(apologies for the gendered title – culture evolves faster than idioms do)

Selfishness versus Altruism

Language makes us think that they’re opposite ideas: The selfish behave like no one matters but them, while the altruists sacrifice themselves for the good of others. But what if selfishness and altruism weren’t opposites, but rather the same phenomena, looked at from different perspectives?

We can see a hint of this perspective shift in the early theories about the evolution of cooperation developed by W.D. Hamilton. Hamilton was concerned with this longstanding puzzle about how a behavior which benefited others, but not the actor, could ever evolve by natural selection. His solution to the puzzle involved a simple equation: rB – C > 0

Inés Dawson from Draw Curiosity made a great video explaining this equation in more detail

The basic idea is that an altruistic behavior will evolve when the benefit that the behavior gives to the recipient (multiplied by the relatedness of the actor to the receiver) is greater than the cost to the actor. The extent of relatedness (sisters > cousins, etc) is critical for this to work. Basically, the equation says that altruism evolves when creatures are able to sacrifice for the good of their family.

In just about any biological example of altruism, there is a perspective that you could take which would show underlying selfish motives. But, reciprocally, most examples of selfishness can prove to have an altruistic silver lining. 

For the Good of the Family…

Since Hamilton’s work in the 60’s, we’ve noticed lots of examples of altruism that don’t actually involve direct familial benefit. But still, even if it doesn’t involve genetically related families per say, altruism always involves helping those with whom the actor has something in common. Altruistic creatures act for the benefit of some entity or quality which both they, and those they are sacrificing for, share.

To put this a bit more poetically, altruists are actually, well, selfish, but the self they are interested in serving isn’t their personal body, but rather some sort of ‘higher self.’ Think of the honey bee, risking its life for the hive, or the martyr, dying “in the name of God.

This perspective shift works in the reverse direction too. Even the most selfish acts creatures perform, like theft, or violence, can only be accomplished by sophisticated altruistic cooperation among that creature’s cells. Skin cells, liver cells, kidney cells all work together, and regularly give up their cellular lives, so that the ‘higher self’ they are a part of can continue acting.

The takeaway here is that what we label as selfish and what we label as altruistic depends on our level of analysis. Selfish animals are made from altruistic cells, and altruistic animals, like bees, enable the selfish behavior of their hive.

This doesn’t mean that altruism doesn’t exist, or that selfishness is just in the eye of the beholder, but rather that these two ideas are intimately related. Here’s another video which explores their connection:

These videos are all part of an ongoing class called the EvoS Seminar Series. Every week a different guest lecture streams a talk, and then the class makes videos about the core concepts. We believe that the best way to learn science is to learn how to tell stories about it.

You can check out Inés Dawson’s excellent talk on the evolution of cooperation on her YouTube channel: Draw Curiosity

The EvoS Seminar Series (and this article) tries to provide a chain of content between science and narrative. We post articles, live lectures, and explainer videos which all present the same ideas from different perspectives and different levels of complexity.

Below is one more video about selfishness and altruism, this one made by artist Max Albee.

Subscribe to the EvoS Seminar Series for even more links in the Science to Narrative chain.

Image: Photoshop mosaic by Maximus Thaler; background image “The Haywain Triptych” by Hieronymus Bosch | Wikipedia

Published On: December 29, 2017

Maximus Thaler

Maximus Thaler

Maximus is a PhD candidate at Binghamton University studying cultural evolution (Wilson Lab). His work focuses on the organismality of intentional communities, in collaboration with the Federation of Egalitarian Communities. Maximus is an activist – founder of The Gleaners’ Kitchen and the Genome Collective. He’s written a cookbook and runs the EvoS Seminar Series Youtube channel.


