Receptive Field Shifts Research Articles

Receptive field shifts in area MT during smooth and rapid eye movements

Receptive field shifts in area MT during smooth and rapid eye movements

The spatial distribution of receptive field changes in a model of peri-saccadic perception: Predictive remapping and shifts towards the saccade target

At the time of an impending saccade receptive fields (RFs) undergo dynamic changes, that is, their spatial profile is altered. This phenomenon has been observed in several monkey visual areas. Although their link to eye movements is obvious, neither the exact pattern nor their function is fully clear. Several RF shifts have been interpreted in terms of predictive remapping mediating visual stability. In particular, even prior to saccade onset some cells become responsive to stimuli presented in their future, post-saccadic RF. In visual area V4, however, the overall effect of RF dynamics consists of a shrinkage and shift of RFs towards the saccade target. These observations have been linked to a pre-saccadically enhanced processing of the future fixation. In order to better understand these seemingly different outcomes, we analyzed the RF shifts predicted by a recently proposed computational model of peri-saccadic perception ( Hamker, Zirnsak, Calow, & Lappe, 2008). This model unifies peri-saccadic compression, pre-saccadic attention shifts, and peri-saccadic receptive field dynamics in a common framework of oculomotor reentry signals in extrastriate visual cortical maps. According to the simulations that we present in the current paper, a spatially selective oculomotor feedback signal leads to RF dynamics which are both consistent with the observations made in studies aiming to investigate predictive remapping and saccade target shifts. Thus, the seemingly distinct experimental observations could be grounded in the same neural mechanism leading to different RF dynamics dependent on the location of the RF in visual space.

Robotic Vision with the Conformal Camera: Modeling Perisaccadic Perception

Humans make about 3 saccades per second at the eyeball's speed of 700 deg/sec to reposition the high-acuity fovea on the targets of interest to build up understanding of a scene. The brain's visuosaccadic circuitry uses the oculomotor command of each impending saccade to shift receptive fields (RFs) to cortical locations before the eyes take them there, giving a continuous and stable view of the world. We have developed a model for image representation based on projective Fourier transform (PFT) intended for robotic vision, which may efficiently process visual information during the motion of a camera with silicon retina that resembles saccadic eye movements. Here, the related neuroscience background is presented, effectiveness of the conformal camera's non-Euclidean geometry in intermediate-level vision is discussed, and the algorithmic steps in modeling perisaccadic perception with PFT are proposed. Our modeling utilizes basic properties of PFT. First, PFT is computable by FFT in complex logarithmic coordinates that also approximate the retinotopy. Second, the shift of RFs in retinotopic (logarithmic) coordinates is modeled by the shift property of discrete Fourier transform. The perisaccadic mislocalization observed by human subjects in laboratory experiments is the consequence of the fact that RFs' shifts are in logarithmic coordinates.

