This perspective piece came about through the Generative Adversarial Collaboration (GAC) series of workshops organized by the Computational Cognitive Neuroscience (CCN) conference in 2020. We brought together a number of experts from the field of theoretical neuroscience to debate emerging issues in our understanding of how learning is implemented in biological recurrent neural networks. Here, we will give a brief review of the common assumptions about biological learning and the corresponding findings from experimental neuroscience and contrast them with the efficiency of gradient-based learning in recurrent neural networks commonly used in artificial intelligence. We will then outline the key issues discussed in the workshop: synaptic plasticity, neural circuits, theory-experiment divide, and objective functions. Finally, we conclude with recommendations for both theoretical and experimental neuroscientists when designing new studies that could help to bring clarity to these issues.