Tim Tyler: Williams, Plan and Purpose in Nature (review)

Tim Tyler: Williams, Plan and Purpose in Nature (review)

Transcript:

Hi. I'm Tim Tyler, and this is a review of this book:

Plan and Purpose in Nature: The Limits of Darwinian Evolution by George C Williams.

George Williams is an expert in Darwinian evolution, and this book is an attempt to condense his knowledge down into a format which is easily digested by those new to the field. It is very readable and entertaining book.

It covers basic issues in evolutionary biology, particularly adaptationism, the unit of selection, sexual reproduction, senescence and medicine.

Williams embraces the term "The adaptationist program" - saying he almost selected it as the title of his book. This phrase originated with Gould and Lewontin - and was intended as a term of derision. No doubt Williams is making a point by using the term, but I would have preferred that adaptation enthusiasts left it alone - making it harder for critics to point at instances of the alleged phenomenon.

The book was published in 1996 - and I thought it was interesting to see what had changed since then. Williams offered an explanation for sexual reproduction that invokes generating diversity to help adapt to novel environments. For example plants like strawberries reproduce using runners and vegetative reproduction locally, but produce sexual seeds for transmission of offspring to a diverse range of remote environments. That is not a completely unreasonable explanation, but the modern way of putting it invokes parasites. If it wasn't for the need to evade parasites, the environment would often not change fast enough to favour sexual recombination. Williams does mention parasites, but they aren't the main feature. Another area where I noticed there were some oddities were in the chapter on senescence. The disposable soma theory and the antagonistic pleiotropy theory of aging are given without being named - and Williams emphasizes how senescence of living bodies is nothing like senescence of machinery - saying:

In thinking of senescence, the analogy to the wear and tear, or corrosion, or other process that ultimately causes an artificial device to fail is utterly misleading.

I think that the relationship here is close enough to not deserve being described as "utterly misleading". Indeed these days we have new modern theories of senescence - like "reliability theory" - that apply to both machinery and bodies, that highlight the relationship between senescence in both kinds of system.

The section about group selection has stood the test of time reasonably well. Williams says that most animal groups are not functionally organized, and that most groups are "just mobs of self-seeking individuals". That's true - though these days people might take more care to mention the possibility of cooperation with kin - i.e. genetic selfishness, not individual selfishness.

The section about medicine is welcome as well. However, while Darwinian medicine sounds nice, we seem firmly embedded in the era of "drug company" medicine, with little sign of an end in sight.

At the end of the book there's a section on philosophical implications. Williams describes the product of natural selection as immoral. He says "although the biological creation process is evil it is also abysmally stupid". His example of the horrors of natural selection is infanticide by males, which he argues is natural and adaptive - though obviously infanticide by males is certainly not adaptive in many modern societies today - since it is likely to result in extended incarceration. Williams endorses Dawkins' proposed rebellion against the selfish replicators and responds to naturalistic moralities with condemnation. This is a common position these days, but human morality sits firmly in the domain of evolutionary theory, and if your theory of evolution doesn't explain it, you need to rethink it. I figure that evolutionary theory offers reasonable explanations of morality that we don't need to apologize for.

Anyway, this is still a fine book - though perhaps some learned readers might find it too simplistic.

Enjoy,

Tim Tyler: Okasha, Evolution and the Levels of Selection

Transcript:

Hi. I'm Tim Tyler, and this is a review of this book:

Evolution and the Levels of Selection by Samir Okasha

I'd previously read Samir's Very Short Introduction to the Philosophy of Science. I thought that was good - and so I had some idea of what this book would be like.

The book contains an interesting and entertaining romp through the territory of group selection. It's what I call a "firehose presentation". In other words, it's a long stream of technical material that doesn't let up. This is a good match for my own preferences in a science book. Samir goes through practically every controversy in the field, and provides insightful opinions and commentary.

The book contains discussions of the Price equation and its significance, causality, emergence, evolutionary transitions, the gene's eye view, species selection, the group selection controversy and kin selection.

I thought the book was interesting and good. However, there were also quite a few parts of it which I disagreed with - or did not like. This is a reflection of the controversial nature of the subject matter.

The book dates from 2006. Throughout most of the history of the field of group selection, many of its advocates considered it to be a super-set of kin selection - often saying things like: relatedness is only one of many ways in which altruists can form groups which are then selected. However in recent years, the quest to find things that group selection explained - and that kin selection did not - seems to have petered out, with many of the most vocal group selection advocates now proclaiming its equivalence to kin selection. Samir's book predates many of these developments - and I suspect anyone writing a book on the subject today would treat the topic rather differently.

The book discusses kin selection only rather briefly. There's a discussion about it in the chapter relating to the group selection controversy, and another one in the chapter about evolutionary transitions. Samir recognises the possibility that kin and group selection might be equivalent, and cites several sources who claim that it is, sometimes approvingly. However, most of this book makes no mention of kin selection.

These days, I think few would approach group selection this way. Kin selection has a rich and successful history, while group selection has spent most of its existence mired in confusion and controversy. Kin selection has been much better studied. So: an obvious approach to many of the topics in this book would be to just use kin selection. However, it is hard to imagine this whole book being written in the language of kin selection. A good number of the issues just seem less important from that perspective. For example, in group selection there's the issue of what counts as a group. This broadly maps onto the issue of what counts as an individual in kin selection - yet this issue seems less controversial. Group selection faces of issue of how to model parly-overlapping groups - since most group selection models feature disjoint groups. Yet the corresponding issue of partly-overlapping families in kin selection seems less contentious. It's hard to escape the impression that the need for this book is partly because group selection is so awkward, difficult to understand and poorly-studied. Since kin selection is much better studied and much more widely used, it seems as though there would be less need for a philosopher to clear up misunderstandings in the field.