The dynamic routing model of visuospatial attention

Event Abstract Back to Event The dynamic routing model of visuospatial attention Bruce Bobier1*, Terry Stewart1 and Chris Eliasmith1 1 University of Waterloo, Canada The dynamic control of information routing in the brain has been most extensively studied in the context of visual attention, although the underlying mechanisms employed by the involved neural populations remain unclear. Several models have suggested that selective routing of attended information may be performed by modulating the gain of stimulus representations or by modifying synaptic connection weights. A problem with many such models is that they do not address how these processes are performed at the cellular level, or do so in a biologically implausible manner. Here we present the Dynamic Routing Model (DRM), which frames the problem of visuospatial attention as the dynamic routing information through a hierarchy of topographically organized cortical regions to an object-centered reference frame. In the DRM, the inferior and lateral pulvinar are hypothesized to encode the retinotopic location at which attention should be engaged, and project this signal to cortical control columns in each visual area. Each control column serves to guide information routing for a small group of adjacent visual columns. At each layer, the pulvinar signal is transformed by the cortical control columns to encode the location from which information should be represented. The dendrites of a neuron in a given column encode a vector containing the control signal and its inputs, and the synaptic weights of the dendrites collectively serve to approximate a function that modulates the gain of each input depending on its distance from the location specified by the cortical control signal. We begin by considering the attentional routing performed in a five layer hierarchy, where each layer is reciprocally connected with pulvinar. For each visual column, we can determine the interactions between the control and visual inputs that are required to optimally route information from the focus of attention to an object-centered reference frame at the top layer. After defining this interaction, we next investigate whether this function can be represented within the dendrites. The routing mechanism is the same for all nodes in the hierarchy, and by using the Neural Engineering Framework to model a subset of this large-scale network, we can use this high level mathematical description to analytically derive the optimal connection weights to perform the routing function. Simulation results show that in a model composed of a single cortical control population and nine visual columns, each containing 200 LIF neurons that project to a single output population, the neurons are able to accurately route visual information. We then show that the model accounts for numerous findings from neurophysiological and psychophysical studies, including attentional effects occurring earlier and more strongly in higher cortical areas, dynamic receptive field shifts, center-surround suppression, as well as impairments observed in pulvinar lesion patients. The DRM also generates several predictions concerning the character of the suppressive annulus surrounding the focus of attention, the role of pulvinar in engaging attention, and the computations performed by cortical neurons under attention. Conference: Computational and Systems Neuroscience 2010, Salt Lake City, UT, United States, 25 Feb - 2 Mar, 2010. Presentation Type: Poster Presentation Topic: Poster session III Citation: Bobier B, Stewart T and Eliasmith C (2010). The dynamic routing model of visuospatial attention. Front. Neurosci. Conference Abstract: Computational and Systems Neuroscience 2010. doi: 10.3389/conf.fnins.2010.03.00121 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 01 Mar 2010; Published Online: 01 Mar 2010. * Correspondence: Bruce Bobier, University of Waterloo, Waterloo, Canada, bbobier@engmail.uwaterloo.ca Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Bruce Bobier Terry Stewart Chris Eliasmith Google Bruce Bobier Terry Stewart Chris Eliasmith Google Scholar Bruce Bobier Terry Stewart Chris Eliasmith PubMed Bruce Bobier Terry Stewart Chris Eliasmith Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Receptive Field Shift and Shrinkage in Macaque Middle Temporal Area through Attentional Gain Modulation

Selective attention is the top-down mechanism to allocate neuronal processing resources to the most relevant subset of the information provided by an organism's sensors. Attentional selection of a spatial location modulates the spatial-tuning characteristics (i.e., the receptive fields of neurons in macaque visual cortex). These tuning changes include a shift of receptive field centers toward the focus of attention and a narrowing of the receptive field when the attentional focus is directed into the receptive field. Here, we report that when attention is directed into versus of receptive fields of neurons in the middle temporal visual area (area MT), the magnitude of the shift of the spatial-tuning functions is positively correlated with a narrowing of spatial tuning around the attentional focus. By developing and applying a general attentional gain model, we show that these nonmultiplicative attentional modulations of basic neuronal-tuning characteristics could be a direct consequence of a spatially distributed multiplicative interaction of a bell-shaped attentional spotlight with the spatially fined-grained sensory inputs of MT neurons. Additionally, the model lets us estimate the spatial spread of the attentional top-down signal impinging on visual cortex. Consistent with psychophysical reports, the estimated size of the "spotlight of attention" indicates a coarse spatial resolution of attention. These results illustrate how spatially specific nonmultiplicative attentional changes of neuronal-tuning functions can be the result of multiplicative gain modulation affecting sensory neurons in a widely distributed region in cortical space.

