Selection of the fittest or selection of the luckiest: the emergence of Goodhart's law in evolution

Biological evolution depends on the passing down to subsequent generations of genetic information encoding beneficial traits, and on the removal of unfit individuals by a selection mechanism. However, selection acts on phenotypes, and is affected by random contingencies. Thus, a combination of fitness and luck determines which individuals will successfully reproduce and give rise to the next generation. To understand how randomness in the selection mechanism affects the long-term patterns of evolution, we studied an idealized evolution model. We show through simulations and mathematical analysis, that the speed of adaptation increases with increasing selection pressure only up to a threshold. Beyond the threshold, any increase of the selection pressure results in more weight given to random effects rather than on genetic fitness in determining which individuals will successfully reproduce. This severely reduces the speed of adaptation and the diversity in the gene pool. Our findings may be considered as a biological instance of Goodhart's law: "When a measure becomes a target, it ceases to be a good measure". Finally, we show that this intricate response of evolution to natural selection can be mathematically explained by a novel phase transition for pulled traveling waves.