Samir offers several digs at the views of Richard Dawkins in the book. He criticises the idea that evolution is based on replicators, offering Hull's comment about them "passing on their structure intact" to claim that the term "replicator" implies high fidelity copying. I think that practically everyone on both sides of this debate agrees that high fidelity copying is not necessary - and it's high fidelity information transfer that matters for cumulative adaptive evolution. No modern users of the term "replicator" in biology use the term in that way - and many of them have objected to this persistent misunderstanding. Of course it's partly Richard Dawkins' fault for assigning an ordinary english word a counter-intuitive technical meaning.

Saimr also criticises the gene's eye view on two grounds. First, he says that it ignores behavioural and environmental inheritance. That isn't true if you adopt an information-theoretic definition of the term "gene" following Williams - since then memes are a type of gene, and the gene's eye view remains valid. Saimr also says that epistasis and "modifier genes" act against the gene's eye view. This is strictly true, but some linearity in the expression of genes is really all that is required to make the gene's eye view useful. Since a linear component in the expression of genes is ubiquitous, this issue seems like a storm in a teacup to me.

Like any complex technical book, there are some mistakes. The most embarrassing one I spotted was where Samir offered an incorrect definition of inclusive fitness - including the "augmenting" but not the "stripping" component - on page 145. Samir's explanations are usually clear - but sometimes the light fades. One such problem comes with the concepts of "MLS1" and "MLS2". Samir introduces these concepts by saying that they represent different focuses of interest on page 56. However on page 59 we hear that "MLS1 and MLS2 are distinct processes" and "whether either occurs in a particular case is a matter of objective fact". At best, this sort of material is very confusing.

Overall, this is a fine book - but I was left wondering if Samir had directed his energies in an appropriate fashion. The book will probably contribute to the modern group selection revival. However that revival seems likely to be accompanied by the usual muddle and confusion that follows group selection around like a black cloud. The problems with group selection at this stage are more sociological than anything else. Yes, groups exhibit reproduction and differential reproductive success, and that affects the course of evolution, but the problem is that practically whenever group selection gets used it results in junk science, or at best, science that is inferior to that which would have been produced by using kin selection. Looking at the mess that group selection has caused in the evolutionary human sciences illustrates this point. Does the world really need more group selection? After reading Samir's book, I was still sceptical. Samir doesn't address sociological questions concerning whether the muddle associated with group selection means that it does more harm than good. Instead, he just wants to clear up the muddle. But in that case: why not use kin selection? It seems much better studied, much less confusing, and has produced much less junk science.

What I think group selection needs most is clearly-articulated reasons to use it in place of kin selection. At the moment, the "why not use kin selection?" question is challenging to answer. Maybe there are reasons - but this book doesn't really provide an answer. It doesn't even ask the question.

Enjoy,

Kin selection's domain

The Price equation pretty clearly implied that any sort of inheritance is likely to be subject to forces associated with kin selection. However, the message seems to have sunk in slowly. Kin selection was applied to DNA-mediated inheritance, but cultural inheritance was almost completely ignored by theorists.

A widespread lack of understanding of the evolutionary significance of cultural transmission could have been responsible. Also, when cultural kin selection was studied, it was often called something else. Tag-based cooperation was one popular term. Tag-based cooperation covers both genetic and cultural kin selection.

Mimicry is another area where kin selection applies. Sysyems involving mimicry may be broadly classified into organic-mimicing-organic, cultural-mimicing-cultural and cultural-mimicing-organic.

Based on the idea of similarity selection, another source of similarity in nature involves convergent evolution. Some convergent evolution involves copying from another organism - for example the parasites of swans may convergently evolve to be white. Other types involves copying from the environment, or from the laws of physics. Kin selection based on convergent evolution may happen - but it isn't easy to think of good examples. Convergent evolution is a research area for kin selection theorists.

Lastly, there's universal Darwinism to consider. Kin selection probably applies to abiotic systems as well - but again, it is not easy to think of good examples. The best cases of high-fidelity copying there are probably crystal growth and positional inheritance. Abiotic systems represent another research area.

Kin selection and mimicry

Mimicry is not normally regarded as being a form of kin selection. However, it is pretty clear that mimics engage in copying of traits - and if the individual which is being copied changes its traits, the mimic tracks the changes. Thus, mimicry can be classified as a form of copying (or inheritance) that crosses species boundaries.

There are concrete examples in nature of cooperation based on mimicry. The classic example is the cuckoo. Cuckoos mimic the eggs of their hosts - often in considerable detail - in order to elicit cooperative feeding behaviour from them. The mimicry is necessary - since the hosts employ kin recognition based on egg shell traits in order to identify their own offspring. Modifications in the host eggs are eventually copied by the cuckoo lineage - proving that genuine copying is going on.

Kin selection is clearly involved in the mimicry of the cuckoo - but most would characterize it as a case of kin selection gone wrong - since the benefits go to non-kin.

An alternative analysis looks at relatedness between the copied egg shell traits. These have managed to extend themselves beyond the host species, by copying themselves into another species - thereby gaining access to the resources of a different niche. Since many host eggs perish for each cuckoo egg, this might not seem like a good deal for the trait - but such spreading between species often turns out to be a smart move in the long run.

Of course, mimicry also happens in human culture. For example, viral videos spawn parodies which are a form of cultural mimicry. There's also mimicry between culture and organic organisms - for example, Kermit the frog mimics an organic frog.

