Human brain activity during mental imagery exhibits signatures of inference in a hierarchical generative model

Humans have long wondered about the function of mental imagery and its relationship to vision. Although visual representations are utilized during imagery, the computations they subserve are unclear. Building on a theory that treats vision as inference about the causes of sensory stimulation in an internal generative model, we propose that mental imagery is inference about the sensory consequences of predicted or remembered causes. The relation between these complementary inferences yields a relation between the brain activity patterns associated with imagery and vision. We show that this relation has the formal structure of an echo that makes encoding of imagined stimuli in low-level visual areas resemble the encoding of seen stimuli in higher areas. To test for evidence of this echo effect we developed imagery encoding models---a new tool for revealing how imagined stimuli are encoded in brain activity. We estimated imagery encoding models from brain activity measured with fMRI while human subjects imagined complex visual stimuli, and then compared these to visual encoding models estimated from a matched viewing experiment. Consistent with an echo effect, imagery encoding models in low-level visual areas exhibited decreased spatial frequency preference and larger, more foveal receptive fields, thus resembling visual encoding models in high-level visual areas where imagery and vision appeared to be almost interchangeable. Our findings support an interpretation of mental imagery as a predictive inference that is conditioned on activity in high-level visual cortex, and is related to vision through shared dependence on an internal model of the visual world.