Minimally sufficient conditions for the evolution of social learning and the emergence of non-genetic evolutionary systems

Social learning, defined as the imitation of behaviors performed by others, is recognized as a distinctive characteristic in humans and several other animal species. Previous work has claimed that the evolutionary fixation of social learning requires decision-making cognitive abilities that result in transmission bias (e.g., discriminatory imitation) and/or guided variation (e.g., adaptive modification of behaviors through individual learning). Here, we present and analyze a simple agent-based model that demonstrates that the transition from instinctive actuators (i.e., non-learning agents whose behavior is hardcoded in their genes) to social learners (i.e., agents that imitate behaviors) can occur without invoking such decision-making abilities. The model shows that the social learning of a trait may evolve and fix in a population if there are many possible behavioral variants of the trait, if it is subject to strong selection pressure for survival (as distinct from reproduction), and if imitation errors occur at a higher rate than genetic mutation. These results demonstrate that the (sometimes implicit) assumption in prior work that decision-making abilities are required is incorrect, thus allowing a more parsimonious explanation for the evolution of social learning that applies to a wider range of organisms. Furthermore, we identify genotype-phenotype disengagement as a signal for the imminent fixation of social learners, and explain the way in which this disengagement leads to the emergence of a basic form of cultural evolution (i.e., a non-genetic evolutionary system).