Mimicry shows that kin selection can still apply between what seem to be non-relatives, provided they share an inherited trait. Cooperation can result because the traits themselves are kin - in the sense that one of them is copied from the other one.

Genetic similarity theory and ethnic nepotism

In the 1980s, genetic similarity theory was proposed:

We present genetic similarity theory (GST), which incorporates the kin selection theory of altruism under a more general principle. GST states that a gene ensures its own survival by acting so as to bring about the reproduction of any organism in which copies of itself are to be found. Rather than behaving altruistically only toward kin, organisms are able to detect other genetically similar organisms and to exhibit favoritism and protective behavior toward these "strangers," as well as toward their own relatives. In order to pursue this general strategy, an organism must, in effect, be able to detect copies of its genes in other organisms. We order several data sets with this theory including (a) kin recognition studies in animals raised apart, (b) assortative mating, (c) intrafamilial relations, (d) human friendship and altruism, and (e) ethnic nepotism.

This theory was frowned upon by most kin selection enthusiasts. As Richard Dawkins put it:

As to distancing myself from the National Front, that I did! The National Front was saying something like this, "kin selection provides the basis for favoring your own race as distinct from other races, as a kind of generalization of favoring your own close family as opposed to other individuals." Kin selection doesn't do that! Kin selection favors nepotism towards your own immediate close family. It does not favor a generalization of nepotism towards millions of other people who happen to be the same color as you.

Some of the points in genetic similarity theory are good. It is true that kin recognition is based on perceived relatedness - and perceived relatedness can be quite variable. It can extend beyond blood relatives. Also, one brother may appear to be more closely related to a father than another one - and this may well affect how they are treated.

However, probably the key point of controversy is that, in kin selection, relatedness drops off relatively quickly as the degree of relatedness increases - 1/2 for a brother, 1/8 for a cousin, etc - while in genetic similarity theory, genetic similarity is portrayed as dropping off more slowly - covering entire ethnic groups, for example.

This is partly an empirical question. Kin selection's metric wins quite convincingly empirically. Yes, all humans are genetically similar, but in Price's formulation, they can be expected to behave slightly spitefully towards half of the population. However it should be acknowledged that there are some effects - such as ethnic nepotism - which do seem to require an expanded version of relatedness to explain. So: what is actually going on?

Let's list some of the possible explanations:

1. Kin recognition mechanisms are mis-firing - producing the effect;
2. Humans have ingroup/outgroup detection, which is largely independent of kin recognition, and which causes this effect;
3. High-level selection (e.g. group selection or species selection) is responsible;

The first theory probably has some truth to it. The fellow from down the street could easily be your brother, while the stranger with the different racial background probably is not. Indeed, they would probably represent an unrelatedness superstimulus. However, the theory doesn't really explain why kin recognition mechanisms would produce an incorrect slope on the niceness/relatedness curve. The idea doesn't seem very plausible as a complete explanation of the phenomenon.

The second theory seems to have a lot going for it. Humans live in groups - and, in ancestral times, group members had bonds involving kinship and reciprocation, while non-group members were likely to screw you over and/or bash your head in with a rock. Not just because of kinship issues, but because they would be unlikely to face sanctions from the rest of the group for bad behaviour against group members. Members of other groups tend to look and act differently - largely because they carry different cultural symbionts. Projected forwards into modern society this "groupishness" could easily resemble ethnic nepotism and xenophobia. A similar idea probably applies to other social animals.

The third theory is probably wrong. Group selection is essentially the same thing as kin selection - and so its effects follow the same "relatedness" curve. Species selection is probably a relatively weak force. High-level selection seems likely to be too weak to explain this effect.

Genetic similarity theory is still alive and kicking in some areas - so it is important to be clear about what is wrong with it.

Kin selection is given as the primary explanation for ethnic nepotism on the Wikipedia page on the topic. However, we should not just consider kin selection based on DNA genes. Cultural kin selection is an important influence on humans. Memes often exaggerate racial division cues for their own ends. There's more to the topic than DNA genes.

---

Update 2014-11-10: Similarity selection seems like essentially the same thing. These days, I think a sympathetic reading of genetic similarity theory is the more useful one.

Neither kin selection nor group selection are much use in explaining altruism

In biology, altruism is typically defined rather like this (from Trivers, 1971):

Altruistic behaviour can be defined as behaviour that benefits another organism not closely related while apparently detrimental to the organism performing the behaviour, benefit and detriment being defined in terms of contribution to inclusive fitness.

Note the last section - about inclusive fitness. In systems whose evolution has been influenced by group selection or kin selection agents act to increase their inclusive fitness. They are not behaving atruistically.

It is possible to define altruism without measuring costs and benefits in terms of contribution to inclusive fitness - but such definitions are not of very much use. Inclusive fitness is the currency of evolution - its natural yardstick.

Reciprocity isn't much use in explaining altruism either. Though many insist on calling it "reciprocal altruism" it isn't a form of altruism at all.

Kin selection without brains

One of the common misunderstandings of kin selection is that recognising relatives requires advanced cognition:

In passing it needs to be remarked that the epistemological problems presented by a lack of linguistic support for calculating, r, coefficients of relationship, amount to a serious defect in the theory of kin selection. Fractions are of very rare occurrence in the world’s languages, appearing in Indo-European and in the archaic civilizations of the Near and Far East, but they are generally lacking among the so-called primitive peoples. Hunters and gatherers generally do not have counting systems beyond one, two and three. I refrain from comment on the even greater problem of how animals are supposed to figure out how that r [ego, first cousins] = 1/8. The failure of sociobiologists to address this problem introduces a considerable mysticism in their theory.