  • Steve Davis says:

    Any discussion of altruism runs into serious problems if based on the work of Hamilton, who used what he called inclusive fitness to explain altruism. The precise nature of the definitions of biological fitness and biological altruism create a serious problem for kin selection/ inclusive fitness. The concept claims to explain the origin of altruism, but when altruism is an act that lowers fitness and fitness is the capacity to produce adult offspring, we see that altruism within kinship groups is becoming difficult to explain. It simply cannot be argued that raising offspring is altruistic. Can a mother feeding a child be seen as altruistic? Feeding a child is a demonstration of fitness and therefore cannot lower fitness. Raising a child cannot be seen as reducing the capacity to raise a child. Parental altruism is therefore impossible.
    The same problem exists for kin altruism. Assisting the survival of cousins is an inclusive fitness act, a demonstration of inclusive fitness, and therefore cannot lower inclusive fitness. Assisting the survival of cousins cannot be seen as reducing the capacity to assist their survival, so kin altruism is a fiction. Kin altruism also fails as it is a function carried out at the group level and there is no nett loss of group fitness.
    Does assisting cousins lower the individual fitness of the actor? Yes, but only in a sense so narrow as to be meaningless. Consider Hamilton’s position that the concept of inclusive fitness is “stripped of all components which can be considered as due to the individual’s social environment”. If we accept the stripping of all social components then it must be acknowledged that we are now dealing with a concept that is separated from reality. In reality, assistance to kin is reciprocated; the loss of individual fitness is regained, so there is no nett loss of fitness for the individual therefore there is no altruism.
    There is therefore no such thing as kin altruism or reciprocal altruism, as there is no loss of fitness involved in both cases. Kin selection is no more than a construct comprised of a jumble of contradictory meaningless concepts, and it certainly does not explain the origin of altruism.

  • Steve Davis says:

    By the way Maximus, I agree with your position that “To put this a bit more poetically, altruists are actually, well, selfish, but the self they are interested in serving isn’t their personal body, but rather some sort of ‘higher self.’ Think of the honey bee, risking its life for the hive, or the martyr, dying “in the name of God.”

    It’s just that I cannot see how that position can be reached from a Hamiltonian perspective. The inclusive fitness explanation for altruism revolves around shared genes; it is rigidly linked to shared genes. Once you move beyond shared genes you have abandoned inclusive fitness and all the comical positions it gives rise to; such as altruism proportionate to relatedness which logically excludes the existence of non-kin altruism, the search for the fabled but non-existent altruism gene and so on.

    You mentioned bees risking their lives for the hive. Worker bee behaviours are incessantly presented as the classic example of biological altruism, yet worker bees, being sterile (for all practical purposes) have no fitness. As they have no fitness they cannot reduce their fitness, and so cannot be altruistic.

  • Steve Davis says:

    Maximus, you have emphasised the importance of Hamilton’s Rule, even linking to an explanatory video, so it’s worth investigation.
    I think you would agree that Alan Grafen is something of an authority on the matter, a fierce defender of inclusive fitness. Here’s how Grafen describes its predictive power.

    In Natural Selection Kin Selection and Group Selection he writes; “Hamilton’s rule holds good only under certain assumptions.” Well, there’s a dent in predictive power for a start. He went on; “There are different definitions of r, and the scope of the rule depends on the definition of r (which is) employed.” Cracks are starting to appear, and it’s looking less and less like a rule.

    Grafen then discussed different scenarios, beginning with; “An animal that is helped m times, and helps n times, should experience a change of nb-nc in its number of offspring as a result. This assumes that effects add. Addition will not always be the best way for costs and benefits to combine. If the trait affects survival, then multiplication may be more appropriate.”

    “May” be more appropriate? We can pick and choose?

    So, it only holds good under certain assumptions, it doesn’t work at all under certain values of relatedness, and we can choose addition or multiplication to achieve a certain outcome. What outcome could they be trying to achieve? This is the heart of the matter. Hamilton and those who followed him had one purpose – to discredit group selection and cooperation as features of evolution. They wanted to establish individual selection as the only force in evolution, and to give respectability to selfishness as the driver of individual selection. They claimed to be motivated by a prevailing group selectionism that excluded individual selection, and if that did exist (which it may well have) then they were correct to attempt to put the record straight. But rather than have the two processes accepted as valid, they began a war on group selection. A war based on black or white is an ideological war.