Electrophysiological correlates of presaccadic remapping in humans

Saccadic eye movements cause rapid displacements of space, yet the visual field is perceived as stable. A mechanism that may contribute to maintaining visual stability is the process of predictive remapping, in which receptive fields shift to their future locations prior to the onset of a saccade. We investigated electrophysiological correlates of remapping in humans using event-related potentials. Subjects made horizontal saccades that caused a visual stimulus to remain within a single visual field or to cross the vertical meridian, shifting between visual hemifields. When an impending saccade would shift the stimulus between visual fields (requiring remapping between cerebral hemispheres), presaccadic potentials showed increased bilaterality, having greater amplitudes over the hemisphere ipsilateral to the grating stimulus. Results are consistent with interhemispheric remapping of visual space in anticipation of an upcoming saccade.

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.

Dynamic shifts in the owl's auditory space map predict moving sound location

The optic tectum of the barn owl contains a map of auditory space. We found that, in response to moving sounds, the locations of receptive fields that make up the map shifted toward the approaching sound. The magnitude of the receptive field shifts increased systematically with increasing stimulus velocity and, therefore, was appropriate to compensate for sensory and motor delays inherent to auditory orienting behavior. Thus, the auditory space map is not static, but shifts adaptively and dynamically in response to stimulus motion. We provide a computational model to account for these results. Because the model derives predictive responses from processes that are known to occur commonly in neural networks, we hypothesize that analogous predictive responses will be found to exist widely in the central nervous system. This hypothesis is consistent with perceptions of stimulus motion in humans for many sensory parameters.

Dynamic shifts of visual receptive fields in cortical area MT by spatial attention

Voluntary attention is the top-down selection process that focuses cortical processing resources on the most relevant sensory information. Spatial attention--that is, selection based on stimulus position--alters neuronal responsiveness throughout primate visual cortex. It has been hypothesized that it also changes receptive field profiles by shifting their centers toward attended locations and by shrinking them around attended stimuli. Here we examined, at high resolution, receptive fields in cortical area MT of rhesus macaque monkeys when their attention was directed to different locations within and outside these receptive fields. We found a shift of receptive fields, even far from the current location of attention, accompanied by a small amount of shrinkage. Thus, already in early extrastriate cortex, receptive fields are not static entities but are highly modifiable, enabling the dynamic allocation of processing resources to attended locations and supporting enhanced perception within the focus of attention by effectively increasing the local cortical magnification.

Spatiotopic Transfer of Visual-Form Adaptation across Saccadic Eye Movements

Although conscious perception is smooth and continuous, the input to the visual system is a series of short, discrete fixations interleaved with rapid shifts of the eye. One possible explanation for visual stability is that internal maps of objects and their visual properties are remapped around the time of saccades, but numerous studies have demonstrated that visual patterns are not combined across saccades. Here, we report that visual-form aftereffects transfer across separate fixations when adaptor and test are presented in the same spatial position. The magnitude of the transsaccadic adaptation increased with stimulus complexity, suggesting a progressive construction of spatiotopic receptive fields along the visual-form pathway. These results demonstrate that basic shape information is combined across saccades, allowing for predictive and consistent information from the past to be incorporated into each new fixation.

Tuning Curve Shift by Attention Modulation in Cortical Neurons: a Computational Study of its Mechanisms

Physiological studies of visual attention have demonstrated that focusing attention near a visual cortical neuron's receptive field (RF) results in enhanced evoked activity and RF shift. In this work, we explored the mechanisms of attention induced RF shifts in cortical network models that receive an attentional 'spotlight'. Our main results are threefold. First, whereas a 'spotlight' input always produces toward-attention shift of the population activity profile, we found that toward-attention shifts in RFs of single cells requires multiplicative gain modulation. Secondly, in a feedforward two-layer model, focal attentional gain modulation in first-layer neurons induces RF shift in second-layer neurons downstream. In contrast to experimental observations, the feedforward model typically fails to produce RF shifts in second-layer neurons when attention is directed beyond RF boundaries. We then show that an additive spotlight input combined with a recurrent network mechanism can produce the observed RF shift. Inhibitory effects in a surround of the attentional focus accentuate this RF shift and induce RF shrinking. Thirdly, we considered interrelationship between visual selective attention and adaptation. Our analysis predicts that the RF size is enlarged (respectively reduced) by attentional signal directed near a cell's RF center in a recurrent network (resp. in a feedforward network); the opposite is true for visual adaptation. Therefore, a refined estimation of the RF size during attention and after adaptation would provide a probe to differentiate recurrent versus feedforward mechanisms for RF shifts.

