Why can't we all just get along?

After the 2010 Nowak, Tarnita, & Wilson affair, Samir Okasha weighed in with a comment in Nature, effectively asking: why can't we all just get along?

The paper was titled: Altruism researchers must cooperate. It is indeed ironic that the science of cooperation has led to so much scientific acrimony.

In the paper, Samir embraces equivalence, writing:

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

He says the persistent rival camps are a puzzle since:

The existence of equivalent formulations of a theory, or of alternative modelling approaches, does not usually lead to rival camps in science. The Lagrangian and Hamiltonian formulations of classical mechanics, for example, or the wave and matrix formulations of quantum mechanics, tend to be useful for tackling different problems, and physicists switch freely between them.

His explanation:

History shows that, despite its enormous empirical success, evolutionary biology is peculiarly susceptible to controversy and infighting. This is particularly true of social evolution theory, in part because of its potential applications to human behaviour.

I think this is essentially correct. People fight over application of evolutionary theory to humans for many reasons, but among them are: "it's complicated", and "it's important to get it right".

Group selection doesn't score well in the latter category. It isn't so much that it is wrong, it's more that it has led to decades of muddle, confusion and poor-quality science. If kin selection and group selection are feeling different parts of the same elephant, kin selection has hold of the trunk, while group selection is groping the left thigh.

It is certainly frustrating to have all the biology papers using kin selection, while all the humanities papers seem to use group selection.

A big part of the problem is misunderstandings surrounding cultural evolution. Cultural evolution has historically lagged by decades behind conventional evolutionary theory. There was a revolution in the 1970s in which group selection fell out of fashion, and kin selection became much more popular. A parsimonious explanation of the penchant for group selection in the social sciences is that the field of cultural evolution has yet to go through this transition. The significance of relatedness between memes and memeplexes has yet to be fully appreciated.

Nowak and Wilson are at it again!

You might think Nowak and Wilson would have had their fingers burned in the reaction to their 2010 Nature paper.

However, it seems as though they are at it again. They have a PNAS paper titled: Limitations of inclusive fitness by Benjamin Allen, Martin A. Nowak, Edward O. Wilson.

They deny that evolution can be usefully viewed as an optimization process, saying:

Thus, evolution does not, in general, lead to the maximization of inclusive fitness or any other quantity.

That's complete nonsense: Allen, Nowak, and Wilson don't know what they are talking about. Optimization models are perfectly general - and can model any dynamical system.

The guts of their paper is a criticism of the application of Price's equation to inclusive fitness.

Their main beef is linearity. They are concerned that the 'adding' and 'subtracting' that goes on in the definition of inclusive fitness limits its generality. They write:

Inclusive fitness assumes that personal fitness is the sum of additive components caused by individual actions. This assumption does not hold for the majority of evolutionary processes or scenarios.

However, this linearity doesn't really cause problems - since you can approximate non-linear curves using a series of line segments.

Inclusive fitness is mostly concerned with the issue of whether some helping behaviour will evolve. It calculates whether the net selective effect of a behaviour on the frequency of a gene responsible for it (some of which may be copies in relatives of the actor) is positive or negative (relative to a set of alternatives) in the current environment. The math of inclusive fitness tells you whether the gene is selectively favoured. That's not quite the same thing as whether the behaviour will evolve (maybe it will drift into extinction) - but it's a good start. Nobody ever thought that the behaviours and genes involved had to not interact. Nobody thought that the rate of change of the frequency of the gene would be fixed over time. Maybe the environment will change in crazy non-linear ways in the future - and you'll have to redo your sums.

Basically, it isn't true that inclusive fitness theories assume that personal fitness is a linear sum of action fitness deltas. An action can change the environment (or the actor) in ways that affect the costs and benefits of future actions - and this is perfectly compatible with Hamilton's rule. It means that you may have to apply the rule more than once, is all.

Allen, Nowak, and Wilson's paper is like saying: I've got this crazy non-linear function, there's no way your linear approximation can match it. Except that: yes, there is - a series of linear functions can approximate any other function arbitrarily closely.

If you think that making this sort of objection is childish, I think that you're right - it is childish.

The paper says it was: "supported by a grant from the John Templeton Foundation". It seems like even more fuel for those who think that the Templeton Foundation is systematically distorting science.

