory — natural selection simply cannot explain out-group altruism, or altruism directed towards individuals in a competing group.

Out-group altruism would be anomalous and unprecedented; it would necessitate uncompromising altruism from both a motivational and biological lens. All except one of the scenarios we’ve examined thus far cannot be definitively reconciled with consequential altruism, and even the case of in-group biological altruism is attributable to a self-sustaining evolutionary process. But loving our opponent, sacrificing for the enemy, pursuing altruism for the outgroup — these make no sense from the evolutionary standpoint, no matter how you look at it.

Perhaps you’re wondering if this even exists. The answer is yes. Some of the most inspiring, courageous demonstrations of out-group altruism to be recorded in humanity’s history were by Holocaust rescuers in Germany. These were people who put their lives and the lives of their families on the line to save strangers, strangers who society regarded as the national enemy. Of their own volition, these rescuers allied themselves with their adversaries and embraced irreparable damage to their evolutionary fitness.

Evolution would say that such drastic, fatal measures must ensure a staggering payoff — one that would ultimately make these sacrifices consequentially selfish. But there is no conceivable benefit here. The rescuers freely chose to gain nothing and lose everything, a reality that is incomprehensible from the evolutionary perspective. With out-group altruism as exemplified by Holocaust rescuers, we’ve run into a question that evolution can’t answer.

Looking Beyond Evolution

If we decide to (even temporarily) entertain the idea that evolution is not enough, where should we search? I believe that we can find something worthwhile in motivational altruism. What exactly motivated non-Jewish Europeans to rescue Jews during the Holocaust?

A 2018 New York Times article chronicled interview responses from rescuers in a Holocaust documentary. One woman said she could not “go to bed with a bad conscience;” a man said that anti-Semitism “would be immoral;” another woman said she “did it for humanity.[12]” Each of these answers — along with many more — invoked an authoritative standard of morality. This was a standard that superseded the black-and-white rules of natural selection; it is one that has been absent from the discussion of all non-human organism behavior. But where did we get this idea of morality from, and why is altruism inherently moral?

Before we answer this question, let’s fast-forward to today. If we examine the events and emotions that have unfolded since George Floyd’s death, we see that similarly, many of our nation are. rallying behind out-group altruism in the name of morality. Indeed, many of us do not identify as Black, yet we unite under a universal moral banner that calls us to fight for justice — justice for the outgroup. To some of us, this manifests as risking our health and possibly even our lives to protest, because the injustices Floyd and so many unnamed others suffered was ethically abominable, because Black lives matter, we say. In a larger sense, a substantial part of our society champions out-group altruism — not as something that is simply “good,” but something we must do. The responsibility of those who profit from white privilege, male privilege, or socioeconomic privilege, in both pandemic and non-pandemic times, is to extend a sacrificial hand of love to the underprivileged — to those who are different from us, to the marginalized. We believe that empathy and altruism — out-group altruism — are morally right, that they create the only path to a more equitable tomorrow. Again, where does this deeply ingrained understanding of justice and morality come from?

It is my conviction that the Christian God provides an answer. The Christian perspective is that there is a God from whom all altruism, morality, justice and love originate. That He redeemed the selfishness of humanity, the out-group, through death, the greatest act of altruism. That He calls us to lay down all of our self-seeking, “survival-of-the-fittest” (group) nature to embody His universal, nondiscriminatory altruism. The Christian viewpoint is that we should be altruistic, moral, and loving — not only to the in-group, but even more so to those who are different from us, to those whom we may perceive as the enemy. This is because God — to whom we owe our lives — did it first for us, and now He’s asking us to do the same for each other.

What I present is not religious doctrine. It is, instead, an invitation to think critically about some big questions that are worth investigating — even if doing so requires challenging deeply ingrained beliefs, those that have been “factual” for as long as we can remember. I believe we can grow immensely through uncovering layer after layer of the “why:” Why is advocating and sacrificing for marginalized communities the right thing to do? Why must we be altruistic to the out-group? Why do we have such a profound belief in justice, and where do justice and morality even come from? In this time of coronavirus, turmoil and brokenness, I believe that questions like these, within a larger process of intimate reflection, will help us anchor new meaning and hope.

[2]West, S. A., Griffin, A. S., & Gardner, A. (2007). Social semantics: altruism, cooperation, mutualism, strong reciprocity and group selection. Journal of evolutionary biology, 20(2), 415–432.

[6] Schloss, J. P. (2002). ‘Love Creation’s Final Law?’: Emerging Evolutionary Accounts of Altruism. Altruism and Altruistic Love: Science, Philosophy, and Religion in Dialogue, edited by Stephen G. Post, Lynn G. Underwood, Jeffrey P. Schloss, and William B. Hurlbut, 212–42.

[7] Fehr, E., & Fischbacher, U. (2003). The nature of human altruism. Nature, 425(6960), 785–791.

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