Receptive Fields Of Cortical Neurons Research Articles

Mir-132/212 is required for maturation of binocular matching of orientation preference and depth perception

MicroRNAs (miRNAs) are known to mediate post-transcriptional gene regulation, but their role in postnatal brain development is still poorly explored. We show that the expression of many miRNAs is dramatically regulated during functional maturation of the mouse visual cortex with miR-132/212 family being one of the top upregulated miRNAs. Age-downregulated transcripts are significantly enriched in miR-132/miR-212 putative targets and in genes upregulated in miR-132/212 null mice. At a functional level, miR-132/212 deletion affects development of receptive fields of cortical neurons determining a specific impairment of binocular matching of orientation preference, but leaving orientation and direction selectivity unaltered. This deficit is associated with reduced depth perception in the visual cliff test. Deletion of miR-132/212 from forebrain excitatory neurons replicates the binocular matching deficits. Thus, miR-132/212 family shapes the age-dependent transcriptome of the visual cortex during a specific developmental window resulting in maturation of binocular cortical cells and depth perception.

Rapid Task-Related Plasticity of Spectrotemporal Receptive Fields in the Auditory Midbrain.

Previous research has demonstrated that auditory cortical neurons can modify their receptive fields when animals engage in auditory detection tasks. We tested for this form of task-related plasticity in the inferior colliculus (IC) of ferrets trained to detect a pure tone target in a sequence of noise distractors that did not overlap in time. During behavior, responses were suppressed at the target tone frequency in approximately half of IC neurons relative to the passive state. This suppression often resulted from a combination of a local tuning change and a global change in overall excitability. Local and global suppression were stronger when the target frequency was aligned to neuronal best frequency. Local suppression in the IC was indistinguishable from that described previously in auditory cortex, while global suppression was unique to the IC. The results demonstrate that engaging in an auditory task can change selectivity for task-relevant features in the midbrain, an area where these effects have not been reported previously. Significance statement: Previous studies have demonstrated that the receptive fields of cortical neurons are modified when animals engage in auditory behaviors, a process that is hypothesized to provide the basis for segregating sound sources in an auditory scene. This study demonstrates for the first time that receptive fields of neurons in the midbrain inferior colliculus are also modified during behavior. The magnitude of the tuning changes is similar to previous reports in cortex. These results support a hierarchical model of behaviorally driven sound segregation that begins in the subcortical auditory network.

Sperm donor pool shrivels when payments cease

The thalamus serves as a gate that regulates the flow of sensory inputs to the neocortex, and this gate is controlled by neuromodulators from the brainstem reticular formation that are released during arousal. We found recently that sensory-evoked responses are suppressed in the neocortex during arousal. This sensory suppression results from the activity-dependent depression of the thalamocortical connection caused by increased tonic firing of thalamocortical cells during arousal. In the present study, the functional consequences of thalamocortical suppression during arousal were investigated using the vibrissae system of rodents. The results show that thalamocortical suppression is associated with a strong reduction in the spread of sensory inputs through the cortex, thus reducing the size of sensory representations. In addition, when the responses of single cells to principal and adjacent whiskers are compared, the response to the adjacent whiskers was found to be strongly suppressed, much more so than that of principal whiskers. Consequently, the receptive fields of cortical neurons become more focused to the principal whisker. The results indicate that thalamocortical suppression during arousal serves to focus sensory inputs to their appropriate representations in neocortex, which may be computationally helpful for the spatial processing of sensory information.

Disruption of Balanced Cortical Excitation and Inhibition by Acoustic Trauma

Sensory deafferentation results in rapid shifts in the receptive fields of cortical neurons, but the synaptic mechanisms underlying these changes remain unknown. The rapidity of these shifts has led to the suggestion that subthreshold inputs may be unmasked by a selective loss of inhibition. To study this, we used in vivo whole cell recordings to directly measure tone-evoked excitatory and inhibitory synaptic inputs in auditory cortical neurons before and after acoustic trauma. Here we report that acute acoustic trauma disrupted the balance of excitation and inhibition by selectively increasing and reducing the strength of inhibition at different positions within the receptive field. Inhibition was abolished for frequencies far below the trauma-tone frequency but was markedly enhanced near the edges of the region of elevated peripheral threshold. These changes occurred for relatively high-level tones. These changes in inhibition led to an expansion of receptive fields but not by a simple unmasking process. Rather, membrane potential responses were delayed and prolonged throughout the receptive field by distinct interactions between synaptic excitation and inhibition. Far below the trauma-tone frequency, decreased inhibition combined with prolonged excitation led to increased responses. Near the edges of the region of elevated peripheral threshold, increased inhibition served to delay rather than abolish responses, which were driven by prolonged excitation. These results show that the rapid receptive field shifts caused by acoustic trauma are caused by distinct mechanisms at different positions within the receptive field, which depend on differential disruption of excitation and inhibition.

