Functional organization of the insula and inner perisylvian regions
- aDipartimento di Neuroscienze, Università di Parma, 43100 Parma, Italy;
- bDepartment of Neurobiology and Behavior, State University of New York, Stony Brook, NY 11794-5230;
- cBrain Center for Social and Motor Cognition, Italian Institute of Technology, 43100 Parma, Italy; and
- dDipartimento di Psicologia Generale, Università di Padova, 35131 Padua, Italy
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Edited* by Charles Gross, Princeton University, Princeton, NJ, and approved May 3, 2012 (received for review January 5, 2012)

Abstract
In the last few years, the insula has been the focus of many brain-imaging studies, mostly devoted to clarify its role in emotions and social communication. Physiological data, however, on which one may ground these correlative findings are almost totally lacking. Here, we investigated the functional properties of the insular cortex in behaving monkeys using intracortical microstimulation. Behavioral responses and heart rate changes were recorded. The results showed that the insula is functionally formed by two main subdivisions: (i) a sensorimotor field occupying the caudal–dorsal portion of the insula and appearing as an extension of the parietal lobe; and (ii) a mosaic of orofacial motor programs located in the anterior and centroventral insula sector. These programs show a progressive shift from dorsally located nonemotional motor programs (ingestive activity) to ventral ones laden with emotional and communicative content. The relationship between ingestive and other behaviors is discussed in an evolutionary perspective.
Footnotes
↵1A.J. and F.C. contributed equally to this work.
- ↵2To whom correspondence may be addressed. E-mail: fausto.caruana{at}iit.it. or vittorio.gallese{at}unipr.it.
Author contributions: V.G. and G.R. designed research; A.J. and F.C. performed research; A.J., F.C., and I.S. analyzed data; and A.J., F.C., and G.R. wrote the paper.
The authors declare no conflict of interest.
↵*This Direct Submission article had a prearranged editor.
This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.1073/pnas.1200143109/-/DCSupplemental.
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