Review
Cognitive and emotional influences in anterior cingulate cortex

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Abstract

Anterior cingulate cortex (ACC) is a part of the brain’s limbic system. Classically, this region has been related to affect, on the basis of lesion studies in humans and in animals. In the late 1980s, neuroimaging research indicated that ACC was active in many studies of cognition. The findings from EEG studies of a focal area of negativity in scalp electrodes following an error response led to the idea that ACC might be the brain’s error detection and correction device. In this article, these various findings are reviewed in relation to the idea that ACC is a part of a circuit involved in a form of attention that serves to regulate both cognitive and emotional processing. Neuroimaging studies showing that separate areas of ACC are involved in cognition and emotion are discussed and related to results showing that the error negativity is influenced by affect and motivation. In addition, the development of the emotional and cognitive roles of ACC are discussed, and how the success of this regulation in controlling responses might be correlated with cingulate size. Finally, some theories are considered about how the different subdivisions of ACC might interact with other cortical structures as a part of the circuits involved in the regulation of mental and emotional activity.

Section snippets

Anatomy and lesion studies

The cingulate gyri were broadly defined by Broca as belonging to the limbic lobe. With the advent of more-precise methods for studying anatomical connectivity, cytoarchitecture and function, it is now clear that cingulate cortex encompasses numerous specialized subdivisions that subserve a vast array of cognitive, emotional, motor, nociceptive and visuospatial functions5.

Anterior cingulate cortex can be differentiated from posterior cingulate cortex on the basis of cytoarchitecture and patterns

Imaging studies

Functional neuroimaging techniques, which include PET and fMRI, have provided invaluable insights into the roles that ACC has in cognitive and emotional processing. The cognitive subdivision is part of a distributed attentional network. It maintains strong reciprocal interconnections with lateral prefrontal cortex (BA 46/9), parietal cortex (BA 7), and premotor and supplementary motor areas6. Various functions have been ascribed to the ACcd, including modulation of attention or executive

Electrical studies of error monitoring

Recent electrophysiological studies using event-related potentials (ERPs) have shown that when subjects make an erroneous response in a behavioral task there is a negative deflection in the stimulus- and response-locked ERP (24, 25) called the error-related negativity (ERN)25. The ERN is sharper and clearer in the response-locked averages and has a centromedial scalp distribution. Its onset is simultaneous with the erroneous response and peaks approximately 100 ms after response onset (see Fig.

Developmental studies

Although the cingulate cortex is an ancient structure, there is evidence that it has undergone changes in recent evolution32. There is no doubt that cingulate cortex, like other frontal areas, undergoes a long developmental process. Conel has shown extensive development of the connectivity of cingulate cortex from infancy to late childhood33. However, few studies to date have related these changes to behavior.

During later childhood, neuroimaging methods can be applied. In one study of 5- to

Functional theories

Before attempting to synthesize and integrate existing theories, it is important to emphasize that ACC does not act in a vacuum. Many researchers have identified ACC as an important component of parallel distributed attentional40, 41, 42, 43 and emotional5, 6 networks. Imaging studies have begun to provide a basis for exploring frontal networks involved in effortful control44. These studies suggest that both dorsal ACC and areas of the lateral prefrontal cortex operate together during tasks

Outstanding questions

  • What is the nature of the interaction between lateral prefrontal cortical areas and the various subdivisions of the cingulate cortex?

  • What types of interaction between cognitive and emotional processes are possible? Do they always work by mutual inhibition?

  • Does the ERN that follows an error and the ERN that follows feedback come from the same or different areas of the cingulate, and how do their locations relate to the cognitive and emotional subdivisions?

  • When do the cognitive and emotional

Acknowledgements

G.B. was supported by NIMH 01611, NARSAD (as the Forrest C. Lattner Foundation Investigator) and the Harvard/MIT Division of Health Sciences and Technology Clinical Investigator Training Program. M.P. was supported by grants from the James S. McDonnell Foundation and the National Science Foundation grant BCS 9907831 to the Sackler Institute. The authors thank B. Vogt for his helpful comments.

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