    But let’s get to the nitty gritty of this. The equation is based on an assumption for which no evidence is given, the kinship requirement for altruism. This is an example of a well-known logical fallacy; that correlation is causation. It goes like this; we mostly cooperate with kin, so kinship is the determining factor obviously. Actually, no. There is a more important factor involved – proximity. We can only cooperate with those close to us. We cooperate with kin more often because it is kin that are closest to us. And as non-kin cooperation is commonplace, even among species other than humans, it is clear that proximity is the determining factor, not kinship. Proximity is essential; kinship is not.

    So why does the equation appear to back up the hypothesis? Because kinship is a near enough substitute for proximity. If the “r” component was replaced with a proximity component the results would be much the same.

  • Steve Davis says:

    The utter irrelevance of inclusive fitness can be seen in Hamilton’s declaration that “Inclusive fitness may be imagined as the personal fitness that an individual actually expresses in its production of adult offspring as it becomes after it has first been stripped and then augmented in a certain way. It is stripped of all components which can be considered as due to the individual’s social environment, leaving the fitness which he would express if not exposed to any of the harms or benefits of that environment.” Arbitrary decisions such as that have no place in science. You cannot put imaginary limits on natural selection then say “This is how evolution works.” But its lack of value is best understood by examination of the history of inclusive fitness.

    The development of the idea was prompted by Hamilton’s fascination with eugenics. He denied that link, but as his interest in eugenics sprang from his fear of overpopulation, and as inclusive fitness is concerned with the maximisation of adult offspring, a denial is unconvincing. Particularly when we consider this evidence of a prior agenda from Hamilton; “As a young man,…I slowly came to realize that there were major unsolved problems about the organisation of life that, until solved, must preclude eugenic prescriptions.” He has clearly linked the unsolved problems to eugenic solutions. On the next page this; “Another thing I wanted to understand, as a preliminary to thinking about what might be “best” for humanity as seen by my limited people-regarding side, was the source of the passions that seem inevitably involved in any mere discussion of such issues as eugenics and population control.” The “source of the passions” referred to was the origin of altruism and selfishness.

    A consistent feature of Hamilton’s work was his refusal to accept data that did not fit his conclusions, as in this; “My ideas about kin selection were at last written down and submitted to a journal. I was pretty sure they were right – that is, that they were correctly argued. If right in this way, it was clear that no amount of evidence from nature would make them wrong;…” This contempt for scientific principles can be seen most clearly in his “Geometry for the Selfish Herd”, where he rejected evidence from the field that did not fit his models, and berated Francis Galton for not extending observations of ruminant herds to cover conclusions about human behaviours!

    His other departure from scientific principles was his habit of making important assumptions, crucial assumptions, without evidence for them. This can be seen in his first published paper, The Evolution of Altruistic Behaviour (1963) in which he assumed a genetic basis to altruism, even referring to a gene for altruism. These unfounded assumptions facilitated a descent into fantasy from which Hamiltonian thought has never recovered, e.g., “To put the matter more vividly, an animal acting on this principle (altruism proportional to relatedness) would sacrifice its life if it could thereby save more than 2 brothers but not for less.” (Narrow Roads of Gene Land Vol 1, 7) Opinions such as this, that are clearly at odds with observed phenomena, have their root in his irrational belief that unless observed phenomena can be explained by a mathematical model, they are not to be taken seriously. As in this; “…this explanation (for concern for group welfare) must be treated with reserve so long as it remains unsupported by mathematical models.” (NRG Vol 1, 6) This is numerology worming its superstitious way into biology. Ever since humans first saw that mathematics could explain natural phenomena, numbers have been assumed to have magical qualities. So it’s no coincidence that because the two pioneers of gene-centrism, R A Fisher and W D Hamilton, were mathematicians before they were biologists, both developed aspects of gene-centrism based on mathematics, and consequently numbers have assumed the highest rank in the gene-centric view of evolution. (When they talk of maximising gene survival, all they are talking about is numbers.)