Commentary

• Nowak and colleagues skewer Inclusive Fitness
• Now I know I am “El Lobo Solitario”: I don’t even agree with Allen, Nowak, and Wilson
• Inclusive fitness, models, and religious evolution

References

• Allen B, Nowak MA, & Wilson EO (2013) Limitations of inclusive fitness. Proceedings of the National Academy of Sciences of the United States of America
• Nowak MA, Tarnita CE, & Wilson EO (2010) The evolution of eusociality. Nature, 466 (7310), 1057-62

Kin selection is very general

Kin selection theory can only be applied when organisms are influenced by their kin. However this happens at least once in every organisms' lifetime - when they are born.

Kin selection applies to calculations of parental investment and cases where there is parent-offspring conflict. Since all multicellular creatures have a parent-offspring asymmetry and provide some kind of resource bolus to their offspring, that means that kin selection applies to all multicellular organisms.

In other organisms, kin selection is also often relevant. Many microscopic organisms exhibit limited dispersal - and so are surrounded by their kin. If your kin are nearby, you should typically curb your pollution, share your food, and commit suicide if you are compromised by pathogens. Many parasites spend much of their time inside their hosts - where they are often surrounded by their kin.

Kin selection also applies to cultural variation - in the form of cultural kin selection - where it is similarly ubiquitous.

The productivity of kin selection and group selection

West Griffin and Gardner wrote a comparison of kin selection and group selection, titled: Social semantics: how useful has group selection been?. Thay basically said that kin selection was the hands-down winner.

I think it is hard to argue that group selection has been anywhere near as productive as kin selection. However, group selection enthusiasts have been massively outnumbered over most of the recent history of the two ideas. Group selection defenders can claim that group selection has been less productive due to fewer man-hours expended on it.

The primary strengths of kin selection as I see them:

• Kin selection makes extensive use of relatedness (a concept group selection enthusiasts rarely mention). This facilitates making quantitative predictions.

• Kin selection has "Kin" in its title, which emphasizes the significance of kinship to the effect. Calling the effect "group selection" fails to to emphasize kinship. In practice this leads to applications where kinship isn't involved - and such applications are more often than not errors and mistakes.

• The field is more mature, established and respectable.

I think that group selection appeals more to those who are attracted to the scientific fringes - in the hope of finding useful things there. That's all very well, but the the scientific fringes have a lot of junk science in them as well. Some of it is to do with group selection. Take care with it.

A brief history of cultural kin selection

Kin selection applies to cultural variation. This is called cultural kin selection. It is an important topic.

Here's a brief history of the concept:

The observation that social and genetic kinskip were different in humans dates back to Lewis Henry Morgan in 1871.

Boyd and Richerson published one of the first articles on the topic in 1980 - connecting cultural kin selection and cultural group selection.0 An early modern article on the topic was written by Kristen Hawkes in 1983 and titled: "Kin Selection and Culture".

David Schneider's 1984 book "Critique of the Study of Kinship" argued that kinship was a different concept in different cultures, that people treating non-relatives as though they were kin was common - and that this reduced the significance of kin theories from biology when considering humans.

The idea of cultural kin selection appears in David Hull's (1988) book. David wrote:

Increasing one's conceptual inclusive fitness in science means increasing the number of replicates of one's contribution in the work of successive generations of other scientists.

...and...

science is a function of conceptual inclusive fitness.

Next, Francis Heylighen's (1992) paper: "Selfish Memes and the Evolution of Cooperation" stated:

The mechanism of kin selection can be extended to memes by redefining inclusive fitness as the fitness of a meme taking into account all its carriers. All individuals carrying the same meme can be viewed as relatives or kin insofar as this meme is concerned. Hence, the tendency to be altruistic towards offspring or close kin that follows from genetic evolution can be generalized to altruistic tendencies towards members of the same memetic or cultural group. The explanation for ultrasociality in the social insects on the basis of genetic inclusive fitness can be readily transposed to the emergence of ultrasociality in humans on the basis of memetic inclusive fitness.

He went on to explain how memes exploited kin-altruism towards relatives to redirect benefits towards cultural relatives:

many ethical systems explicitly refer to the ideal of "fraternity", and sometimes members of the same cultural group (e.g. monks or Freemasons) are supposed to call each other "brother". Though these are not brothers in the biological sense, the meme attempts to harness the innate tendency to behave altruistically towards kin and to use it for purposes different from the increase of genetic inclusive fitness.