- Sahlins (1977) The Use and Abuse of Biology: An Anthropological Critique of Sociobiology.

Please give me an example where kin selection has been conclusively demonstrated in any species other than animals. I'm not interested in theoretical speculations or models. I'm interested in actual evidence.

- Moran, Larry (2012) Squirrels, Dawkins, and Evolution

Er, kin selection is not confined to animals. See, for example:

• Gilbert, O. M., Strassmann, J.E., & Queller, D.C. (2012) High relatedness in a social amoebae: the role of kin discrimination.

• Kuzdzal-Fick, J.A., Fox, S.A., Strassmann, J.E. & Queller, D. C. (2011) High relatedness is necessary and sufficient to maintain multicellularity in Dictyostelium.

• Gilbert, O.M., Foster, K.R., Mehdiabadi, N.J., Strassmann, J.E., Queller, D.C.(2007) High relatedness maintains multicellular cooperation in a social amoeba by controlling cheater mutants.

• Donohue, K. (2003) The influence of neighbor relatedness on multilevel selection in the Great Lakes sea rocket.

• Kelly, J.K. (1996) Kin selection in the annual plant Impatiens capensis.

• Dudley, Susan A. and File, Amanda L. (2007) Kin recognition in an annual plant.

• Rumbaugh K.P., Trivedi U., Watters C., Burton-Chellew M.N., Diggle S.P., West S.A. (2012) Kin selection, quorum sensing and virulence in pathogenic bacteria.

• Strassmann, J.E., Gilbert, O.M., Queller, D.C. (2011) Kin discrimination and cooperation in microbes.

• Chao, L., Hanley, K.A., Burch, C.L., Dahlberg, C. and Turner, P.E. (2011) Kin selection and parasite evolution: higher and lower virulence with hard and soft selection.

Equivalence naysayers

Not everyone is on board with kin selection and group selection turning out to be the same thing. Some quotes from the naysayers:

The non-equivalence of group selection and kin selection is therefore not only an important finding in itself, but also a case where the use of the Price equation leads to a claim that is not correct.

- Van Veelen M, García J, Sabelis MW, Egas M. (2012) Group selection and inclusive fitness are not equivalent; the Price equation vs. models and statistics.

Moreover, the claim that group selection is kin selection is certainly wrong.

- Nowak, Martin A., Tarnita, Corina E., and Wilson, Edward O. (2010) THE EVOLUTION OF EUSOCIALITY.

The concept of Group Selection has no useful role to play in psychology or social science.

- Pinker, Steven (2012) THE FALSE ALLURE OF GROUP SELECTION

Group selection', even in the rare cases where it is not actually wrong, is a cumbersome, time-wasting, distracting impediment to what would otherwise be a clear and straightforward understanding of what is going on in natural selection.

- Dawkins, Richard (2012) "Group Selection" Is A Cumbersome, Time-Wasting Distraction

He treated kin selection as a special case of group selection, an error which I was later to highlight in my paper on “Twelve Misunderstandings of Kin Selection” as Misunderstanding Number Two. Kin may or may not cling together in a group. Kin selection works whether they do or not.

- Dawkins, Richard (2012) The descent of Edward Wilson

the kin selection model is merely a special case of the multilevel selection theory

- Turchin, Peter (2011) Warfare and the Evolution of Social Complexity: A Multilevel-Selection Approach

Maybe, neither inclusive fitness nor evolutionary game theory can cover all mathematical intricacies of the other approach.

- Arne Traulsen (2009) MATHEMATICS OF KIN- AND GROUP-SELECTION: FORMALLY EQUIVALENT?

Ever since the publication of G. C. Williams' 1966 classic Adaptation and Natural Selection, biologists have joined with social scientists to form an altruism debunkery society. Any human or animal act that appears altruistic has been explained away as selfishness in disguise, linked ultimately to kin selection (genes help copies of themselves), or reciprocal altruism (agents help only to the extent that they can expect a positive return, including to their reputations). But in the last few years there's been a growing acceptance of the fact that "Life is a self-replicating hierarchy of levels," and natural selection operates on multiple levels simultaneously, as Bert Hölldobler and E. O. Wilson put it in their recent book, The Superorganism.

- Jonathan Haidt (2012) Contingent Superorganism

bees, ants and termites did not evolve their social behavior by group selection, but by a different mechanism known as kin selection

- Massimo Pigliucci (2011) Jonathan Haidt does it again, unfortunately

The altruism that evolves by group selection is "genuine" because it entails real self- sacrifice, while the altruism that evolves by kin selection is only "apparent" because it is just genes promoting copies of themselves in other individuals.

- David Sloane Wilson (1994) INTRODUCING GROUP SELECTION TO THE HUMAN BEHAVIORAL SCIENCES

Multilevel selection is gaining in favor among evolutionary biologists because of a recent mathematical proof that kin selection can arise only under special conditions that demonstrably do not exist, and the better fit of multilevel selection to all of the two dozen known animal cases of eusocial evolution.

- E. O. Wilson (2013) The Riddle of the Human Species

To his credit, David Sloane Wilson has subsequently reversed his position on this issue.

However, as of 2015, he still denies that kin selection and group selection cover the same turf. In Challenge To Kin Selectionists. Explain This! he claims that group selection is a more general theory.

Modern kin selection and group selection concepts have turned out to have an enormous overlap. Even if they are not identical, they are certainly very close concepts.

