In my last post, I discussed the framework for cultural evolution laid out by Claire El Mouden and colleagues in a new article in the Journal of Evolutionary Biology (ungated pdf and supporting information). By setting out clear definitions for the analysis of cultural evolution, such as cultural relatedness and fitness, a workable framework using evolutionary biology’s Price equation can be developed.

As I noted in that post, it is when the biological and genetic frameworks are laid on top of each other, as is the focus of the dual-inheritance literature, that things get interesting. With their framework, El Mouden and friends tackle a couple of the prominent gene-culture evolution questions.

The first question is to what extent cultural evolution increases or decreases genetic fitness. The authors note that a theme of the gene-culture evolution literature is that cultural evolution made the scale and distribution of today’s human population possible. How does this work?

For there to be a positive correlation between cultural and genetic fitness, those who have the most cultural influence must also leave the most offspring. It is easy to see circumstances where this holds, with those of high social status tending to have both cultural influence and more offspring (look at the number of partners of rock stars).

An example to the contrary is low fertility in developed countries. El Mouden and colleagues reference work by Richerson and Boyd, who suggest that high status is given to professions with high investment in education, which would allow the behaviour to spread despite the negative relationship between education and fertility.

However, El Mouden and friends show that this mismatch between cultural and genetic fitness is evolutionarily unstable as genetic natural selection acts to align genetic and cultural fitness. They suggest two reasons for this, one involving transmission and the other selection.

Their transmission explanation relies on the already evolved propensity for people to avoid behaviours that are harmful to their genetic interests. For example, the nausea produced when consuming toxins would fight against any cultural pressure to eat toxic food. However, it is an interesting question as to how general this transmission effect would be, as many cultural forces are novel and without precedent in our evolutionary history. Transmission may be unlikely to constrain our desire for high status professions that require large investments in education any time soon.

The genetic selection explanation would seem to have broader power. In the supplementary information, the authors ask us to consider a population where cultural and genetic fitness were not aligned. Now, imagine a mutant in that population that causes people to pay attention to a cultural trait that is more highly correlated with genetic fitness. As these mutants have higher genetic fitness, they increase in proportion of the population, and cultural and genetic fitness are now more correlated. Cultural fitness now promotes genetic fitness. In the long-run, the two will be perfectly correlated (the exception being where cultural traits are neutral to genetic fitness).

The catch in that last sentence is the “long-run”. As cultural evolution can be so much faster than genetic evolution, systems can be far from genetic equilibrium until the genetic response evolves. Fertility in developed countries would be an example of this. There may also be some constraints that prevent perfect alignment, such as the presence of appropriate learning mechanisms.

This interaction of genetic and cultural evolution gets most interesting is when we turn to the evolution of altruism. In examining this question we must remember that genetic and cultural fitness are distinct. Cultural altruism reduces the altruist’s cultural fitness; that is, their influence. As a result, the claim that cultural evolution increases genetic altruism (the more common claim in the gene-culture evolution literature) needs to be made carefully.

As an illustration, consider this interesting example from the paper. A stranger is being attacked, so a good Samaritan steps in to defend them and dies as a result. Whether this is culturally altruistic would depend on whether the Samaritan’s deed was copied. If so, then the Samaritan’s act would actually have been increase cultural fitness as it would have increased their influence in respect of that cultural trait.

Conversely, their death is genetically altruistic. As a result, genetic selection would tend to act against it. Those who ignore this cultural trait will have higher genetic fitness, grow in proportion of the population and eventually bring cultural and genetic fitness into alignment.

So what of behaviours that are both culturally and genetically altruistic? Whether the behaviour spreads will depend on the degree of cultural and genetic relatedness.

Evidence suggests that cultural relatedness within ethnic groups is higher than genetic relatedness (although it is still not high in absolute terms, with more within group than between group variation). This means that there are a wider range of circumstances for which cultural altruism can emerge than for genetic altruism. However, that domain in which cultural but not genetic altruism is likely to emerge will be subject to the forces described above to align cultural and genetic fitness.

Another important point is that each cultural trait should be considered separately. Even though a group may have the same language, giving them high relatedness for this cultural trait, this does not mean that they have the same views on giving their lives for strangers, for which they may have low cultural relatedness. Consideration of the conditions for altruism need to consider the specific cultural trait.

There are many other interesting points in the article - I recommend reading the whole thing - but I will close with a point on the practicality of modelling cultural evolution in this way. El Mouden and friends note that there is a host of complications not present in the genetic case. Cultural relatedness can vary wildly across cultural traits, whereas the nature of genetic transmission means that relatedness is similar across most of the genome. Recognising the pattern of inheritance is also a challenge, as ancestor numbers can vary in number and be of vastly different biological ages. In that context, there is no such thing as a standard length of generation.

So although this paper presents a nice approach to cultural evolution, it does not present an approach that is easily applied to empirical observation. However, given the lack of clarity across much of the gene-culture evolution literature, particularly when examined across authors and papers, it is nice to see an attempt to achieve some conceptual coherence.