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The Journal of Neurophysiology Vol. 84 No. 6 December 2000, pp. 3026-3035
Copyright ©2000 by the American Physiological Society
Wellcome Department of Cognitive Neurology, Institute of Neurology, London WC1N 3BG, United Kingdom
Ramnani, N.,
I. Toni,
O. Josephs,
J. Ashburner, and
R. E. Passingham.
Learning- and Expectation-Related Changes in the Human Brain
During Motor Learning. J. Neurophysiol. 84: 3026-3035, 2000. We have studied a simple form of motor learning
in the human brain so as to isolate activity related to motor learning
and the prediction of sensory events. Whole-brain, event-related
functional magnetic resonance imaging (fMRI) was used to record
activity during classical discriminative delay eyeblink conditioning.
Auditory conditioned stimulus (CS+) trials were presented either
with a corneal airpuff unconditioned stimulus (US,
paired), or without a US (unpaired). Auditory CS
trials were never reinforced with a US. Trials were presented
pseudorandomly, 66 times each. The subjects gradually produced
conditioned responses to CS+ trials, while increasingly differentiating
between CS+ and CS
trials. The increasing difference between
hemodynamic responses for unpaired CS+ and for CS
trials evolved
slowly during conditioning in the ipsilateral cerebellar cortex (Crus
I/Lobule HVI), contralateral motor cortex and hippocampus. To localize
changes that were related to sensory prediction, we compared trials on
which the expected airpuff US failed to occur (Unpaired CS+) with
trials on which it occurred as expected (Paired CS+). Error-related
signals in the contralateral cerebellum and somatosensory cortex were
seen to increase during learning as the sensory prediction became
stronger. The changes seen in the ipsilateral cerebellar cortex may be
due either to the violations of sensory predictions, or to
learning-related increases in the excitability of cerebellar neurons to
presentations of the CS+.
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