Around the same time, Paul Allison published a series of papers on the topic - most notably "The Cultural Evolution of Beneficent Norms".

Heylighen revisited the idea of cultural kin in 1997, writing:

The genetic argument for altruism towards individuals carrying the same genes (kin selection) generalizes to altruism towards carriers of the same memes.

In 1997, David Hales wrote in "Memetic Evolution & Suboptimisation":

I now want to propose that two previously described extensions to basic natural selection can be applied subtlety within a meme framework: kin altruism and group selection.

There's a paper by John Evers (1998) called "A justification of societal altruism according to the memetic application of Hamilton's Rule", which attempts to directly adapt Hamilton's Rule to memetics - based on the idea of the "fraction of shared memes". John doesn't really go into the difficulties associated with this idea, though.

Also in 1998, "Cultural Software" by J. M. Balkin came out. Balkin clearly understood cultural kin selection, writing:

We often see people energetically promulgating their memes in the forms of beliefs, behaviors, artifacts, and customs while struggling with others who resist or disagree. Just as competition between biological kin groups can lead to strife, so can competition between cultural kin groups.

...and...

Just as individuals have varying degrees of genetic kinship, they also have varying degrees of memetic kinship. The two forms of kinship are cross-cutting: people can have many of the same memes even if they are completely unrelated. If the analogy to evolutionary arguments about kin-based altruism holds, then we would predict considerable altruistic behavior between people with lots of similar cultural software-for example, people of the same religion or culture, teachers and students, members of the same fraternity or club.

In 2000, David Hull showed that he understood the idea that kin selection extended into the cultural realm, writing: "One final example of similar processes operating in biological and memetic change is kin selection". He went on to give an example:

In science, scientists also distinguish between kin and non-kin, but the relevant genealogy is conceptual. The issue is not who holds similar ideas but who is conceptually connected to whom. The best way to increase the likelihood that you will be a successful scientist is to work under a successful scientist (Hull 1988).

Momme Von Sydow in a 2000 thesis wrote:

Corresponding to kin selection on the biological level, memes might ‘egoistically’ support their relatives in the same brain or in different brains, if they were identical (or similar enough). One might formulate a mathematical inequality analogous to Hamilton’s, where memes would replace genes.

In 2001, an important paper on tag-based cooperation came out - by Riolo, Cohen and Axelrod. It was followed by a large number of other papers on the topic. Riolo had come up with the idea much earlier - and had performed computer simulations of tag-based cooperation in 1992.

In 2001, Karl Sigmund and Martin Nowak, pointed out that cultural tag-based cooperation was a form of cultural kin selection.

Gordon Rakita (2003) rechristened cultural kin selection "kith selection". The term "kith selection" is kind-of cute - but it seems as though it is unnecessary jargon - and it doesn't fit in very well with "cultural group selection".

The idea of "cultural kin selection" appears again in Francis Heylighen's (2008) article: "Cultural Evolution and Memetics" - where he says:

Horizontal transmission of cooperation norms solves this problem, since the members of a cultural group are all memetically related to each other, sharing their memes rather than their genes. Therefore, cultural kin selection will extend to all members of the group (Evers, 1998). This entails a selective pressure for memes to support the fitness of the whole group of their carriers, e.g. by promoting cooperation.

In 2009, Gordon Rakita spelled out some of the implications of the idea of "kith selection" in detail. He started out by introducing a class of behaviours that kin selection acting on genes appears to explain poorly:

Kin selection does not seem to offer a suitable explanation for instances in which soldiers give their lives for their comrades. The conditions necessary for the model are not met, for example, soldiers are rarely genetically related. Given this, how are such acts to be explained?

He then offered a fine explanation - involving selection acting on memes:

A meme that encourages a soldier to throw himself on a grenade to save his comrades will negatively impact its future prospects of being transmitted to others. (This behavior is similar to that of the worker ant who forgoes reproduction in favour of the reproductive potential of the queen.) However, if the soldier’s comrades also carry the meme for grenade smothering, then the soldier’s behavior will indirectly impact the future replicative success of those other copies of the altruistic meme. This mimetic model of kin selection I term kith selection, kith being unrelated but like-minded acquaintances

Recently Mark Pagel's Wired For Culture book offered an explanation of human cooperation in terms of cultural "green beard" effects. In the process, he pointed out that such green beard effects were a kind of kin selection.

