Learning at Variable Attentional Load Requires Cooperation of Working Memory, Meta-learning, and Attention-augmented Reinforcement Learning.

Flexible learning of changing reward contingencies can be realized with different strategies. A fast learning strategy involves using working memory of recently rewarded objects to guide choices. A slower learning strategy uses prediction errors to gradually update value expectations to improve choices. How the fast and slow strategies work together in scenarios with real-world stimulus complexity is not well known. Here, we aim to disentangle their relative contributions in rhesus monkeys while they learned the relevance of object features at variable attentional load. We found that learning behavior across six monkeys is consistently best predicted with a model combining (i) fast working memory and (ii) slower reinforcement learning from differently weighted positive and negative prediction errors as well as (iii) selective suppression of nonchosen feature values and (iv) a meta-learning mechanism that enhances exploration rates based on a memory trace of recent errors. The optimal model parameter settings suggest that these mechanisms cooperate differently at low and high attentional loads. Whereas working memory was essential for efficient learning at lower attentional loads, enhanced weighting of negative prediction errors and meta-learning were essential for efficient learning at higher attentional loads. Together, these findings pinpoint a canonical set of learning mechanisms and suggest how they may cooperate when subjects flexibly adjust to environments with variable real-world attentional demands.