Primary visual coding can be characterized by the receptive field (RF) properties of single neurons. Subject of this paper is our search for a global, second coding step beyond the RF-concept that links related features in a visual scene. In recent models of visual coding, oscillatory activities have been proposed to constitute such linking signals. We tested the neurophysiological relevance of this hypothesis for the visual system. Single and multiple spikes as well as local field potentials were recorded simultaneously from several locations in the primary visual cortex (A17 and A18) using 7 or 19 individually advanceable fiber-microelectrodes (250 or 330 microns apart). Stimulus-evoked (SE)-resonances of 35-85 Hz were found in these three types of signals throughout the visual cortex when the primary coding channels were activated by their specific stimuli. Stimulus position, orientation, movement direction and velocity, ocularity and stationary flicker caused specific SE-resonances. Coherent SE-resonances were found at distant cortical positions when at least one of the primary coding properties was similar. Coherence was found 1) within a vertical cortex column, 2) between neighbouring hypercolumns, and 3) between two different cortical areas. We assume that the coherence of SE-resonances is mediated by recurrent excitatory intra- and inter-areal connections via phase locking between assemblies that represent the linking features of the actual visual scene. Visually related activities are, thus, transiently labelled by a temporal code that signalizes their momentary association.

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.

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Event-related potentials (ERPs) to changes in the visual environment were recorded in rabbits. In the oddball condition, infrequently presented (deviant) stimuli occurred in a series of frequently presented (standard) stimuli. In the deviant-alone condition, standards were omitted. ERPs to oddball-deviants differed from those to standards in all recording sites (cerebellar cortex, visual cortex, dentate gyrus). No corresponding differences were found between ERPs to deviants in the oddball condition and those in the deviant-alone condition. However, because ERPs to deviants in the deviant-alone condition and those to standards did not differ either, ERPs to stimulus changes in the oddball condition seemed to be dependent on the presence of standards, thus representing an analogue to mismatch negativity (MMN) in humans.