Effects of spatial separation between stimuli in whole report from brief visual displays

Direct measurements of the effects of spatial separation between stimuli in whole report from brief visual displays are reported. The stimuli were presented on the periphery of an imaginary circle centered on fixation. In Experiment 1, each display showed two capital letters (letter height approximately equal 1.3 degrees, width approximately equal 0.9 degree, eccentricity approximately equal 5.5 degrees). The proportion of correctly reported letters was a strictly increasing, decelerating function of the spatial separation between the letters for center-to-center separations ranging from less than 2 degrees to more than 10 degrees of visual angle. Experiment 2 yielded similar results with triples of letters. Experiment 3 showed that accuracy increased with spatial separation for report of two short words, and Experiment 4 showed the same result for words presented upside down. The results are explained by a model of lateral masking (crowding) based on competitive interactions within receptive fields of cortical neurons.

Responses of Neurons in Primary Visual Cortex to Transient Changes in Local Contrast and Luminance

During normal saccadic inspection of natural images, the receptive fields of cortical neurons are bombarded with frequent simultaneous changes in local mean luminance and contrast, yet there have been no systematic studies of how cortical neurons respond to such stimulation. The responses of single neurons in the primary visual cortex of the cat were measured for 200 ms presentations of sine-wave gratings confined to the conventional receptive field. Both local mean luminance and contrast were parametrically and randomly varied over the 1-1.5 log unit ranges that are typical of natural images. We find that responses are strongly modulated by both the local mean luminance and contrast, but in an approximately separable manner: the contrast response function is approximately invariant except for a scale factor that depends on the local mean luminance. The shape of the temporal response profiles were found to be approximately invariant with contrast, but were strongly affected by the local mean luminance. The results suggest that most, if not all, cortical neurons carry substantial local luminance information.

Auditory cortical plasticity: Does it provide evidence for cognitive processing in the auditory cortex?

The past 20 years have seen substantial changes in our view of the nature of the processing carried out in auditory cortex. Some processing of a cognitive nature, previously attributed to higher-order “association” areas, is now considered to take place in auditory cortex itself. One argument adduced in support of this view is the evidence indicating a remarkable degree of plasticity in the auditory cortex of adult animals. Such plasticity has been demonstrated in a wide range of paradigms, in which auditory input or the behavioural significance of particular inputs is manipulated. Changes over the same time period in our conceptualization of the receptive fields of cortical neurons, and well-established mechanisms for use-related changes in synaptic function, can account for many forms of auditory cortical plasticity. On the basis of a review of auditory cortical plasticity and its probable mechanisms, it is argued that only plasticity associated with learning tasks provides a strong case for cognitive processing in auditory cortex. Even in this case the evidence is indirect, in that it has not yet been established that the changes in auditory cortex are necessary for behavioural learning and memory. Although other lines of evidence provide convincing support for cognitive processing in auditory cortex, that provided by auditory cortical plasticity remains equivocal.

The Role of Feedback in Shaping the Extra-Classical Receptive Field of Cortical Neurons: A Recurrent Network Model

The responses of neurons in sensory cortices are affected by the spatial context within which stimuli are embedded. In the primary visual cortex (V1), orientation-selective responses to stimuli in the receptive field (RF) center are suppressed by similarly oriented stimuli in the RF surround. Surround suppression, a likely neural correlate of perceptual figure-ground segregation, is traditionally thought to be generated within V1 by long-range horizontal connections. Recently however, it has been shown that these connections are too short and too slow to mediate fast suppression from distant regions of the RF surround. We use an anatomically and physiologically constrained recurrent network model of macaque V1 to show how interareal feedback connections, which are faster and longer-range than horizontal connections, can generate "far" surround suppression. We provide a novel solution to the puzzle of how surround suppression can arise from excitatory feedback axons contacting predominantly excitatory neurons in V1. The basic mechanism involves divergent feedback connections from the far surround targeting pyramidal neurons sending monosynaptic horizontal connections to excitatory and inhibitory neurons in the RF center. One of several predictions of our model is that the "suppressive far surround" is not always suppressive, but can facilitate the response of the RF center, depending on the amount of excitatory drive to the local inhibitors. Our model provides a general mechanism of how top-down feedback signals directly contribute to generating cortical neuron responses to simple sensory stimuli.