    The problem with this is that gene numbers are not evolution, just as a map is not a landscape, and the number of grains in a bag of sugar is not sugar. The spread of genes in a population is just one outcome of natural selection. It is not an explanation of evolution, just as ash is one outcome of fire but is not an explanation of fire. Here’s a definition, the significance of which will be immediately recognised; “Numerology is any study of the purported divine, mystical or other special relationship between a number and some coinciding observed (or perceived) events….The term numerologist is also used derogatorily for those perceived to place excess faith in numerical patterns and draw scientifically unsound inferences from them, even if those people do not practice traditional numerology.” Inclusive fitness is the West’s version of Lysenkoism; biology in the service of ideology.

  • Steve Davis says:

    The silence that has greeted my criticisms of inclusive fitness is rather disturbing. It brings to mind an answer given by Noam Chomsky during an interview in Australia a few years back. When asked for his thoughts on the role of academic institutions, he gave a lengthy reply that included this;

    “The students are expected in a good scientific university to not just repeat what they’re told; they’re expected to challenge it. In fact a lot of the new ideas and innovation come from that, so students are expected to say, you know, I don’t believe what you’re saying, here’s an argument to the contrary. That’s good, that’s what you try to encourage and if you didn’t try to encourage that, the sciences would die.”

    When obviously untrue statements such as “individuals act so as to maximise their inclusive fitness” and “altruism is proportional to relatedness” can flow in an endless stream from inclusive fitness without being challenged, then what we are seeing is the death of evolutionary biology as a science field with merit.

    My criticisms of inclusive fitness are either wrong; in which case someone should have the nerve to correct me, or they are valid, in which case someone should have the nerve to encourage me.

  • Steve Davis says:

    I think it was D S Wilson who wrote a few years back that the contribution of cooperation to evolution has been undervalued because cooperation is the ocean we swim in; it is omnipresent. This is true, but the situation is worse than this. There has been a campaign to discredit cooperation in evolution, a campaign that has culminated in Hamiltonian biology.

    There has been a strong element of individualism within British culture for over a thousand years. It began with the Norman invasion, but first found written expression in the pseudo-philosophy of Thomas Hobbes, who began a tradition of armchair philosophy based on untested assumptions, and that refuses to consider evidence that does not fit with preconceived ideas. This can be seen in Hamilton’s Geometry For The Selfish Herd, a paper littered with abuses of the scientific method. Hobbes also began the fascination with mathematical models that we see in Hamilton’s work today.

    The cult of individualism can be recognised in the artificial constraints Hamilton imposed on inclusive fitness; that is, that the concept involves “personal fitness… stripped of all components which can be considered as due to the individual’s social environment, leaving the fitness which he would express if not exposed to any of the harms or benefits of that environment.” As though society does not exist or can be ignored. As though social cooperation does not contribute to fitness. This is biology teetering on the edge of insanity.
    This tenuous hold on reality can be seen here; “First, there had come the realisation that the genome wasn’t the monolithic data bank plus executive team devoted to one project – keeping oneself alive, having babies – that I had hitherto imagined it to be. Instead, it was beginning to seem more a company boardroom, a theatre for a power struggle of egotists and factions. I was having to imagine, in parallel with others, a kind of parliament of the genes….My own conscious and seemingly indivisible self was turning out far from what I had imagined…I was an ambassador ordered abroad by some fragile coalition, a bearer of conflicting orders from the uneasy masters of a divided empire…As I write these words, even so as to be able to write them, I am pretending to a unity that, deep inside myself, I know does not exist.” (Hamilton Narrow Roads of Gene Land Vol. 1, 133-135) It might be assumed that the final sentence quoted is a rejection of individualism, but that is not the case. It is individualism taken to its logical but impossible conclusion – an individualism in which genes battle for survival. An imaginary individualism that has no connection to reality. The genome is the seat of cooperation, not conflict.
    How did this nonsense ever get traction?

    If you think my connecting inclusive fitness to a cult mentality is a step too far, consider the hysterical reaction to the Nowak Tarnita Wilson paper a few years back. Scientists with a genuine regard for their profession would not have responded with anger, they would have been delighted by the intellectual challenge. That is how science advances.

