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Unto Others: The Evolution and Psychology of Unselfish Behavior

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Altruism has always been a problem for evolutionists. How does one explain a creature giving up something for another, sometimes its very life? Why, for example, will a monkey give a warning cry that alerts other members of the troop, but that gives away its own position? How could genes governing such behavior persist in the relentless competition for a place in the genome?

The kinds of reasoning used to explain behavior that is good for the group but perhaps not so good for the individual performing it is as old as Darwin. Until George Williams demolished whole classes of argument in his lovely 1966 book, "Adaptation and Natural Selection", it was common to invoke "group selection" as an analog to individual selection, and explain, in a vague, hand-waving sort of way, how altruistic behavior could arise by enhancing the survival of the herd, or school, or flock. And after Dawkins, both the individual and the group were banished from consideration, and the selfish gene reigned supreme.

Only one category of altruism has been taken as consonant with the unit of replication being the gene, namely "kin selection". This is the favoring of relatives: since relatives share genes, helping a gene-mate helps one's own genes, whether or not it benefits one's self. Yet much altruism in nature goes unexplained by kin selection. Think of the soldier who falls on the hand grenade so his (unrelated) buddies can live. There are many more examples from the lives of many creatures, most of whom never saw a war movie. How does one explain the clear patterns of altruistic behavior in animals at all levels of consciousness and cuddliness? Wilson, a biologist, and Sober, a philosopher, dare to think the unthinkable, or at least the unfashionable: is it possible that individuals or groups really do play a replicator role in evolution? They believe that group selection deserves another chance, but this time more rigorously specified.

I was very impressed with the first half of the book, in which they justify a group-selection model for adaptive evolution that can explain a persistent strain of altruism. What they show is that selection can take place at the level of a group of individuals in many more sorts of situations than were thought possible. (A nice bonus of this approach is that kin selection can be explained more simply using this more general context of the group.) Groups, however ephemeral, do have a role to play in selection.

The second half of the book is less convincing, as it involves psychological and philosophical arguments for "psychological altruism" in humans (that is, you not only behave unselfishly, but "want" to behave unselfishly), which, by its very nature, is hard (or very hard) to tease out in experiments, or to introspect to. However, the authors are reasonably convincing that nature would most likely not employ some Rube Goldberg-type of mental devices that depended on hedonism (pleasure-and-pain-driven behavior) to accomplish important tasks, such as child-rearing, but rather build in directly the mechanism to make a parent care to care for its child. In that way, the care of its child would be a primary motivation, rather than an intrumental one (sorry about the jargon!) on the way to getting pleasure or avoiding pain. Parents will find this convincing, as the desire to take care of one's children seems not to depend on how much we "enjoy" doing it.

This book is detailed, conscientious and well-written, but it covers a lot of ground and many of its arguments, especially in the second part, are subtle. So I recommend reading it more than once: this is contentious material. While the authors do not make anything of the political and social implications of their work, these are always waiting in the wings. Altruism, after all, is in direct opposition to selfishness. Many people see in this a political point, and a social point. Those issues are not properly a part of such a work, but do give great interest to its arguments and conclusions. And whether or not its conclusions finally survive intact, this book's arguments and approach seem exemplary and fruitful.

For more than a generation now, students of evolutionary biology have been taught that natural selection is a process that works on individuals. Where there is a conflict between the good of the individual and the good of the community, the selfish almost always prevails. There are good theoretical reasons to believe this should be so. Most of the work that has been done in the last century to turn Darwin's theory into a quantitative science seems to point in that direction. Individual selection should be fast and efficient; group selection slow and unreliable. Yet the biological world that we see seems to fly in the face of this conclusion. So much of the adaptation we see in the natural world looks like it benefits the community or the species, often at the expense of the individual. So the pure individual selectionists (99% of evolutionary biologists today) have had to concoct a series of excuses, kluges, and workarounds. There are a multitude of reasons! that what looks like a group adaptation is really an individual adaptation. Most of our community has unthinkingly adopted the view that the "selfish gene" perspective holds a key to understanding the "illusion" of group selection. Wilson has been working for 20 years to reform this situation, and to restore common sense. If it looks like a group adaptation, it probably is a group adaptation. No surprise here - except to that 99% of the academic community who has been raised to think that "group selection" is a dirty word - something like "Lamarckism" or "Creationism". Wilson's book is just the kick in the pants that the 99% of us need. It is readable, yet meticulously documented. He traces the history of our prejudice against group selection, and exposes the faulty logic in those kluges and workarounds. Group selection really is necessary to explain what we observe in nature. Then, he goes on to offer us the th! eoretical foundation we need to make group selection plausi! ble. There are mechanisms overlooked by the quantitative theorists that make group selection a far more viable process than they give it credit for. If you're a lay person, you may think "of course - what's the big deal." But if you're an academic evolutionist educated in the last 30 years, you need this book; your thinking about altruism and fitness of communities will be changed forever. All this is in the first half of the book. The second half, presumably contributed by Sober, is much less focused and scientific, more apt to dwell on definitions and philosophical distinctions. The attempt to connect the sound conclusions of the book's first half to attitudes about human cultures is both more speculative and somehow less ambitious and important than the book's first half.

