We recently discovered stimulus-specific interactions between cell assemblies in cat primary visual cortex that could constitute a global linking principle for feature associations in sensory and motor systems: stimulus-induced oscillatory activities (35-80 Hz) in remote cell assemblies of the same and of different visual cortex areas mutually synchronize, if common stimulus features drive the assemblies simultaneously. Based on our neurophysiological findings we simulated feature linking via synchronizations in networks of model neurons. The networks consisted of two one-dimensional layers of neurons, coupled in a forward direction via feeding connections and in lateral and backward directions via modulatory linking connections. The models' performance is demonstrated in examples of region linking with spatiotemporally varying inputs, where the rhythmic activities in response to an input, that initially are uncorrelated, become phase locked. We propose that synchronization is a general principle for the coding of associations in and among sensory systems and that at least two distinct types of synchronization do exist: stimulus-forced (event-locked) synchronizations support “crude instantaneous” associations and stimulus-induced (oscillatory) synchronizations support more complex iterative association processes. In order to bring neural linking mechanisms into correspondence with perceptual feature linking, we introduce the concept of the linking field (association field) of a local assembly of visual neurons. The linking field extends the concept of the invariant receptive field (RF) of single neurons to the flexible association of RFs in neural assemblies.
The analysis of a peripheral visual location can be improved in two ways: either by orienting one's gaze (usually by making a foveating saccade) or by 'covertly' shifting one's attention to the peripheral location without making an eye movement. The premotor theory of attention holds that saccades and spatial shifts of attention share a common functional module with a distinct neuronal basis. Using single-unit recording from the brains of trained rhesus monkeys, we investigated whether the superior colliculus, the major subcortical center for the control of saccades, is part of this shared network for attention and saccades. Here we show that a distinct type of neuron in the intermediate layer of the superior colliculus, the visuomotor neuron, which is known to be centrally involved in the preparation of saccades, is also active during covert shifts of attention.
One of the earliest computational principles attributed to the cerebellum was the measurement of time. This idea was originally suggested on anatomical grounds, and was taken up again to explain some of the deficits in cerebellar patients. The contribution of the cerebellum to eye movements, in contrast, has traditionally been discussed in the context of motor learning. This view has received support from the loss of saccade adaptation, one of the key examples of motor learning, following lesions of the posterior cerebellar vermis. However, the relationship between the properties of saccade-related vermal Purkinje cells and the behavioural deficits that follow lesions is unclear. Here we report results from single-unit recording experiments on monkeys that reconcile the seemingly unrelated concepts of timing and motor learning. We report that, unlike individual Purkinje cells, the population response of larger groups of Purkinje cells gives a precise temporal signature of saccade onset and offset. Thus a vermal population response may help to determine saccade duration. Modifying the time course of the population response by changing the weights of the contributing individual Purkinje cells, discharging at different times relative to the saccade, would directly translate into changes in saccade amplitude.
The improvement of motor behavior, based on experience, is a form of learning that is critically dependent on the cerebellum. A well studied example of cerebellar motor learning is short-term saccadic adaptation (STSA). In STSA, information on saccadic errors is used to improve future saccades. The information optimizing saccade metrics is conveyed by Purkinje cells simple spikes (PC-SS) because they are the critical input to the premotor circuits for saccades. We recorded PC-SS of monkeys undergoing STSA to reveal the code used for improving behavior. We found that the discharge of individual PC-SS was unable to account for the behavioral changes. The PC-SS population burst (PB), however, exhibited changes that closely paralleled the qualitatively different changes of saccade kinematics associated with gain-increase and gain-decrease STSA, respectively. Gain-increase STSA, characterized by an increase in saccade duration, replicates the relationship between saccade duration and the end of the PB valid for unadapted saccades. In contrast, gain-decrease STSA, which sports normal saccade duration but reduced saccadic velocity, is characterized by a PB that ends well before the adapted saccade. This suggests that the duration of normal as well as gain-increased saccades is determined by appropriately setting the end of PB end. However, the duration of gain-decreased saccades is apparently not modified by the cerebellum because the PB signals ends too early to determine saccade end. In summary, STSA, and most probably cerebellar-dependent learning in general, is based on optimizing the shape of a PC-SS population response.
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