Retinal Correspondence of Monocular Receptive Fields in Disparity-Sensitive Complex Cells from Area V1 in the Awake Monkey

To explore the neural mechanisms underlying disparity sensitivity in complex cells of the macaque visual cortex, the relationship between interocular receptive field (RF) positional shift and disparity sensitivity was studied in area V1. Single-unit recordings were made from area V1 of awake Macaca mulatta. Monocular RFs were mapped by means of a reverse cross-correlation technique, and their centers were determined after performing a bidimensional Gaussian function fitting. Interocular RF shifts were calculated for both bright and dark stimuli. Similarly, Gabor adjustments were obtained from disparity profiles to bright and dark dynamic random-dot stereograms (RDSs). Twenty-five complex cells were studied. The response profiles to disparity were similar for bright and dark RDSs. Interocular RF positional shift correlated significantly with both the peaks of Gabor fittings of disparity-sensitivity profiles and the peaks of the Gaussian envelopes of these Gabor fittings. Correlation between interocular RF positional shift and the peaks of the Gaussian envelopes was stronger than correlation between interocular RF positional shift and peaks of Gabor fittings. Interocular shift of monocular RFs is more related to the center of the range of disparities to which the cell is sensitive, than to the preferred disparity of the cell.

Effect of eye position on saccades and neuronal responses to acoustic stimuli in the superior colliculus of the behaving cat.

We examined the motor error hypothesis of visual and auditory interaction in the superior colliculus (SC), first tested by Jay and Sparks in the monkey. We trained cats to direct their eyes to the location of acoustic sources and studied the effects of eye position on both the ability of cats to localize sounds and the auditory responses of SC neurons with the head restrained. Sound localization accuracy was generally not affected by initial eye position, i.e., accuracy was not proportionally affected by the deviation of the eyes from the primary position at the time of stimulus presentation, showing that eye position is taken into account when orienting to acoustic targets. The responses of most single SC neurons to acoustic stimuli in the intact cat were modulated by eye position in the direction consistent with the predictions of the "motor error" hypothesis, but the shift accounted for only two-thirds of the initial deviation of the eyes. However, when the average horizontal sound localization error, which was approximately 35% of the target amplitude, was taken into account, the magnitude of the horizontal shifts in the SC auditory receptive fields matched the observed behavior. The modulation by eye position was not due to concomitant movements of the external ears, as confirmed by recordings carried out after immobilizing the pinnae of one cat. However, the pattern of modulation after pinnae immobilization was inconsistent with the observations in the intact cat, suggesting that, in the intact animal, information about the position of the pinnae may be taken into account.

Receptive field plasticity of neurons in rat auditory cortex

Using conventional electrophysiological technique, we investigated the plasticity of the frequency receptive fields (RF) of auditory cortex (AC) neurons in rats. In the AC, when the frequency difference between conditioning stimulus frequency (CSF) and the best frequency (BF) was in the range of 1-4 kHz, the frequency RF of AC neurons shifted. The smaller the differences between CSF and BF, the higher the probability of the RF shift and the greater the degree of the RF shift. To some extent, the plasticity of RF was dependent on the duration of the session of conditioning stimulus (CS). When the frequency difference between CSF and BF was bigger, the duration of the CS session needed to induce the plasticity was longer. The recovery time course of the frequency RF showed opposite changes after CS cessation. The RF shift could be induced by the frequency that was either higher or lower than the control BF, demonstrating no clear directional preference. The frequency RF of some neurons showed bidirectional shift, and the RF of other neurons showed single directional shift. The results suggest that the frequency RF plasticity of AC neurons could be considered as an ideal model for studying plasticity mechanism. The present study also provides important evidence for further study of learning and memory in auditory system.