I am speaking from the point of view of graduate student studying evolutionary biology.

The first section of this book is one of the clearest written accounts of group selection theory. Group level selection happens when traits are selected BECAUSE they are helpful to that group (usually a group of organisms, but some point out that individuals themselves are groups of cells). Sometimes this level of selection can be antagonistic to traits selected for at the individual level. It is well known that genetic inheritance passes through individuals. Hence the reason why existence of altruism, where an individual sacrifices its fitness for the good of group, is such a controversial topic. Of specific importance to this topic is Wilson and Sober's presentation of the Price equation, which outlines the neccessary states for group level selection to occur. Essentially, variation BETWEEN groups must be greater than variation WITHIN groups. This equation is elegant and fundamentaly irrefutable. Another important topic that Wilson and Sober present in this section is the averaging fallacy, averaging out fitness without regard to group structure. People have used this type of averaging to say that because indivduals do better when they help their group that selection is really happening at the individual level. The use of this average DOES say that those individuals are more fit but it DOES NOT say anying about levels of selection, because the averaging method ignores group structure from the outset and thus excludes the possibility that group selection could explain the results!

As the above comments reflect, Wilson and Sober have gathered together a very comprehensive set of solid theory on levels of selection. Where the book goes wrong is when Wilson and Sober start telling stories (after having promised not to do exactly that). The last two chapters of the first section deal with cultural inheritance, and such things the low cost of punishment within a group. The book degenerates severely from there in terms of scientific quality, and relies on anecdotal evidence (stories). Some of this is fun to think about, but it should be taken with many grains of salt.

Overall, Wilson and Sober have presented and communicated a set contemporary work that shows very clearly that group level selection can and probably does occur. Unfortunately, they have muddied this important contribution by trying to overreach their thesis by creating too tenuous a link between altruism based on genetic inheritence and human culture, and then using human culture as a metaphor for the evolution of altuism in other organisms. Read this book for the real and important contributions, but read it critically and do not swallow every argument whole. As John Maynard Smith said of the book, "to do so would be disaterous."

Is there empirical, biological, and evolutionary justification that mankind acts with unselfish behavior? The authors approach the subject of human altruism and the biological advantages of multilevel (group) selection vis-a-vis human egoism, hedonism, anti-functionalism, and individual functionalism from an interdisciplinary, but primarily evolutionary, approach.

The first half of the book deals with biology, genetics, and anthropology that provide the empirical grounds and logical inferences for believing that multi-level functionalism (groups and stratification) as opposed to individual-only and anti-functionalism evolved through natural selection by rewarding the fittest group selection, social norms, group adaptation, and cultural evolution, just as it rewards the fittest individual. Ergo, just as natural selection favors the fittest individuals, so it favors those individuals who cooperate in the traits of the fittest groups that survive over many generations.

The second section of the book takes the multi-level functionalism and altruism of the first half and evaluates arguments for and against it from psychological, motivational, and philosophical perspectives. While largely armchair speculation (due to lack of empirical studies confined to products of evolution rather than the actual process of evolution), the authors conclude again that natural selection again favors the fittest group, multi-level functionalism, and altruism over egoism, hedonism, selfishness, and individual selection only.

The authors' evidence and arguments are elegant, persuasive, and rigorous, but as the authors admit, much of the arguments are speculative, as no large scale studies have been done to prove or disprove their theses, because the whole subject had been largely abandoned for decades. Still, the cogent and coherent arguments make a convincing case for the rehabilitation of group-altruistic natural selection that is every bit as effectual as individual-selfish natural selection, just as Darwin presciently observed in the "Descent of Man." The conclusion is that mankind is naturally disposed by evolution to work altruistically in groups and that certain groups adapt to their environment better than others increases the significance of natural selection of the group as well as the individual. What the authors prove is that we can no longer ignore group dynamics in the evolutionary process. Altruism benefits both the individual and the group in natural selection. Highly recommended.

Sober and Wilson attempt to reduce the perception of evolution as necessarily leading to individualism and selfishness by resurrecting group selection as the means by which altruism has also evolved. Unfortunately they have produced a book which not only fails but is a laborious read.

