What modulates the Mirror Neuron System during action observation? Multiple factors involving the action, the actor, the observer, the relationship between actor and observer, and the context

Seeing an agent perform an action typically triggers a motor simulation of that action in the observer's Mirror Neuron System (MNS). Over the past few years, it has become increasingly clear that during action observation the patterns and strengths of responses in the MNS are modulated by multiple factors. The first aim of this paper is therefore to provide the most comprehensive survey to date of these factors. To that end, 22 distinct factors are described, broken down into the following sets: six involving the action; two involving the actor; nine involving the observer; four involving the relationship between actor and observer; and one involving the context. The second aim is to consider the implications of these findings for four prominent theoretical models of the MNS: the Direct Matching Model; the Predictive Coding Model; the Value-Driven Model; and the Associative Model. These assessments suggest that although each model is supported by a wide range of findings, each one is also challenged by other findings and relatively unaffected by still others. Hence, there is now a pressing need for a richer, more inclusive model that is better able to account for all of the modulatory factors that have been identified so far.