Mirror neurons (and beyond) in the macaque brain: An overview of 20 years of research

Mirror neurons are a class of neurons in the ventral pre-motor cortex (area F5) and inferior parietal lobule (area PFG) that respond during the execution as well as the observation of goal-directed motor acts. These intriguing response properties stirred an intense debate in the scientific community with respect to the possible cognitive role of mirror neurons. The aim of the present review is to contribute to this debate by providing, in a single paper, an extended summary of 20 years of neurophysiological research on mirror neurons in the macaque. To this end, I provide a comprehensive description of the methodology and the main results of each paper about mirror neurons published since their first report in 1992. Particular care was devoted in reporting the different response characteristics and the percentages of neurons exhibiting them in relation to the total number of studied neurons. Furthermore, I also discuss recent results indicating that mirror neurons might not be confined to areas F5 and PFG and that "mirroring" might not be limited to action observation. Finally, I offer a unifying framework for many of the results discussed here by speculating that a potential functional role of mirror neurons might be, during action observation, to generalize from the particular grasping movement being observed to the "concept" of grasping. (C) 2012 Elsevier Ireland Ltd. All rights reserved.