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1 University of Wales, Bangor, United Kingdom
* To whom correspondence should be addressed. E-mail: Edmund.Rolls{at}psy.ox.ac.uk.
In an event-related fMRI study in humans it was shown, first, that water produces activations in cortical taste areas (in particular the frontal operculum/anterior insula which is the primate primary taste cortex, and the caudal orbitofrontal / secondary taste cortex) comparable to those produced by the prototypical tastants salt and glucose. Second, the activations in the frontal operculum/anterior insula produced by water when thirsty were still as large after the subjects had consumed water to satiety. Third, in contrast, the responses to water in the caudal orbitofrontal cortex were modulated by the physiological state of the body, in that responses to the oral delivery of water in this region were not found after the subjects had drunk water to satiety. Fourth, further evidence that the reward value or pleasantness of water is represented in the orbitofrontal cortex was that a positive correlation with the subjective ratings of the pleasantness and the refreshing value of the water was found with activations in the caudal and anterior orbitofrontal cortex, and also in the anterior cingulate cortex. Fifth, it was found that a region of the middle part of the insula was also activated by water in the mouth, and further that this activation only occurred when thirsty. Sixth, analyses comparing pre- and post-satiety periods (i.e. when thirsty and when not thirsty) independently of stimulus delivery revealed higher activity levels in the rostral anterior cingulate cortex. The activity of the rostral anterior cingulate cortex thus appears to reflect the thirst level or motivational state of the subjects.
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