The authors clearly accept kin-selection and the gene's-eye view yet fail to show how these methods already explain altruism and cooperation. Their attempts to show that there is altruism beyond this only leads to misrepresentations and a contrived argument.

Their definition of a group is 'a set of individuals that influence each other's fitness with respect to a certain trait' and groups are therefore 'defined on a trait by trait basis'. This sounds to me as if it is actually one of the interpretations of inclusive fitness or, perhaps, something along the lines of Dawkins' extended phenotype, which is fine but is not how people understand groups and group selection.

The main problem with group selection is the lack of stable entities that can be called groups in the first place. The authors try to leave out sexual reproduction to make it easier but this is unacceptable. One or both sexes move between groups to breed so groups are always fluctuating entities - and within a species there can be separated groups of males only or even solitary males. In humans females moved between groups either willingly or by force. Though cooperation within groups is important it is a major omission to overlook the importance of actual numbers of fertile females and female fertility in the successful expansion of groups and how this leads to male-male competition within groups and between male kin groups.

The authors do raise the relevance of empathy but other writers make a much better case of how the biological mother-infant bond has expanded to some degree to the human male and beyond parental feelings to group feelings (eg Jonathan Haidt in 'The Happiness Hypothesis').

Sober and Wilson have contrived evidence for 'pure' altruism, contrived a definition of 'group' and contrived models of 'group-selection' that are frankly impossible to relate to reality. Though well intentioned they have unfortunately merely produced a book that to read is like wading through mud and does nothing but confuse the debate. A very disappointing book.

I avoided reading this book for years because I thought it would advocate some kind of fuzzy minded group selectionism. Well, they advocate group selectionism all right, but in anything but a fuzzy minded way. They explain clearly the exact conditions under which group selection can occur and provide several examples in nature where it has undoubtably occurred. Then they go on to discuss group selectionism as it applies to human beings and the evolution by group selectionism of moral or altruistic feelings in human beings. This section of the book was EXTREMELY interesting, and brought up alot of points I had never considered before.

I wouldn't recommend this as your first book on human evolution, but if you have already read 'The Selfish Gene' and 'The Blind Watchman' by Dawkins, and 'Passions Within Reason' by Frank, and 'The Blank Slate' by Pinker, and are still excited to learn more, this book is definitely the next step. This book doesn't have the sparkling prose style of the other authors in this list, unfortunately, but it is still worth it. At least the first half of the book is.


The second half of the book was, unfortunately, a complete waste of time. It is on a completely different subject matter, the underlying psycological mechanism by which peoples feelings of altruism are produced, and they spend endless time trying to prove that people's feelings of altruism arise by one underlying psychological mechanism than by another. They are completely unsuccessful in their attempts to do so, and unfortunately it was a pretty boring journey along the way. Make that a really boring journay along the way. I struggled through the whole second half of the book because I thought that they would get over their obsession and start discussing something interesting again. But alas they never do.


I wish they had spent more time fleshing out the subject matter of the first half of the book, group selectionism, giving more examples of it in action, and discussing its implications, etc, instead of trying to tackle another subject. Either that or that they had tried to tackle a subject that was more interesting.

Nevertheless, I would still highly recommend this book. It has alot of insights in it, and it is definitely worth your time.

This book is a continuation of those books that keep moving us closer to where we came from. After decades of wandering in the jungle of postmodernism, we are finally emerging to find our roots. This book is not for the casual reader. But it is an important contribution in understanding the evolution of groupism, why humans go to war, and why belonging to the human race is not enough to bring forth altruism. Altruism evolved as a means of group consolidation of the ingroup, and genocide towards all other groups. This book should be read along with "Demonic Males" to get a good understanding of how altruism evolved.

Group selection. Suppose a population is divided into two groups: one with many altruists and one with few. Within each group altruists are (by definition) less fit than non-altruists so they will decrease in frequency. But altruists can still increase in frequency in the population as a whole, due to the altruist group expanding more rapidly. So while the altruists will die out by individual selection if the groups are completely isolated, If the groups remix or compete then the altruists can use this net-frequency advantage to win in the long run (group selection). "Altruism is maladaptive with respect to individual selection but adaptive with respect to group selection." Individual selection here means within-group selection. It may be tempting to instead define individual selection as selection for the best population-wide average fitness; but this definition (the "averaging fallacy") covers all selection, so it cannot be used to argue against group selection, only to define it away.

Example: sex ratios. Many alleged instances of group selection can be explained in terms of individual or gene selection, and it is often virtually impossible to decide between them on empirical grounds. But a case which seems to allow for empirical test is sex ratios. A female-biased sex ratio makes for faster population growth, since females can have only a limited number of offspring whereas there is no such limit for men. Thus group selection should produce a female-biased sex ratio (assuming that there are plentiful resources; conversely otherwise). Individual selection on the other hand does not want this to happen, since giving birth to a male means more future offspring. There are indeed many species with female-biased sex ratios that make good sense from a group selection point of view (e.g., parasites, who generally do not lack sustenance and are naturally divided into groups (per host)).