Similarity selection

Both Price's formalism and tag-based cooperation strongly suggest that it is trait similarity that is important to evolving cooperation - and that relatedness isn't the sole cause of similar identifying markers leading to cooperation.

This suggests that kin selection's name is dubious - and that the actual effect might be better described as a form of "similarity selection".

Reviewing the causes of similarity in biology, we find:

• Similarity based on selection
  • Similarity based on mimicry
  • Similarity based on convergent evolution
• Similarity based on chance
• Similarity based on inheritance
  • Similarity based on organic inheritance
  • Similarity based on cultural inheritance
  • Similarity based on environmental inheritance

However, looking at where cooperation arises in nature, we see it between relatives, but there's relatively little sign of cooperation based on mimicry, convergent evolution or chance resemblances.

There are some examples of mimicry leading to cooperation. For example, cuckoos generate cooperative behaviour in their hosts by employing egg mimicry. Some orchids mimic the abdomens of their pollinators, to encourage insects to attempt to have sex with their flowers. These examples are instructive: mimicry seems to be associated with producing cooperative behaviour via deception and manipulation.

Chance and convergent evolution seem to result in cooperation much less frequently. The resulting resemblances are rarely close enough.

Convergent evolution may explain the similarity between the wings of bats and birds - however, neither the bats not the birds regard each other as kin, and they don't cooperate with each other especially frequently. Much the same goes for sharks and whales - which are another case of convergent evolution - but not much cooperation arises from the resulting resemblance.

The best example I can think of for similarity which is not caused by some kind of relatedness resulting in cooperation arises in a variant of the prisoner's dilemma. Imagine a group of unrelated agents of various types playing an iterated prisoner's dilemma game with each other - under circumstances where both agents can choose whether to continue with their current partner or pick another one at random from the pool of agents without partners. No agents reproduce, but the worst agents may die. In such a game, the "nice" agents will gradually find "nice" partners, and stick with them - gaining higher payoffs than other agents. They will do this regardless of whether they are related by kinship, or not. These kind of dynamics might help to explain some forms of mutualism. However, there's a problem with describing the cooperation in such models in terms of kin selection. An alternative explanation for the resulting cooperation is reciprocity.

Whether similarity selection really adds anything to the idea of selection based on relatedness is a somewhat contentious point.

The moral of this story is that in practice most, but not all forms of similarity selection take place as a result of relatedness or kinship.

Note that we are not talking about blood relatendess here. Organisms which are memetically related may also come to cooperate - as a result of the related memes manipulating their hosts.

Group selection and kin selection: formally equivalent

These days, there's a scientific consensus about group selection and kin selection.

As Marek Kohn said in 2008:

There is widespread agreement that group selection and kin selection — the post-1960s orthodoxy that identifies shared interests with shared genes — are formally equivalent.

As Michael Wade, 2009 put it:

It is remarkable that kin selection has been widely accepted and group selection widely disparaged when, for simple genetic models, they are actually equivalent mathematically.

As Peter Richerson, 2012 put it:

I think most evolutionists now agree that kin and group selection are the same thing.

Such observations date back to Hamilton (1975). Queller (1992) is another important paper on the topic.

Modern paper titles in the area include: "Group selection and kin selection: two concepts but one process" and "Group selection and kin selection: formally equivalent approaches".

In "Social semantics: how useful has group selection been?", West, Griffin and Gardner (2009) state:

There is no theoretical or empirical example of group selection that cannot be explained with kin selection.

The theoretical equivalence of kin selection models with those of the new group selection seems to be fairly widely recognized. Wilson and Wilson (2007) seem to agree, saying:

The theories that were originally regarded as alternatives, such that one might be right and another wrong, are now seen as equivalent in the sense that they all correctly predict what evolves in the total population. They differ, however, in how they partition selection into component vectors along the way. The frameworks are largely intertranslatable and broadly overlap in the kinds of traits and population structures that they consider.

Formal models of "group selection" and "kin selection" are now widely regarded as producing the same results. Gardner and Grafen (2008) say:

group selection has already been incorporated into social evolution theory, and is found to be exactly equivalent to kin selection: the two approaches are simply different ways of describing the same evolutionary process and both lead to the prediction that individuals should maximize their inclusive fitness

Here's Gardner, West and Wild (2011):

it has long been understood that the kin selection and multilevel (group) selection approaches to social evolution are mathematically equivalent, and merely represent different partitions of the same evolutionary process (i.e. natural selection; Hamilton, 1975; Grafen, 1984, 2006a; Wade, 1985; Frank, 1986, 1995; Queller, 1992b; Rousset, 2004; Gardner et al. , 2007; Lehmann et al., 2007b; Gardner & Grafen, 2009). No model of multilevel selection has ever delivered a (correct) prediction that could not be reformulated in terms of kin selection – despite repeated claims to the contrary.

Kerr and Godfrey-Smith (2002) recommend switching between the two perspectives - saying:

we also argue that each type of model can have heuristic advantages over the other. Indeed, it can be positively useful to engage in a kind of back-and-forth switching between two different perspectives on the evolutionary role of groups. So the position we defend is a “gestalt-switching pluralism.”

Group selection enthusiast Samir Okasha endorsed equivalence in a 2010 editiorial titled Altruism researchers must cooperate writing:

Lastly, kin and multi-level selection are not alternative theories; they simply offer different takes on the question of how social behaviour evolved. Proponents of kin selection, for example, explain sterile workers in insect colonies by saying that the workers are helping the queen to reproduce, and thus boosting their own inclusive fitness. Proponents of multi-level selection argue that the workers are providing a benefit to the colony as a whole, thus making the colony fitter than other colonies. These explanations may seem different, but mathematical models show that they are in fact equivalent.