For references, see the cultural kin selection bibliography.

Productive chickens: group selection vs kin selection

Group selection advocates often cite laboratory studies on breeding chickens to maximise egg production.

What they don't mention is that the birds involved were housed by sire family, were close relatives - and the whole process was conceived at the time in terms of kin selection. Check out this quote:

Craig (1982) advocated kin selection, in which pullets are housed in cages together as families with mean performance used as the criterion of selection. He hypothesized that families that perform best tend to have those physiological and behavioral characteristics most appropriate for group well-being and productivity. Muir (1985), Craig and Muir (1993), and Muir (1994) hypothesized that kin selection would favor cooperative tolerant behavior and concluded that selection on family means, when families are kept together as family groups, provides a method of improving traits in which behavioral interactions influence overall well-being and productivity.

The challenge for group selection has always been distinguishing itself from its widely-accepted rival, kin selection. The case of the chickens surely represents a failure to do this.

References

• Muir, W. M. and Crig, J. V. (1998) Improving Animal Well-Being Through Genetic Selection

Tree trunks are not wasteful selfishness

I re-read the Dawkins essay "Gods Utility Function" recently. Dawkins offers an argument against high-level selection in it. However, the article gives a string of dodgy examples. He argues that tree trunks are wasteful:

Why are forest trees so tall? Simply to overtop rival trees. A “sensible” utility function would see to it that they were all short. They would get exactly the same amount of sunlight, with far less expenditure on thick trucks and massive supporting buttresses. But if they were all short, natural selection couldn’t help favoring a variant individual that grew a little taller. The ante having been upped, others would have to follow suit. Nothing can stop the whole game escalating until all trees are ludicrously and wastefully tall. It is ludicrous and wasteful only from the point of view of a rational economic planner thinking in terms of maximizing efficiency. But it all makes sense once you understand the true utility function-genes are maximizing their own survival.

This is wrong. Very tall tree trunks are found in the most efficient dissipating regions of the earth: rain forests. These extract far more energy from incident sunlight that the grasslands that resemble the type of ecosystem which Dawkins is describing as more efficient. Those tree trunks pay off in efficiency terms. If you have a flat surface, it heats up - and the incident energy radiates back into space. You need a lot of surface area to allow water evaporation to keep you cool. The depth produced by tall trees helps to provides such a surface.

In the essay, Dawkins also describes elephant seal sex ratios as "inefficient":

The sex ratio-the proportion of males to females-in wild populations is usually 50:50. This seems to make no economic sense in those many species in which a minority of males has an unfair monopoly of the females: the harem system. In one well-studied population of elephant seals, 4 percent of the males accounted for 88 percent of all the copulations. Never mind that God’s Utility Function in this case seems so unfair for the bachelor majority. What is worse, a cost-cutting, efficiency-minded deity would be bound to spot that the deprived 96 percent are consuming half the population’s food resources (actually more than half, because adult male elephant seals are much bigger than females). The surplus bachelors do nothing except wait for an opportunity to displace one of the lucky 4 percent of harem masters. How can the existence of these unconscionable bachelor herds possibly be justified? Any utility function that paid even a little attention to the economic efficiency of the community would dispense with the bachelors. Instead, there would be just enough males born to fertilize the females.

Unfortunately for this argument, the surplus of males helps the population weed out parasite-ridden and mutation-loaded individuals. A population with fewer males would mean males of lower quality breeding - with the quality of individuals cumulatively declining over the generations. Dawkins' economic argument about how the makes are redundant is simply mistaken. The deaths of the failed males helps the population to adapt by marking out the adjacent gene-space that represent reduced quality - helping the population flee from its parasites and decrease its mutational load.

Perhaps it is possible for trees to be too tall - and for there to be too many elephant seal males. However tall trees and numerous male elephant seals proves very little.

Of course, the basic theme of this essay - that "the true utility function of life, that which is being maximized in the natural world, is DNA survival" is hopelessly mistaken.

Dawkins' argument against multi-level selection in this essay is also incoherent.