  • Steve Davis says:

    I suppose that because I have rejected the inclusive fitness explanation for the origin of altruism, I am obliged to provide an alternative view. This is not too difficult.

    The first life forms emerged through the coalescing and cooperation of pairs or groups of complex molecules. It might seem strange at first to think of molecules cooperating, because we, as conscious entities, see cooperation as intentional actions for a future benefit, but as long as a joint action has a future benefit of any kind, intent is not necessary – it is still cooperation. In the case of the first life forms, the benefits of cooperation were stability and longevity, i.e. increased survival rates, and eventually reproduction.

    Because cooperation was involved in the formation of first life, it preceded evolution, as evolution could not begin until reproduction produced variations upon which natural selection could work. Yet biologists still agonise over how cooperation could evolve in a system which is seemingly driven by ruthless competition. The lack of thought involved in evolutionary biology is staggering.

    As life forms became more complex, the forms of cooperation that assisted survival evolved with them, so that finally societies of conscious entities developed who saw that social interaction assisted individual survival. The stage was set for altruism. When individuals see themselves as part of a greater entity, transfers of material or energy to other members of the group are not seen as a loss, they are seen as internal.

    Robert Ardrey in The Social Contract explained that the healthiest societies and groups are those that facilitate both the internal conformity necessary for group stability, and the internal diversity necessary for the group to withstand environmental challenges. Which also explains why group selection and individual selection not only exist side by side, but are actually complementary. Ardrey presented such a convincing argument that Dawkins was prompted to respond with The Selfish Gene, a self-indulgent drift into scientific fantasyland.

  • Steve Davis says:

    The irrelevance of kinship to cooperation was actually conceded by Hamilton, (Narrow Roads of Gene Land Vol. 1, 51)

    “It may be, for instance, that in respect of a certain social action performed towards neighbours indiscriminately, an individual is only just breaking even in terms of inclusive fitness. If he could learn to recognise those of his neighbours who really were close relatives and could devote his beneficial actions to them alone an advantage to inclusive fitness would at once appear. Thus a mutation causing such discriminatory behaviour itself benefits inclusive fitness and would be selected. In fact, the individual may not need to perform any discrimination so sophisticated as we suggest here; a difference in generosity according to whether the situations evoking it were encountered near to, or far from his own home might occasion an advantage of a similar kind.”

    In other words, the equation on which Hamilton’s Rule is based is utterly meaningless. But more importantly, the quoted passage illustrates the incredible lack of rigour in the thinking that gave rise to inclusive fitness.

    For a start, “generosity”. Does the word even have a science based meaning? We can assume he means altruism, but precision in language is vitally important.
    Also, this short passage is littered with uncertainties. We have an “it may be”, an “only just breaking even,” an “if he could learn,” a “may not need,” and a “might occasion.” This is not science, it is not even a thought experiment; it’s idle self-indulgent speculation.

    Then there’s the assumptions.
    A “mutation causing discriminatory behaviour”? Simplistic nonsense. Keep in mind that the paper which introduced inclusive fitness to an unthinking world was titled “The Genetical Evolution of Social Behaviour”. Social behaviours are to a large extent formed by social influences; in short, by environment. To attempt to explain behaviours as purely genetic in origin by excluding social influences was not just evidence of an appalling lack of rigour; it was dishonest.

    Then to imply a mutation causing “a difference in generosity according to whether the situations evoking it were encountered near to, or far from his own home” shows how outrageous suggestions and conclusions, teetering on the edge of insanity, can flow from assumptions that are not challenged. No evidence was given for altruism proportional to distance, or for altruism proportional to relatedness. The paradox here, is that altruism proportional to distance and to relatedness do exist in fact, which might explain their enduring fascination for those with uncritical faculties. But they exist not for the reasons given by Hamilton. Decreasing levels of altruism occur with increasing distance because the opportunities for altruism decrease. Social animals conduct most social interactions close to home with familiar partners. Interactions far from home are simply less frequent because less frequented; that is where proportionality comes in. Distance and relatedness have no bearing on the level of particular altruistic actions. We do not need a mathematical formula to work this stuff out. All we need to do is think.

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