To his credit, group selection enthusiast David Sloane Wilson is now on the correct side in this debate. So is group selection enthusiast Peter Richerson and group selection enthusiast Michael Wade.

For naysayers in this area, see here.

References

• Marshall, J.A. (2011) Group selection and kin selection: formally equivalent approaches.
• Godfrey-Smith, Peter (2002) Gestalt-Switching and the Evolutionary Transitions
• West, Griffin and Gardner (2009) Social semantics: how useful has group selection been?
• West, S.A., Griffin, A.S. & Gardner, A. (2007) Social semantics: altruism, cooperation, mutualism, strong reciprocity and group selection.
• Gardner, A. and Grafen, A. (2008) Capturing the superorganism: a formal theory of group adaptation
• Lehmann, L (2007) Group selection and kin selection: Two concepts but one process
• Kohn, Marek (2008) Darwin 200: The needs of the many
• Traulsen, A. (2010) Mathematics of kin- and group-selection: formally equivalent?
• Leigh, E.G. Jr. (2010) The group selection controversy.
• Queller, David C. (2012) Two languages, one reality
• Queller, David C. (1992) Quantitative genetics, inclusive fitness, and group selection.
• Hamilton, W. D. (1975) Innate Social Aptitudes of Man: an Approach from Evolutionary Genetics
• David Sloane Wilson and E. O Wilson (2007) Rethinking the theoretical foundation of sociobiology.
• Grafen, Alan (2007) Natural Selection, Kin Selection and Group Selection.
• Johnson, Eric Michael (2012) The Good Fight.
• Wilson, David Sloan (2012) Clash of Paradigms: Why Proponents of Multilevel Selection Theory and Inclusive Fitness Theory Sometimes (But Not Always) Misunderstand Each Other
• Wilson, David Sloan (2012) Richard Dawkins, Edward O. Wilson, and the Consensus of the Many
• Wade, Michael J., Wilson, David Sloane, Goodnight, Charles, Taylor, Doug, Bar-Yam, Yaneer, de Aguiar, Marcus A. M., Stacey, Blake, Werfel, Justin, Hoelzer, Guy A., Brodie, Edmund D. III, Fields, Peter, Breden, Felix, Linksvayer, Timothy A., Fletcher, Jeffrey A., Richerson, Peter J., Bever, James D., Van Dyken, J. David and Zee, Peter (2009) Multilevel and kin selection in a connected world
• Bijma, P. and Aanen, D. K. (2009) Assortment, Hamilton's rule and multilevel selection.
• Breden, F. (1990) Partitioning of covariance as a method for studying kin selection.
• Wade, M. J. (1980) Kin selection: its components.
• Frank, S. A. (1994) Genetics of mutualism: the evolution of altruism between species
• Okasha, Samir (2010) Altruism researchers must cooperate

Group selection turns out to be kin selection

It was observed early on that most of the evidence that was cited as favouring group selection seemed to be explained rather well by kin selection.

Slime mold aggregation typically consisted of identical clones - the highest level of relatedness it is possible to get. The ant and termite families that group selection enthusiasts loved got a long list of nice quantitative predictions from kin selection theory. The evolution of chickens that supposedly counted as evidence for group selection was explained neatly by kin selection: the chickens were couped up with their close relatives.

It was argued that group selection was needed to explain cooperation between strangers who were not genetically related. However, this line of argument ignored cultural kin selection - much cooperation between strangers could be explained neatly in terms of shared memes - rather than shared genes.

There were other factors that explained cooperation too: including reciprocity, reputations, virtue signalling and the Allee effect. It seemed as though group selection was trying to take credit for the moves of other theories which were already well-established.

Was there anything group selection was needed for? Fortunately, this question now seems to have been answered...

The evolution of group selection

Group selection enthusiasts were dissatisfied with the consensus that group selection couldn't compete with individual-level selection. They counter-attacked in a number of ways, saying that:

• The insistence by critics on groups being disjoint was mistaken - groups could still be targets of selection even if they overlapped.

• Kin selection was actually a type of group selection - acting on family groups.
• Reciprocal altruism was actually a type of group selection - involving small groups of reciprocators.
• Some major evolutionary transitions showed group selection in action - today's organisms were once yesterday's groups.
• Group selection had firm mathematical foundations - in the form of the Price equation.
• Laboratory experiments on group selection showed that it was potentially a powerful force.

• There are circumstances where between-group migration was very low - for example with parasites and symbiotes that spent most of their time inside other organisms.

• Group selection explained altruism, religion, senescence, sexual recombination - etc.

They also pointed out that a number of criticisms that had been leveled at group selection were wrong:

• The idea that between-group variation would be destroyed by migration and/or within-group selection was wrong - since substantial between group variation could be observed empirically - for example in a species such as our own.

• The idea that groups are not replicators - and so aren't a valid "level of selection" - was bunk.
• The widespread 50:50 sex ratio was not an argument against group selection for that trait - group selection could easily favour a 50:50 sex ratio.

A number of these points seemed quite reasonable - however...

Tag-Based Cooperation

Price's insight about the generality of selection made slow progress into the mainstream. Universal Darwinism is still struggling for acceptance in the mainstream, four decades after Price published. Another important aspect of the generality of selection - cultural kin selection - also laboured in obscurity for a long time. Some memetics enthusiasts understood it, but few others understood them. It wasn't until the year 2000 that the idea began to gather steam in academia - under the term "Tag-Based Cooperation".