The influence of cortical feature maps on the encoding of the orientation of a short line

The inhomogeneous distribution of the receptive fields of cortical neurons influences the cortical representation of the orientation of short lines seen in visual images. We construct a model of the response of populations of neurons in the human primary visual cortex by combining realistic response properties of individual neurons and cortical maps of orientation and location preferences. The encoding error, which characterizes the difference between the parameters of a visual stimulus and their cortical representation, is calculated using Fisher information as the square root of the variance of a statistically efficient estimator. The error of encoding orientation varies considerably with the location and orientation of the short line stimulus as modulated by the underlying orientation preference map. The average encoding error depends only weakly on the structure of the orientation preference map and is much smaller than the human error of estimating orientation measured psychophysically. From this comparison we conclude that the actual mechanism of orientation perception does not make efficient use of all the information available in the neuronal responses and that it is the decoding of visual information from neuronal responses that limits psychophysical performance.

Human sound localization at near-threshold levels

Physiological studies of spatial hearing show that the spatial receptive fields of cortical neurons typically are narrow at near-threshold levels, broadening at moderate levels. The apparent loss of neuronal spatial selectivity at increasing sound levels conflicts with the accurate performance of human subjects localizing at moderate sound levels. In the present study, human sound localization was evaluated across a wide range of sensation levels, extending down to the detection threshold. Listeners reported whether they heard each target sound and, if the target was audible, turned their heads to face the apparent source direction. Head orientation was tracked electromagnetically. At near-threshold levels, the lateral (left/right) components of responses were highly variable and slightly biased towards the midline, and front vertical components consistently exhibited a strong bias towards the horizontal plane. Stimulus levels were specified relative to the detection threshold for a front-positioned source, so low-level rear targets often were inaudible. As the sound level increased, first lateral and then vertical localization neared asymptotic levels. The improvement of localization over a range of increasing levels, in which neural spatial receptive fields presumably are broadening, indicates that sound localization does not depend on narrow spatial receptive fields of cortical neurons.

Role of Thalamocortical Sensory Suppression during Arousal: Focusing Sensory Inputs in Neocortex

The thalamus serves as a gate that regulates the flow of sensory inputs to the neocortex, and this gate is controlled by neuromodulators from the brainstem reticular formation that are released during arousal. We found recently that sensory-evoked responses are suppressed in the neocortex during arousal. This sensory suppression results from the activity-dependent depression of the thalamocortical connection caused by increased tonic firing of thalamocortical cells during arousal. In the present study, the functional consequences of thalamocortical suppression during arousal were investigated using the vibrissae system of rodents. The results show that thalamocortical suppression is associated with a strong reduction in the spread of sensory inputs through the cortex, thus reducing the size of sensory representations. In addition, when the responses of single cells to principal and adjacent whiskers are compared, the response to the adjacent whiskers was found to be strongly suppressed, much more so than that of principal whiskers. Consequently, the receptive fields of cortical neurons become more focused to the principal whisker. The results indicate that thalamocortical suppression during arousal serves to focus sensory inputs to their appropriate representations in neocortex, which may be computationally helpful for the spatial processing of sensory information.

A structure preserving image transformation as the goal of visual sensory coding

Neuronal selectivity is often characterized by stimuli that evoke maximal responses. But early in the visual system neurons are known to respond to a richer class of stimuli than it can be expected by a feed-forward architecture. To explain this finding we present a more general functional framework for the exploration of neuronal responses. To quantify the structure or ‘interestingness’ of an image patch we introduce a measure of the symmetries that can be found in image patches at the size of local receptive fields of cortical neurons. We show how this transformation can explain strong responses to complex stimuli as well as responses to simple grating stimuli as found for neurons in V1 and V2.

Corticoreticular pathways in the cat. I. Projection patterns and collaterization.

