One of the most fundamental functions of the brain is to predict upcoming events on the basis of the recent past. A closely related function is to signal when a prediction has been violated. The identity of the brain regions that mediate these functions is not known. We set out to determine whether they are implemented at the level of single neurons in the visual system. We gave monkeys prolonged exposure to pairs of images presented in fixed sequence so that each leading image became a strong predictor for the corresponding trailing image. We then monitored the responses of neurons in the inferotemporal cortex to image sequences that obeyed or violated the transitional rules imposed during training. Inferotemporal neurons exhibited a transitional surprise effect, responding much more strongly to unpredicted transitions than to predicted transitions. Thus, neurons even in the visual system make experience-based predictions and react when they fail.macaque | vision | plasticity T he inferotemporal cortex (ITC), the terminus of the ventral stream of visual areas (1), plays a critical role in object vision (2, 3). ITC neurons respond with individual patterns of selectivity to complex images (4). Training monkeys to discriminate between images (5-7), categorize them (8, 9), or form associations between them (10-12) induces functional changes among neurons in the ITC which have the effect of strengthening the representation of image attributes relevant to task performance. Even passive viewing causes changes in neuronal visual responsiveness. Repeated viewing of a single image leads to a weakening of responses to it (13,14). Repeated viewing of two images close together in time leads to pair coding: a tendency for neurons responsive to one image also to respond to the other (15-17). The effects of passive viewing, because they do not depend on task demands, fall into the category of unsupervised statistical learning.An important form of unsupervised learning not previously studied at the level of single neurons concerns transitional statistics. The learning of transitional statistics has been the focus of much behavioral study in humans because it is thought to underlie the development during infancy of the ability to perceive event boundaries including word boundaries in speech (18,19). Human infants passively exposed to a stimulus stream in which certain visual images always follow certain others automatically register the transitional rules as evidenced by their orienting preferentially to a test stream containing novel transitions (20). The adult human brain is sensitive to transitional probabilities, as evidenced by its generating strong responses to improbable transitions at the level of scalp potential and blood oxygenation measures (21-27). Monkeys, like human infants, have been reported to learn transitional probabilities and to orient preferentially to improbable transitions (28). No effort has been made as yet to characterize the underlying neuronal mechanisms (29). We hypothesized that neurons in the ITC ...