A pioneering paper in 2001 - titled "Evolution of cooperation without reciprocity" - introduced the idea. The abstract said:

Here we use computer simulations to show that cooperation can arise when agents donate to others who are sufficiently similar to themselves in some arbitrary characteristic. Such a characteristic, or `tag', can be a marking, display, or other observable trait. Tag-based donation can lead to the emergence of cooperation among agents who have only rudimentary ability to detect environmental signals and, unlike models of direct or indirect reciprocity, no memory of past encounters is required.

"Tag-Based Cooperation" was recognised as a form of cultural kin selection by Sigmund and Nowak in 2001 - as follows:

the mechanism that leads to cooperation is a form of kin selection — either classical (if traits are inherited genetically) or social (if they are inherited culturally, like a dress code).

References

• Holland, John H. (1993) The Effect of Labels (Tags) on Social Interactions.
• Eshel, Ilan, Sansone,Emilia and Shaked, Avner (1999) The emergence of kinship behavior in structured populations of unrelated individuals.
• Hales, David (2000) Cooperation without Space or Memory: Tags, Groups and the Prisoner's Dilemma.
• Riolo, R.L., Cohen, M.D., Axelrod, R. (2001) Evolution of cooperation without reciprocity.
• Sigmund, Karl and Nowak, Martin (2001) Tides of tolerance.
• Hales, David (2001) Tag Based Cooperation in Artificial Societies.
• Hales, David (2002a) Cooperation and Specialisation without Kin Selection using Tags.
• Hales, David (2002b) Smart Agents Don’t Need Kin – Evolving Specialisation and Cooperation with Tags.
• Traulsen, Arne and Schuster, Heinz Georg (2003) Minimal model for tag-based cooperation
• Axelrod, Robert, Hammond, R.A, Grafen, A (2004) Altruism via kin-selection strategies that rely on arbitrary tags with which they coevolve.
• Hales, David (2004) Tags for All! – Understanding and Engineering Tag Systems.
• Axelrod, R, Hammond, R. A. & Grafen, A. (2004) Altruism via kin-selection strategies that rely on arbitrary tags with which they coevolve.
• Hales, David and Edmonds, B. (2005a) Applying a socially-inspired technique (tags) to improve cooperation in P2P Networks.
• Hales, David (2005b) Change Your Tags Fast! – A Necessary Condition for Cooperation?
• Marcozzi, Andrea, Hales, David, Jesi, Gian Paolo, Arteconi, Stefano and Babaoglu, Ozalp (2005) Tag-Based Cooperation in Peer-to-Peer Networks with Newscast.
• Jansen VA, van Baalen M. (2006) Altruism through beard chromodynamics.
• Tanimoto J. (2007) Does a tag system effectively support emerging cooperation? A study of indirect reciprocity involving a reputation system or a simple tag system in a one-shot, multi-player game.
• Antal, Tibor, Ohtsuki, Hisashi, Wakeley, John, Taylor, Peter D. and Nowak, Martin A. (2008) Evolution of cooperation by phenotypic similarity [alt]
• Sigmund, Karl (2009) Sympathy and similarity: The evolutionary dynamics of cooperation.
• Griffiths, Nathan and Luck, Michael (2010) Changing Neighbours: Improving Tag-Based Cooperation.
• Griffiths, Nathan and Luck, Michael (2010) Norm Emergence in Tag-Based Cooperation.
• Howley, Enda and Duggan, Jim (2011) Tag-Based Cooperation in N-Player Dilemmas.
• Laird, Robert A. (2011) Green-beard effect predicts the evolution of traitorousness in the two-tag Prisoner's dilemma.
• Uitdehaag JC (2011) Bet hedging based cooperation can limit kin selection and form a basis for mutualism.

Enter George Price

One of the more significant developments in the early history of kin selection was the contribution of George Price. Price had read Hamilton's papers, and contributed a mathematical model of kin selection that proved superior to Hamilton. Hamilton had worked with the concept of shared genes. Price developed a largely-equivalent formulation based on correlations. Price's maths was better than Hamilton's - and it raised the possibility of spiteful behaviour.

Hamilton had predicted that close relatives would cooperate - since helping relatives helps copies of your genes inside them. Price pointed out that another way of helping your genes was killing off the least-closely related individuals in the population - that kin selection had an dark side: spite.

Spite was an interesting idea - but not a terribly practical one. The effort and risk involved in harming large numbers of non-relatives seemed highly unlikely to result in a payoff to the actor.

Price's equation allowed for a fairly simple derivation of Hamilton's rule. It was rather like the relationship between Newton's model and Einstein's. Price had a more general model that made almost all the same predictions as Hamilton's model - but handled a few extreme cases better.

However, Price's equation was to have other effects - besides its predictions about spite. Essentially Price modeled the effect of natural selection and persistence on the frequency of a trait in a population. It had two terms, one representing the persistence of the trait and the other representing the change caused by selection. It more-or-less ignored drift by dealing with expected values: stochastic forces would average out. Price had provided the logic of Universal Darwinism - and showed how the mathematics of selection could be applied to practically anything. Its generality reinforced the idea that blood relatedness was only one factor that could be responsible for trait correlations between generations. Many interpreted this to mean that the idea of "kin" was out - and that we needed a more general theory.

One of the things that the Price equation could be applied to was entire groups. You could split a population into groups (however you liked) and apply the Price equation to those groups. You could also apply the equation to individuals within those groups. That quickly led to a model of group selection. The change in frequency of a trait could be modeled as being divided into a component due to natural selection within groups, and a component due to natural selection between groups.