This paper summarizes and compares the projection patterns and the receptive fields of cortical neurons in areas 4 and 6 that project to the pontomedullary reticular formation (PMRF). A total of 326 neurons were recorded in area 4 and 129 in area 6 in four awake, unrestrained cats that were chronically implanted with arrays of electrodes in the PMRF and the pyramidal tract (PT). In area 4, 47% of the neurons projected to the caudal PT but not to the PMRF (PTNs); 19% were activated only from the PMRF [corticoreticular neurons (CRNs)], whereas 27% were activated from both the PT and the PMRF (PTN/CRNs). More PTN/CRNs conducted at velocities >20 m/s (82%) than did CRNs (23%). In area 6, only 19% of the neurons were identified as PTNs, 12% were PTN/CRNs and 31% were CRNs; a further 38% could not be activated from either structure. Collateral branches within the PMRF conducted at maximum velocities of 20 m/s (average = 6.5 m/s). No significant differences in the conduction velocities of the collateral branches were found either between fast and slow PTNs or between area 4 and area 6 neurons. A large proportion of neurons in area 4 (85/173, 49%) were activated by passive manipulation of the more distal, contralateral forelimb, with approximately equal numbers being classed as PTNs, PTN/CRNs and CRNs. Most neurons in area 6 for which a receptive field could be found were excited by lightly touching or tapping the face and neck; a receptive field could not be determined for 39% of the area 6 neurons compared with only 5% of those in area 4. Finally, there was evidence that neurons in quite widespread areas of the pericruciate cortex, including both areas 4 and 6 projected onto similar, restricted regions of the PMRF. The fact that the cortical projection from area 4 to the PMRF includes a high percentage of fast PTNs with a receptive field on the distal forelimb is consistent with the view that this projection may serve to integrate movement and the dynamic postural adjustments that accompany them. The fact that the cortical projection from area 6 to the PMRF is primarily from slow PTNs with receptive fields on the face, neck and back is consistent with a role for this cortical area in adjusting the general posture of the animal on which movements are superimposed.

Progressive changes in cutaneous trigger zones for sensation referred to a phantom hand: a case report and review with implications for cortical reorganization

The dominant model of cortical plasticity induced by peripheral denervation suggests that a physiologically-reorganized cortical area can acquire new perceptual meaning, including a change in the peripheral referral of sensation. An alternative view is that such an area may retain its former perceptual significance, even though it becomes responsive to new peripheral inputs. To examine evidence related to this issue, a clinical case is presented documenting the time course of changes in phantom limb sensation in a patient with accidental amputation of a hand. About 24 h after injury, a vivid phantom hand was present; tactile stimulation revealed cutaneous trigger zones on the arm, stimulation of which elicited sensation referred to specific fingers of the phantom. While the phantom hand percept remained fairly stable over time, the trigger zones expanded progressively in size during the next 1-8 weeks but had contracted and changed location considerably about one year later. At all times studied, the trigger zones were topographically related to specific fingers and other parts of the phantom hand. The implications of these and other recent clinical findings for cortical reorganization are discussed, and the following tentative conclusions are drawn. (1) A phantom percept is mediated by central neural networks which remain functionally intact after amputation. (2) Cutaneous trigger zones mapped in humans correspond to novel receptive fields of cortical neurons mapped in animals following peripheral denervation. (3) Cortical reorganization induced by denervation does not produce a major change in perceptual meaning or peripheral reference. In the present case, stimulation of new trigger zones (receptive fields) on the patient's arm presumably activated a reorganized cortical hand area but evoked sensation still referred to the (now missing) hand. Hence, physiological cortical remapping is not necessarily accompanied by functional respecification.

An intracellular study of space and time representation in primary visual cortical receptive fields

In contrast with previous knowledge based on extracellular recordings, the recent development of intracellular techniques in vivo (sharp electrode or ‘blind patch’) ideally allows experimenters to analyze and dissect the contribution of feedforward and lateral connectivity in the functional expression of a synaptic ‘integration field’. We will present recent data which demonstrate that the visual receptive field of cortical neurons described at the level of subthreshold synaptic events extends over much larger regions of the visual field than previously thought, and that the capacity of cells to amplify subthreshold responses depends on the immediate past history of their membrane potential. Our data suggest that visual cortical receptive fields should not be considered as a fixed entity but more as a dynamic field of integration and association. Two types of dynamics can be argued for: 1) the spatial structure of the minimal discharge field (defined by suprathreshold activation of the cell) can be profoundly reorganized at least during development and most probably during selective phases of learning under the control of activity-dependent mechanisms. Adaptive changes in visual responses are thought to reflect long-lasting potentiation and/or depression of synaptic efficacies conveying ON- and OFF-center information; and 2) during sensory processing, reconfiguration of synaptic weights may be achieved on a much faster time-scale and linke to non-linear properties of the postsynaptic membrane as well as that of recruited networks. Association of information available in the central part of the receptive field (RF) and of input coming from the reputedly ‘unresponsive’ regions surrounding it, or arising simultaneously from different parts of the visual field, might be suppressive in certain cases and capable of boosting hidden responses in other cases, depending on the global stimulus configuration.

