Learning is a widespread ability among animals and, like physical traits, is subject to evolution. But how did learning first arise? What selection pressures and phenotypic preconditions fostered its evolution? Neither the fossil record nor phylogenetic comparative studies provide answers to these questions. Here, we take a novel approach by studying digital organisms in environments that promote the evolution of navigation and associative learning. Starting with a nonlearning sessile ancestor, we evolve multiple populations in four different environments, each consisting of nutrient trails with various layouts. Trail nutrients cue organisms on which direction to follow, provided they evolve to acquire and use those cues. Thus, each organism is tested on how well it navigates a randomly selected trail before reproducing. We find that behavior evolves modularly and in a predictable sequence, where simpler behaviors are necessary precursors for more complex ones. Associative learning is only one of many successful behaviors to evolve, and its origin depends on the environment possessing certain information patterns that organisms can exploit. Environmental patterns that are stable across generations foster the evolution of reflexive behavior, while environmental patterns that vary across generations but remain consistent for periods within an organism's lifetime foster the evolution of learning behavior. Both types of environmental patterns are necessary, since the prior evolution of simple reflexive behaviors provides the building blocks for learning to arise. Finally, we observe that an intrinsic value system evolves alongside behavior and supports associative learning by providing reinforcement for behavior conditioning.

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