Processing of transient stimuli by the visual system of the rat
Master Thesis
1993
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University of Cape Town
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While three decades of intensive cortical electrophysiology using a variety of sustained visual stimuli has made a significant contribution to many aspects of visual function, it has not supported the existence of intracortical circuit operations in cortical processing. This study investigated cortical processing by a comparison of the response of primary visual cortical neurones to transient electrical and strobe-flash stimulation. Experiments were performed on 74 anaesthetised Long Evans rats. Standard stereotaxic and extracellular electrophysiological techniques were employed. Continuous (on-line) raster plots and peri-stimulus time histograms (PSTHs) of the extracellular spikes from 81 visual cortical and 55 lateral geniculate nucleus (LGN) neurones were compiled. The strobe-flash stimuli (0.05 ms) were applied to the contralateral eye while the monopolar or bipolar electrical stimuli (0.2 ms, 80-400 μA) were applied to the ipsilateral LGN. 60 of the 81 (74%) tested cortical units were found to be responsive to visual stimuli. A distinct and consistent difference in the cortical response to the two types of transient stimuli was found: (a) Electrical stimulation evoked a prolonged period (197 ± 61 ms) of inhibition in all cortical neurones tested (n=20). This was the case even in those cortical units that were completely unresponsive to visual stimulation. The protracted inhibition was usually followed by a 100-200 ms phase of rebound excitation. (b) Flash stimulation evoked a prominent excitatory discharge (5-30 ms duration) after a latency of 30-60 ms from the onset of the stimulus (n = 59). This was followed by either moderate inhibition or return to a firing rate similar to control activity, for a maximum of 40 ms. Thereafter, cortical neurones showed a sustained increased level of activity with superimposed secondary excitatory phases. The duration of this late re-excitatory phase was 200-300 ms. In 17 of 20 (85%) tested units, the temporal profile of the cortical response to flash stimulation was modulated by small changes in the level of background illumination. In 16 of the 17 units, this sensitivity was reflected primarily as an emergence of a brief secondary inhibitory phase at the lowest level of background illumination (0 lux). Only 1 of the 17 cortical units displayed a flash-evoked primary inhibitory phase at O lux. We explored the possibility that neurones in the lateral geniculate nucleus (LGN) of the thalamus were responsible for the late phase of cortical reexcitation. 49 of the 55 (89%) LGN neurones could be classified as either of the "ON type" i.e. excited by visual stimuli, or the "OFF type" i.e. inhibited by visual stimuli. The response of ON-like LGN neurones to strobe-flash stimulation of the contralateral eye was characterised by a primary excitatory or early discharge (ED) phase after a latency of 25-40 ms. Thereafter, a 200- 400 ms period of inhibition was observed. In 57% of the sample, a rebound excitatory or late discharge (LD) phase completed the response. OFF-like LGN neurones were inhibited by the strobe-flash stimuli after a latency of 30- 35 ms. This flash-evoked inhibition was maintained for 200-400 ms. The sensitivity of the flash-evoked LGN response to the level of background illumination was tested in 11 ON-like and 10 OFF-like neurones. No sustained secondary excitatory events, as observed in visual cortical neurones, were found in any of the ON- and OFF-like LGN neurones, irrespective of the level of background illumination. In conclusion, the data show that the late re-excitatory phase evoked in cortical neurones upon strobe-flash stimulation, is not due to sustained LGN (thalamic) input. Rather, it suggests that these re-excitatory phases are due to intracortical processing of the transient stimuli. These findings emphasize the independent role of the cortex in computing the response to visual stimuli, and cast doubt on traditional theories that have emphasised the role of the thalamus in shaping cortical responses. The difference in the flash and electrically evoked cortical response suggests that even though substantial inhibition is available to the cortex, only a small fraction of this inhibitory capacity is utilised during natural stimulation.
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Kara, P. 1993. Processing of transient stimuli by the visual system of the rat. University of Cape Town.