Backward conditioned connection and inhibitory reorganization of the receptive fields of cortical neurons as the basis of subconscious change in the thresholds of visual recognition and detection

Three groups of facts are compared in this study: the significant adaptive and adaptational modification of the receptive fields of neurons of the visual cortex of the cat, the conditioned, selective, subsensory change in the threshold of perception (detection and recognition) by an individual of a letter in relation to two control letters, and the role of spatially-specialized cortical inhibition in the formation and adaptive modifications of the receptive fields and detector properties of neurons of the visual cortex. The relationship of the described phenomena and the commonality of their mechanisms is discussed.

A computational model of the vertical anatomical organization of primary visual cortex

A method for modeling anatomical connectivity for a vertically organized slab of cortical tissue in mammalian primary visual cortex has been developed. The modeled slab covers 500 x 500 microns of cortical surface and extends vertically throughout the full depth of the cortex. The model slab was divided into 6 laminae and neuronal somata were distributed in three dimensions through the slab in accordance with experimentally derived cell densities. Axonal and dendritic arborizations were modeled as line segments. A total of 17 morphological types of neurons were included. Connectivity was established based on proximity between axonal and dendritic arbors. There is good general agreement between the vertical distribution of connections generated by the model and the vertical distribution of synapses observed for cat area 17. In all layers, fewer connections were generated in the model than synapses in cat area 17. This is due, at least in part, to the exclusion of long range intracortical projections and sources of afferent input other than the dorsal lateral geniculate nucleus from the model. The connection scheme described here will be used in conjunction with a physiology model to model vertical signal flow, and will be expanded further to model receptive fields of cortical neurons.

C-fibres provide a source of masking inhibition to primary somatosensory cortex

Capsaicin was applied to the exposed radial nerve of adult flying foxes (n = 5) and cats (n = 2) while recording in primary somatosensory cortex from a single neuron with a receptive field on digits 1 or 2. Within four minutes of application of capsaicin the borders of these receptive fields dramatically expanded. In a further four flying foxes it was shown, with subcutaneous delivery just proximal to the receptive fields, that capsaicin need affect only afferents from the region of a neuron's receptive field to induce expansion. Capsaicin applied directly to a nerve, or subcutaneously in high concentrations, is a selective neurotoxin that rapidly prevents the propagation of action potentials in most C-fibres. The result provides a partial explanation for experiments involving the specific and complete denervation of receptive fields of neurons in primary somatosensory cortex. Such denervation does not lead to unresponsiveness but to immediate sensitivity to stimulation of areas surrounding the original fields. Thus it appears that some subclass of capsaicin-sensitive C-fibres provides a primary source for the masking inhibition that normally limits the extent of the receptive fields of cortical neurons.

Dynamic stabilization of receptive fields of cortical neurons (VI) during fixation of gaze in the macaque

The positions of receptive field borders of striate cortical neurons were measured repeatedly in awake monkeys during attentive fixation of a small target. The border position, as marked by the onset of evoked activity in response to a moving stimulus, did not show the variability expected from previous measures of eye position variability during fixation. Measured variability was smaller than expected. Trial-by-trial comparisons suggest that receptive field borders are not shifted by the small eye movements occurring during attentive fixation. It is our hypothesis that attentive fixation engages a mechanism that gates incoming information to achieve a stabilization of the receptive field relative to the external world. Such a dynamic positional compensation may underlie preliminary evidence showing that the response of stereo-sensitive neurons in striate cortex is consistent with stimulus disparity measures and, within limits, does not reflect the retinal disparities produced by the changes in binocular alignment during fixation.

Spatial organization of axons in turtle visual cortex: Intralamellar and interlamellar projections

The spatial pattern of projections within turtle visual cortex was studied by using focal injections of horseradish peroxidase into visual cortex in an in vitro wholebrain preparation. Injections anterogradely filled the axons of many layer 2 neurons, which could be followed for 200-500 microns from the injection sites. Axons were typically unbranched, relatively straight, and bore small varicosities at irregular intervals. They radiated from the injection sites in all directions, but showed some preference toward orientations along the lateral-medial axis of the cortex. Earlier work (Mulligan and Ulinski, '90) had demonstrated that turtle visual cortex contains a series of isoazimuth lamellae, each representing an individual azimuth of visual space and oriented perpendicular to the rostral-caudal axis of the cortex. The present study provides evidence for intrinsic projections both along isoazimuth lamellae and between adjacent lamellae. These projections may play roles in the elaboration of wide receptive fields of cortical neurons.