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152 WILLIAM H. MERIGAN AND TATIANA PASTERNAK

hand-reaching movements, and during memory tasks requiring withholding a movement (Snyder et al. 1997, 1998). Moreover, the representation of space generated in parietal neurons is continuously updated in conjunction with the impending motor action, and these dynamic transformations are modulated by attention (Colby and Goldberg 1999). In addition, many parietal neurons respond to various types of optic flow motion (Read and Siegel 1997; Siegel and Read 1997; Phinney and Siegel 2000) as well as three-dimensional features of objects (Sakata et al. 1994; Taira et al. 2000). These properties point to a role for parietal neurons in the integration of sensory signals, and in constructing a representation of extrapersonal space in preparation for motor action.

A relatively small number of lesion studies have been performed in non-human primates to examine the functional contribution of these areas to behaviour. An important role of posterior parietal cortex in spatial aspects of task performance was first shown by Ridley and Ettlinger (1975) and Ratcliff et al. (1977). Subsequently, Latto (1986), tested monkeys after bilateral lesions of area 7a and found deficits in the performance of a spatial landmark test. Further evidence for a role of parietal cortex in space coding was reported by Quintana and Fuster (l993), who applied cooling to areas 5 and 7 of the parietal cortex. They found that the monkeys showed decreased speed and accuracy in both reaching and eye movements during the performance of tasks that required the processing and retention of spatial information. More recently, Li et al. (1999) reported deficits in memory-guided saccadic eye movements after reversible inactivation of LIP, produced by injections of muscimol. They concluded that LIP neurons play a direct role in processing incoming sensory information to programme saccadic eye movements. Inaccuracy in reaching towards visual targets after lesions of areas 7a, 7b, and LIP was observed by Rushworth et al. (1997).

Thus, lesions of parietal cortex appear to disrupt the ability of a subject to coordinate movements relative to him/herself or to other locations in extrapersonal space.

General summary and conclusions

Lesions of cortical visual areas in the macaque have proved to be a powerful and informative method for elucidating the functional specializations of visual cortical areas. Lesions of primary visual cortex cause profound visual loss, whereas lesions higher in the visual pathways cause less dramatic, but selective visual loss. Within the ventral pathway, lesions cause selective loss of orientation-based discriminations, especially those involving borders defined by texture or illusory contour, as well as impaired colour vision and discrimination of complex forms. Lesions of the dorsal pathway disrupt direction and speed discriminations, and there is now evidence that some components of this pathway play a role not only in processing this information, but also in retaining it in memory. Dorsal pathway lesions have also disrupted speed and accuracy in reaching and making eye movements, orienting in space, and discriminations involving motion shear and structure from motion. A number of lesion studies have also produced disruptions of more complex behaviours such as reaching, eye movements, and learning and attention.

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Many studies have demonstrated a remarkable degree of recovery after lesions in both the ventral and the dorsal pathways, although the recovery appears to be much more pronounced in the dorsal than in the ventral pathway. This difference may not be as large as it currently seems, because the studies of dorsal pathway lesions that have shown postlesion improvements were better designed to reveal such recovery and to study their time course. However, both the transitory and permanent behavioural effects of lesions are consistent with the description of primate visual cortex as two relatively parallel streams of processing. Ventral pathway lesions disrupt primarily the perception of form and colour, while dorsal pathway lesions affect largely the perception of motion and space.

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