Most strabismic observers do not suffer from double vision because of suppression from conscious perception of 1 of the 2 eyes' conflicting views. Direct evidence for the site and neural substrate of strabismic suppression has not been available so far, although psychophysical data suggest a cortical origin. On the other hand, cross-orientation suppression among conflicting stimuli presented monocularly has recently been shown to have a strong thalamic component. Here we present evidence, using both visual stimulation and pharmacological techniques, that strabismic suppression occurs in the primary visual cortex and involves gamma-amino butyric acid (GABA)-mediated inhibition. We show that its dependency on the drift rate of the suppressing stimulus is consistent with a cortical origin; unlike monocular cross-orientation suppression, it cannot be evoked by very fast-moving stimuli. Furthermore, strabismic suppression is greatly reduced when GABAergic inhibition is locally blocked by the GABA(A) antagonist bicuculline.
Receptive fields (RFs) in the visual cortex are characterized by spatiotemporal profiles that have been described in detail for area 17 simple cells. In this study, we analyse spatial and temporal RF properties of simple and complex cells in layer II/III of area 18 of the anaesthetized adult cat, using the reverse correlation method with brief 50 ms presentations of flashing bright and dark bars. Stimuli were presented with preferred orientation as previously determined by moving bars. Simple cell RFs were characterized by spatially and temporally separable ON and OFF subfields, while in complex cells ON and OFF subfields were superimposed. To discriminate possible contributions of GABAergic inhibition to RF structure and response dynamics in area 18, we have used three-barrelled micropipettes for single cell recordings and microiontophoresis, and have documented ON and OFF responses before, during and after application of bicuculline methiodide for blockade of GABAA receptors. During blockade of GABAergic inhibition, the stimulus-induced and resting discharge frequency increased, and in about 50% of the cells both ON and OFF subfields changed significantly in space and/or time in a reversible manner. In space, blockade of inhibition widened RF subfields, whereas in time, it shortened the duration of the excitatory cell response in simple and complex cells. ON and OFF subfields separated in space and time (simple cells), or time (complex cells) became less isolated or even superimposed. The results indicate substantial local inhibitory processing contributing to spatiotemporal RF properties in layers II/III of area 18 of the cat.
The role of GABAergic inhibition in orientation and direction selectivity has been investigated with the GABA(A)-Blocker bicuculline in the cat visual cortex, and results indicated a region specific difference of functional contributions of GABAergic inhibition in areas 17 and 18. In area 17 inhibition appeared mainly involved in sculpturing orientation and direction tuning, while in area 18 inhibition seemed more closely associated with temporal receptive field properties. However, different types of stimuli were used to test areas 17 and 18 and further studies performed in area 17 suggested an important influence of the stimulus type (single light bars vs. moving gratings) on the evoked responses (transient vs. sustained) and inhibitory mechanisms (GABA(A) vs. GABA(B)) which in turn might be more decisive for the specific results than the cortical region. To insert the missing link in this chain of arguments it was necessary to study GABAergic inhibition in area 18 with moving light bars, which has not been done so far. Therefore, in the present study we investigated area 18 cells responding to oriented moving light bars with extracellular recordings and reversible microiontophoretic blockade of GABAergig inhibition with bicuculline methiodide. The majority of neurons was characterized by a pronounced orientation specificity and variable degrees of direction selectivity. GABA(A)ergic inhibition significantly influenced preferred orientation and preferred direction in area 18. During the action of bicuculline orientation tuning width increased and orientation and direction selectivity indices decreased. Our results obtained in area 18 with moving bar stimuli, although in the proportion of affected cells similar to those described in area 17, quantitatively matched the findings for direction and orientation specificity obtained with moving gratings in area 18. Accordingly, stimulus type is not decisive in area 18 and the GABA(A) dependent, inhibitory intracortical computations involved in orientation specificity are indeed region-specific and in comparison to area 17 less effective in area 18.
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