Cortical OFF Research Articles

More than the end: OFF response plasticity as a mnemonic signature of a sound's behavioral salience.

In studying how neural populations in sensory cortex code dynamically varying stimuli to guide behavior, the role of spiking after stimuli have ended has been underappreciated. This is despite growing evidence that such activity can be tuned, experience-and context-dependent and necessary for sensory decisions that play out on a slower timescale. Here we review recent studies, focusing on the auditory modality, demonstrating that this so-called OFF activity can have a more complex temporal structure than the purely phasic firing that has often been interpreted as just marking the end of stimuli. While diverse and still incompletely understood mechanisms are likely involved in generating phasic and tonic OFF firing, more studies point to the continuing post-stimulus activity serving a short-term, stimulus-specific mnemonic function that is enhanced when the stimuli are particularly salient. We summarize these results with a conceptual model highlighting how more neurons within the auditory cortical population fire for longer duration after a sound’s termination during an active behavior and can continue to do so even while passively listening to behaviorally salient stimuli. Overall, these studies increasingly suggest that tonic auditory cortical OFF activity holds an echoic memory of specific, salient sounds to guide behavioral decisions.

A generic deviance detection principle for cortical On/Off responses, omission response, and mismatch negativity

Neural responses to sudden changes can be observed in many parts of the sensory pathways at different organizational levels. For example, deviants that violate regularity at various levels of abstraction can be observed as simple On/Off responses of individual neurons or as cumulative responses of neural populations. The cortical deviance-related responses supporting different functionalities (e.g., gap detection, chunking, etc.) seem unlikely to arise from different function-specific neural circuits, given the relatively uniform and self-similar wiring patterns across cortical areas and spatial scales. Additionally, reciprocal wiring patterns (with heterogeneous combinations of excitatory and inhibitory connections) in the cortex naturally speak in favor of a generic deviance detection principle. Based on this concept, we propose a network model consisting of reciprocally coupled neural masses as a blueprint of a universal change detector. Simulation examples reproduce properties of cortical deviance-related responses including the On/Off responses, the omitted-stimulus response (OSR), and the mismatch negativity (MMN). We propose that the emergence of change detectors relies on the involvement of disinhibition. An analysis of network connection settings further suggests a supportive effect of synaptic adaptation and a destructive effect of N-methyl-d-aspartate receptor (NMDA-r) antagonists on change detection. We conclude that the nature of cortical reciprocal wiring gives rise to a whole range of local change detectors supporting the notion of a generic deviance detection principle. Several testable predictions are provided based on the network model. Notably, we predict that the NMDA-r antagonists would generally dampen the cortical Off response, the cortical OSR, and the MMN.

Cortical Balance Between ON and OFF Visual Responses Is Modulated by the Spatial Properties of the Visual Stimulus.

The primary visual cortex of carnivores and primates is dominated by the OFF visual pathway and responds more strongly to dark than light stimuli. Here, we demonstrate that this cortical OFF dominance is modulated by the size and spatial frequency of the stimulus in awake primates and we uncover a main neuronal mechanism underlying this modulation. We show that large grating patterns with low spatial frequencies drive five times more OFF-dominated than ON-dominated neurons, but this pronounced cortical OFF dominance is strongly reduced when the grating size decreases and the spatial frequency increases, as when the stimulus moves away from the observer. We demonstrate that the reduction in cortical OFF dominance is not caused by a selective reduction of visual responses in OFF-dominated neurons but by a change in the ON/OFF response balance of neurons with diverse receptive field properties that can be ON or OFF dominated, simple, or complex. We conclude that cortical OFF dominance is continuously adjusted by a neuronal mechanism that modulates ON/OFF response balance in multiple cortical neurons when the spatial properties of the visual stimulus change with viewing distance and/or optical blur.

Neuronal and perceptual differences in the temporal processing of darks and lights.

Visual information is mediated by two major thalamic pathways that signal light decrements (OFF) and increments (ON) in visual scenes, the OFF pathway being faster than the ON. Here, we demonstrate that this OFF temporal advantage is transferred to visual cortex and has a correlate in human perception. OFF-dominated cortical neurons in cats responded ∼3 ms faster to visual stimuli than ON-dominated cortical neurons, and dark-mediated suppression in ON-dominated neurons peaked ∼14 ms faster than light-mediated suppression in OFF-dominated neurons. Consistent with the neuronal differences, human observers were 6-14 ms faster at detecting darks than lights and better at discriminating dark than light flickers. Neuronal and perceptual differences both vanished if backgrounds were biased toward darks. Our results suggest that the cortical OFF pathway is faster than the ON pathway at increasing and suppressing visual responses, and these differences have parallels in the human visual perception of lights and darks.

Cortical Off Response Tuning for Stimulus Duration

To answer the question whether OFF response amplitude (firing rate) of visual cortical cells varies as a function of stimulus duration, a series of such cells from areas 17 and 18 of the cat were investigated with a stationary light bar, presented for different durations (10–3200 msec) over the receptive field. Out of a sample of 174 cells tested, 58 cells were found for which the OFF responses varied as a function of ON stimulus duration. Of these cells, 29 showed a continuous increase up to the longest duration tested, six showed a sharp tuning for medium range durations (between 50 and 400 msec) and the remaining 23 cells had an intermediate profile (increase to optimum followed by slight decrease). For some of these cells this tuning could be used to predict adequately the velocity tuning. Similar recordings in 13 lateral geniculate nucleus cells failed to show duration tuning. Copyright © 1996 Elsevier Science Ltd.