Kin selection. As we saw above, altruists may enjoy a temporary net-frequency gain despite declining within each group. But they need to act to take advantage of this gain (or else they will ultimately succumb to subversion from within). One way of doing so is to send out the altruists produced by the successful group as "colonists" to form new groups. Precisely this happens when the groups in questions are families. Therefore kin selection is not an alternative to group selection but rather a special case of it.

Evolutionary game theory. Game theory has been presented as an alternative to group selection which explains altruism as a strategy devised by the individual for his benefit. For example, the tit-for-tat strategy (start out altruistic and then mimic the opponent) is effective in a population with repeated prisoner's-dillema-style interactions. Clearly this is individual selection (the successful individual are those who have figured out how to win) rather than group selection (there aren't even any groups involved at all). But no: the pairs can be seen as groups. Now everything becomes group selection just as above: the selfish player is fittest within any pair and can be beaten only by the altruist pair growing faster (the entries in the payoff matrix are number of offspring as a result of this interaction). This example illustrates the general point that the relevant notion of group is that of a "trait group" (i.e., a group interacting with respect to a particular trait). This is the type of group needed for group selection as defined above. Spatial compartmentalisation, for example, is irrelevant.

Gene's eye view. This theory is also presented as an alternative to group selection but it is in fact nothing of the sort. Nothing have ever been explained or predicted by merely staring at genes. The real work when investigating any given evolutionary scenario is being done by tag-along concepts such as "vehicles." It is commonly claimed that individuals but not groups are vehicles, but whether this is so or not has nothing to do with the gene's eye view as such, and so does nothing to promote this theory as an alternative to group selection.

Some American scientists will stretch to amazing extremes in their efforts to overcome Darwin. Gould, one of the worst in this regard, set a tone of erosion of natural selection with "punk eek" or "evolution by jerks." Sober and Wilson, less original than Gould, have attempted to resurrect a long-outdated thesis with this work. As they candidly admit in their Introduction, "group selection" as an evolutionary mechanism, was declared moribund over thirty years ago. However, the remainder of the book is an attempt to revive the corpse. In their view, "altruism", although poorly defined by the authors, shifts natural selection from Darwin's original premise, to group interaction leading to optimal survival. It's a feeble effort, self-refuted on nearly every page.

Sober and Wilson offer two major themes: "Evolutionary Altruism" and "Psychological Altruism". The first part nods to the critics of group selection with some deftly selected quotes made to appear as if surrendering to the notion after all. This is a creationist ploy unexpected in a book purporting to be a serious scientific presentation. The authors attempt to temper the old view of group selection with what they term "multilevel selection theory" which they claim offers "new insights". These insights remain vague or missing altogether. Relying on human-based premises makes it immediately suspect. Using our societal standards to explain the behaviour of insects is not insight - it's misapplication of biology.

The second part, by its very title, shifts the focus to humans alone. Psychology of the remainder of the animal kingdom has so far eluded study. The authors focus on the contrast between "hedonism" and "altruism." These would appear self-explanatory, but in the hands of the authors, you are left in great doubt about their intent. Biologically, they concede these are not absolutes. Hence, we are given thoughts on "desires", "thoughts" and "beliefs" which fail to assemble into anything coherent.

Altruism bothered Darwin. In a world of individual "fitness" within an environment, survival and reproduction seemed the sole driving forces. Although "survival" has taken on a wider definition than in Darwin's day, Sober and Wilson seem to have missed the news. They even go so far as to categorise Darwin as "the first group selectionist"! Although the authors confront the reader with ponderous chapter titles ["Motives As Proximate Mechanisms"], appearing to have deep insights, a moment's reflection would have demonstrated to the authors that their thesis is untenable. No matter how much it bothered Darwin himself, altruism isn't fundamental to evolution's process. Their repeated example, the brain worm, is a prime example instead of kin selection which gives the appearance of altruism [the brain worm sibling group sacrifices one of their number that the remainder can propagate. They are 75% genetically the same].

What is most disturbing about this book is using our species to assess all life. If altruism is an evolutionary issue, why do the authors frame their concept on the human condition? They reveal their secret toward the end of the book in discussing "morality". Sober and Wilson struggle to place "morality" as an issue standing apart from natural selection, yet disclose it lies at the foundation of their presentation. There are many good questions offered for thought in this book, but no valid conclusions to adhere to. [stephen a. haines - Ottawa, Canada]