Cortical neurons exhibit tremendous variability in the number and temporal distribution of spikes in their discharge patterns. Furthermore, this variability appears to be conserved over large regions of the cerebral cortex, suggesting that it is neither reduced nor expanded from stage to stage within a processing pathway. To investigate the principles underlying such statistical homogeneity, we have analyzed a model of synaptic integration incorporating a highly simplified integrate and fire mechanism with decay. We analyzed a “high-input regime” in which neurons receive hundreds of excitatory synaptic inputs during each interspike interval. To produce a graded response in this regime, the neuron must balance excitation with inhibition. We find that a simple integrate and fire mechanism with balanced excitation and inhibition produces a highly variable interspike interval, consistent with experimental data. Detailed information about the temporal pattern of synaptic inputs cannot be recovered from the pattern of output spikes, and we infer that cortical neurons are unlikely to transmit information in the temporal pattern of spike discharge. Rather, we suggest that quantities are represented as rate codes in ensembles of 50‐ 100 neurons. These column-like ensembles tolerate large fractions of common synaptic input and yet covary only weakly in their spike discharge. We find that an ensemble of 100 neurons provides a reliable estimate of rate in just one interspike interval (10‐50 msec). Finally, we derived an expression for the variance of the neural spike count that leads to a stable propagation of signal and noise in networks of neurons—that is, conditions that do not impose an accumulation or diminution of noise. The solution implies that single neurons perform simple algebra resembling averaging, and that more sophisticated computations arise by virtue of the anatomical convergence of novel combinations of inputs to the cortical column from external sources.

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The variable discharge of cortical neurons: implications for connectivity, computation, and information coding

http://yael-nissan.weebly.com/uploads/5/2/9/2/5292348/the_introduction_to_pain_an_integrative_review_-_millan.pdf

The induction of pain: an integrative review

How Inhibition Shapes Cortical Activity

GABAergic Interneurons in the Neocortex: From Cellular Properties to Circuits

https://www.shadlenlab.columbia.edu/publications/publications/mike/shadlen_newsome_Noise94.pdf

Noise, neural codes and cortical organization

Blindfolded rats were trained to stretch across a gap to palpate rough or smooth surfaces with their mystacial vibrissae. Animals learned to discriminate reliably a smooth surface from a rough surface having shallow (approximately 30 microns) grooves spaced at 90 microns intervals. Field-by-field video analyses confirmed that rats used only their vibrissae to contact the discriminanda. The whiskers swept across the surfaces at 696 degrees/sec during forward movements and 1106 degrees/sec for retracting movements. Mean amplitudes, which were 32 degrees, were considerably smaller than the total arc through which whiskers can move. Rats maintained whisker contact with discriminanda for several hundreds of msec, during which time the animals repetitively swept their vibrissae across the surface at a dominant frequency of 8 Hz. The range extended from 1 to 20 Hz, and the frequencies utilized varied within and among subjects. Whiskers contacted the discriminanda along the hair shaft, not at the whisker tips. The hair shafts were bent continually but to varying degrees as an animal palpated the surface, and more than one of the large caudal whiskers were almost always in contact with it. Thus, whiskers are not used independently as rigid levers. Results indicate that the capacity of the rodent whisker system to distinguish a smooth surface from a rough one is comparable to that of primates using their fingertips and suggest common strategies for active touch in the mammalian somatomotor system.

/pdf/biometric-analyses-of-vibrissal-tactile-discrimination-in-11mvpm7c9r.pdf

Biometric analyses of vibrissal tactile discrimination in the rat

1 Extracellular single-unit recordings and controlled whisker stimuli were used to compare response properties between cells in the "barreloids" of the thalamic ventrobasal complex and those in th

Thalamocortical response transformation in the rat vibrissa/barrel system.

Axonal tracing techniques were used to examine the distribution of corticothalamic projection neurons in relation to the organization of the thalamocortical recipient zones in the whisker representation of the rat first somatic sensory cortex. Following injection of horseradish peroxidase into the physiologically defined vibrissa area in the ventrobasal complex of the thalamus, labeling in the cortex had a columnar appearance. Dense patches of anterograde labeling were located within the centers of the layer IV barrels and extended superficially through lamina III; the septa between barrels contained considerably less reaction product. Retrogradely labeled neurons were observed in lower layer V and layer VI where they were concentrated preferentially deep to the barrel centers. Regions deep to the septa displayed less overall labeling and a lower relative number of thalamic projecting neurons. Zones having the larger numbers of retrogradely labeled cells also contained terminallike labeling of either corticothalamic or thalamocortical origin. Following an injection that included the posterior group medial to the ventrobasal complex, anterograde labeling in layer IV was located largely in the septa. In conjunction with previous findings concerning the origin and termination of other projection systems in the barrel cortex, these results suggest that a vibrissal column contains a central core zone intimately linked with the ventrobasal thalamus that is bounded by narrower regions of more diverse inputs and outputs that form an interface between adjacent cortical columns.

Spatial organization of thalamocortical and corticothalamic projection systems in the rat SmI barrel cortex.

The response properties of 123 trigeminal ganglion neurons were studied, using controlled whisker deflections in different directions. When the distal end of the whisker was initially displaced 5.7° (1 mm) from its neutral position, 81% of the cells responded with statistically more spikes/ stimulus to movements in one to three of eight cardinal (45° increment) directions than to the others. The more directionally selective the cell, the more vigorous was its response. On the basis of statistical criteria, 75% of the cells were classified as slowly adapting, 25% as rapidly adapting. A number of quantitative analyses indicated that slowly adapting units respond more selectively than rapidly adapting cells to the direction of whisker movement. Differences in directional sensitivities of rapidly and slowly adapting cells appear to parallel differences between their putative mechanoreceptive endings and the relationships between those endings and the vibrissa follicle's structure. Comparisons between the resp...

Responses of rat trigeminal ganglion neurons to movements of vibrissae in different directions.

Rats explore objects by rhythmically whisking them with their mystacial vibrissae. On two types of tactile discrimination tasks, macrogeometric and microgeometric, better performers palpated the discrimnanda for longer periods of time and used movement patterns that appeared to optimize whisking frequency bandwidth and the extent to which the vibrissae would be bent by object contact. On a task involving finely textured surfaces, good and poor performers differed in the temporal components of their whisking patterns, whereas the spatial domain was more important for animals palpating surfaces with widely separated features. These findings are consistent with increasing neurophysiological evidence that the central representation of the tactile periphery, in rodents and other mammals, is both integrative and dynamic.

George E. Carvell

Papers

Biometric analyses of vibrissal tactile discrimination in the rat

Thalamocortical response transformation in the rat vibrissa/barrel system.

Spatial organization of thalamocortical and corticothalamic projection systems in the rat SmI barrel cortex.

Responses of rat trigeminal ganglion neurons to movements of vibrissae in different directions.

Task- and Subject-Related Differences in Sensorimotor Behavior during Active Touch