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Daniel A Pollen

Publications and source records attributed to Daniel A Pollen.

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Brain stimulation and conscious experience: electrical stimulation of the cortical surface at a threshold current evokes sustained neuronal activity only after a prolonged latency.

Libet demonstrated that a substantial duration (>0.5-1.0 s) of direct electrical stimulation of the surface of a sensory cortex at a threshold or liminal current is required before a subject can experience a percept. Libet and his co-workers originally proposed that the result could be due either to spatial and temporal facilitation of the underlying neurons or additionally to a prolonged central processing time. However, over the next four decades, Libet chose to attribute the prolonged latency for evoking conscious experience to a prolonged central processing time. In my view, Libet has not provided any convincing evidence, either on the basis of his own past work or in his critique of my paper, for his hypothesis of a central processing time exceeding 0.5s before conscious experience emerges following direct electrical threshold stimulation of the cortical surface. I stand by my original results and conclusion that such prolonged latencies are largely the consequence of a dynamically increasing cortical facilitatory process that begins hundreds of milliseconds before there is a sustained neuronal activation. In some cases, the facilitatory process must overcome an initial stimulus-induced inhibition before neuronal firing commences.

Consciousness↗

Brain stimulation and conscious experience.

Libet discovered that a substantial duration (> 0.5-1.0 s) of direct electrical stimulation of the surface of the somatosensory cortex at threshold currents is required before human subjects can report that a conscious somatosensory experience had occurred. Using a reaction time method we confirm that a similarly long stimulation duration at threshold currents is required for activation of elementary visual experiences (phosphenes) in human subjects following stimulation of the surface of the striate cortex. However, the reaction times for the subject to respond to the cessation of the visual experience after the end of electrical stimulation could be as brief as 225-242 ms. We also carried out extensive studies in cats under a variety of anesthetic conditions using the same electrodes and parameters of stimulation employed in the human studies to study the patterns of neuronal activity beneath the stimulating surface electrode. Whereas sufficiently strong currents can activate neurons within milliseconds, stimulating currents close to threshold activate sustained neural activity only after at least 350-500 ms. When currents are close to threshold, some neurons are inhibited for several hundreds of millisecond before the balance between inhibition and excitation shifts towards excitation. These results suggest that the prolonged latencies, i.e., latencies beyond 200-250 ms, for the emergence of conscious experience following direct cortical stimulation result from a delay in the sustained activation of underlying cortical neurons at threshold currents rather than being due to any unusually long duration in central processing time. Intracellular records from cortical neurological cells during repetitive electrical stimulation of the surface of the feline striate cortex demonstrate that such stimulation induces a profound depolarizing shift in membrane potential that may persist after each stimulus train. Such a depolarization is evidence that extracellular K+ concentrations have increased during electrical stimulation. Such an increase in extracellular K+ progressively increases cortical excitability until the threshold for sustained activation of cortical neurons is reached and then exceeded. Consequently, the long latency for threshold activation of cortical neurons depends upon a dynamically increasing cortical facilatory process that begins hundreds of milliseconds before there is sustained activation of such neurons. In some cases, this facilatory process must overcome an initial stimulus-induced inhibition before neuronal firing commences.

Adult↗

Explicit neural representations, recursive neural networks and conscious visual perception.

The fundamental question as to whether the neural correlates of any given conscious visual experience are expressed locally within a given cortical area or more globally within some widely distributed network remains unresolved. We inquire as to whether recursive processing-by which we mean the combined flow and integrated outcome of afferent and recurrent activity across a series of cortical areas-is essential for the emergence of conscious visual experience. If so, we further inquire as to whether such recursive processing is essential only for loops between extrastriate cortical areas explicitly representing experiences such as color or motion back to V1 or whether it is processing between still higher levels and the areas computing such explicit representations that is exclusively or additionally essential for visual experience. If recursive processing is not essential for the emergence of conscious visual experience, then it should also be possible to determine whether it is only the intracortical sensory processing within areas computing explicit sensory representations that is required for perceptual experience or whether it is the subsequent processing of the output of such areas within more anterior cortical regions that engenders perception. The present analysis suggests that the questions posed here may ultimately become experimentally resolvable. Whatever the outcome, the results will likely open new approaches to identify the neural correlates of conscious visual perception.

Animals↗

Spatial receptive field organization of macaque V4 neurons.

Subfield analysis of the receptive fields (RFs) of parafoveal V4 complex cells demonstrates directly that most RFs are tiled by overlapping second-order excitatory inputs that for any given V4 cell are predominantly selective to the same preferred values of spatial frequency and orientation. These results extend hierarchical principles of RF organization in the spatial, orientation and spatial frequency domains, first recognized in V1, to an intermediate extrastriate cortex. Spatial interaction studies across subfields demonstrate that the responses of V4 neurons to paired stimuli may either decrease or increase as a function of inter-stimulus distance across the width axis. These intra-RF suppressions and facilitations vary independently in magnitude and spatial extent from cell to cell. These results taken together with the relatively large RF sizes of V4 neurons - as compared with RF sizes of their afferent inputs - lead us to hypothesize a novel property, namely that classes of stimulus configurations that enhance areal summation while reducing suppressive interactions between excitatory inputs will evoke especially robust responses. We tested, and found support for, this hypothesis by presenting stimuli consisting of optimally tuned sine-wave gratings visible only within an annular region and found that such stimuli vigorously activate V4 neurons at firing rates far higher than those evoked by comparable stimuli to either the full-field or central core. On the basis of these results we propose a framework for a new class of neural network models for the spatial RF organizations of prototypic V4 neurons.

Animals↗