Price's maths could have led to a rapid reconciliation between kin selection and group selection proponents. However, that isn't quite what happened, as we will see.

References

• Gardner, A. (2008) The Price equation.
• Jones, James Holland (2008) Notes on the Price Equation
• Frank, Steven A. (2012) The Price Equation.
• Frank, Steven A. (1995) George Price's contributions to Evolutionary Genetics.
• Grafen, A. (2000) Developments of the Price equation and natural selection under uncertainty.

Kin selection and group selection: early rivals

The dust-up between kin selection and group selection began in 1964 with a paper by John Maynard Smith titled "Group Selection and Kin Selection". Maynard Smith wrote this after reading Hamilton's 1964 paper on the topic and delaying its publication. That caused some sour grapes - but we won't go into that story further here. The paper introduced and christened kin selection - and contrasted kin and group selection. Kin selection offered a promising explanation for altruism, while group selection seemed to have demanding prerequisites, and was probably not very significant. The paper established a pattern of thought on the topic that Maynard Smith would perpetuate throughout his career.

Hamilton wrote mostly about kin selection. Williams (1966) came out against group selection. Wilson (1975) promoted kin selection, and Dawkins (1976) was critical of group selection. Kin selection become popular, group selection became neglected and was believed to be an insignificant force.

Frank's 1998 book "Foundations of social evolution" barely mentions group selection. Alexander's "Darwinism and Human Affairs" (1979) says:

We can now say with conviction that Williams' argument against group selection was right.

Group selection was theoretically dubious, biologists seemed to rarely need to invoke it - and there wasn't any evidence for adaptations for the good of the group.

However, group selection refused to die quietly. Over the years its advocates became increasingly vocal in its defense. The "levels of selection" issue became one of the longest-running battles in evolutionary biology. The issue still seems to be alive and kicking in 2012.

On inclusive fitness

Inclusive fitness is an important concept in kin selection theory. It describes a concept which is an augmented version of "personal" fitness. Here is Hamilton's 1964 description:

Inclusive fitness may be imagined as the personal fitness which an individual actually expresses in its production of adult offspring as it becomes after it has been first 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. This quantity is then augmented by certain fractions of the quantities of harm and benefit which the individual himself causes to the fitnesses of his neighbors. The fractions in question are simply the coefficients of relationship appropriate to the neighbors whom he affects; unit for clonal individuals, one-half for sibs, one-quarter for half-sibs, one-eighth for cousins [...] and finally zero for all neighbors whose relationship can be considered negligibly small.

Inclusive fitness can be seen as an attempt to save the concept of organism-level fitness in the aftermath of the development of the idea of kin selection. The concept and its usefulness can be rather challenging to understand.

The term "inclusive fitness theory" has become a rival to the terminology used by kin selection theorists.

The term "inclusive fitness theory" was endorsed by Hamilton - the originator of the idea of kin selection. Hamilton (1975) wrote:

The usefulness of the 'inclusive fitness' approach to social behaviour [...] is that it is more general than the 'group selection', 'kin selection', or 'reciprocal altruism' approaches and so provides an overview even where regression coefficients and fitness effects are not easy to estimate or specify.

...and...

the foregoing discussion shows that kinship should be considered just one way of getting positive regression of genotype in the recipient, and that it is this positive regression that is vitally necessary for altruism. Thus the inclusive-fitness concept is more general than ‘kin selection’.

However, here we will use the term 'kin selection' in perference to 'inclusive fitness theory' - and treat them as synonyms.

Much is made by advocates of using "inclusive fitness" terminology of the virtues of identity - as opposed to "identity by descent". It is true that it is identity that matters in kin selection. However, in biology, most identity is identity by descent. The alternatives to identity by descent are identity by chance and identity by convergent evolution. However for complex biological objects, identity by chance would be a miracle, and identity by convergent evolution depends on convergent evolution being a very powerful and specific force - which it mostly isn't.

In the realm of organic evolution, genetic identity is identity by descent. If two stretches of DNA of a non-trivial size are identical, it is because they are descended from a common ancestor.

In cultural evolution, identity by convergent evolution is more likely - but still so rare as to be a big non-issue. Convergent evolution usually produces similarity - not identity.

The term "kin selection" is short and neat. It fits in neatly with the term "group selection". "Inclusive fitness theory" is a mouthful. It refers to an esoteric technical concept. It makes no mention of kinship or family.

In a nutshell, the terminology of "inclusive fitness theory" totally sucks.

An introduction

The term "kin selection" refers to a type of selection in which heritable elements increase in frequency by causing their bearers assist their relatives.

Genes coding for strategies favouring assisting relatives can spread in populations if the benefit to the relatives multiplied by a coefficient of relatedness is greater than the cost to the actor. This idea is formalised in Hamilton's equation: rB > C.

The theory of kin selection successfully models much of the apparently-altruistic behaviour in the world. It explains the love of mothers for their children. It shows that an individual-centric perspective on the natural world is incorrect - organisms are built to care about others besides themselves.

The theory of kin selection has roots dating back to Darwin, but it was first formalised by Hamilton in 1964. George Price contributed a slightly improved formalisation in the 1970s - which showed how kin selection could favour spite - as well as altruistic behavior.

Kin selection has been a tremendously productive theory. However in modern times much mud slinging against kin selection has taken place by advocates of a supposedly alternative theory of the evolution of social behaviour: group selection. Here we will attempt to set the record straight on the topic.