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The ability to predict the consequences of one's behavior in a particular environment is a mechanism for adaptation. In the absence of any cost to this activity, we might expect agents to choose behaviors that maximize their fitness, an example of directed innovation. This is in contrast to blind mutation, where the probability of becoming a new genotype is independent of the fitness of the new genotypes. Here, we show that under environments punctuated by rapid reversals, a system with both genetic and cultural inheritance should not always maximize fitness through directed innovation. This is because populations highly accurate at selecting the fittest innovations tend to over-fit the environment during its stable phase, to the point that a rapid environmental reversal can cause extinction. A less accurate population, on the other hand, can track long term trends in environmental change, keeping closer to the time-average of the environment. We use both analytical and agent-based models to explore when this mechanism is expected to occur.

Previous work has shown that mutation bias can direct evolutionary trends in genotypic space under strong selection and rare mutation. We present an extension of this work to general traits of the organism. We do this by allowing many different genotypes, with different fitnesses, to have the same trait value. This approach makes novel predictions and shows that the outcome of evolution for a trait is influenced by mutation bias as well as the fitness distribution of the genotypes that have the same trait value. This distribution can alter evolution in interesting ways, depending on the likelihood of generating high fitness mutants. We also show that mutation bias can direct evolution when many mutants are present at any one time. We demonstrate that mutation bias can drive longterm evolutionary trends when the environment is constantly changing. Under biologically realistic conditions, we show that mutation bias can counter strong gradients of environmental selection over time. We conclude that evolutionary trends can be quite independent of the environment, even when they depress population fitness. Finally, we show that entropy can be a powerful source of mutation bias and can drive evolutionary trends. V C 2015 Wiley Periodicals, Inc. Complexity 21: [331][332][333][334][335][336][337][338][339][340][341][342][343][344][345] 2016

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A process akin to biological evolution is one of the most promising candidates today for producing a general theory of cultural evolution. Current understanding of this process focuses on both drift as well as the selection of cultural variation as the primary vehicles of cultural change. Here, I show that natural selection can produce cultural change in the direction of the generation of cultural variation. I show how this mechanism can result in long-term cultural trends and how it adds to known mechanisms. I present examples to show how this theory is compatible with documented cultural and historical change.