Information optimization in coupled audio-visual cortical maps.

Barn owls hunt in the dark by using cues from both sight and sound to locate their prey. This task is facilitated by topographic maps of the external space formed by neurons (e.g., in the optic tectum) that respond to visual or aural signals from a specific direction. Plasticity of these maps has been studied in owls forced to wear prismatic spectacles that shift their visual field. Adaptive behavior in young owls is accompanied by a compensating shift in the response of (mapped) neurons to auditory signals. We model the receptive fields of such neurons by linear filters that sample correlated audio-visual signals and search for filters that maximize the gathered information while subject to the costs of rewiring neurons. Assuming a higher fidelity of visual information, we find that the corresponding receptive fields are robust and unchanged by artificial shifts. The shape of the aural receptive field, however, is controlled by correlations between sight and sound. In response to prismatic glasses, the aural receptive fields shift in the compensating direction, although their shape is modified due to the costs of rewiring.

The time course of perisaccadic receptive field shifts in the lateral intraparietal area of the monkey.

Neurons in the lateral intraparietal area of the monkey (LIP) have visual receptive fields in retinotopic coordinates when studied in a fixation task. However, in the period immediately surrounding a saccade these receptive fields often shift, so that a briefly flashed stimulus outside the receptive field will drive the neurons if the eye movement will bring the spatial location of that vanished stimulus into the receptive field. This is equivalent to a transient shift of the retinal receptive field. The process enables the monkey brain to process a stimulus in a spatially accurate manner after a saccade, even though the stimulus appeared only before the saccade. We studied the time course of this receptive field shift by flashing a task-irrelevant stimulus for 100 ms before, during, or after a saccade. The stimulus could appear in receptive field as defined by the fixation before the saccade (the current receptive field) or the receptive field as defined by the fixation after the saccade (the future receptive field). We recorded the activity of 48 visually responsive neurons in LIP of three hemispheres of two rhesus monkeys. We studied 45 neurons in the current receptive field task, in which the saccade removed the stimulus from the receptive field. Of these neurons 29/45 (64%) showed a significant decrement of response when the stimulus appeared 250 ms or less before the saccade, as compared with their activity during fixation. The average response decrement was 38% for those cells showing a significant (P < 0.05 by t-test) decrement. We studied 39 neurons in the future receptive field task, in which the saccade brought the spatial location of a recently vanished stimulus into the receptive field. Of these 32/39 (82%) had a significant response to stimuli flashed for 100 ms in the future receptive field, even 400 ms before the saccade. Neurons never responded to stimuli moved by the saccade from a point outside the receptive field to another point outside the receptive field. Neurons did not necessarily show any saccadic suppression for stimuli moved from one part of the receptive field to another by the saccade. Stimuli flashed <250 ms before the saccade-evoked responses in both the presaccadic and the postsaccadic receptive fields, resulting in an increase in the effective receptive field size, an effect that we suggest is responsible for perisaccadic perceptual inaccuracies.

Temporal specificity in the cortical plasticity of visual space representation.

The circuitry and function of mammalian visual cortex are shaped by patterns of visual stimuli, a plasticity likely mediated by synaptic modifications. In the adult cat, asynchronous visual stimuli in two adjacent retinal regions controlled the relative spike timing of two groups of cortical neurons with high precision. This asynchronous pairing induced rapid modifications of intracortical connections and shifts in receptive fields. These changes depended on the temporal order and interval between visual stimuli in a manner consistent with spike timing-dependent synaptic plasticity. Parallel to the cortical modifications found in the cat, such asynchronous visual stimuli also induced shifts in human spatial perception.

Motion processing in the auditory cortex of the rufous horseshoe bat: role of GABAergic inhibition.

This study examined the influence of inhibition on motion-direction-sensitive responses of neurons in the dorsal fields of auditory cortex of the rufous horseshoe bat. Responses to auditory apparent motion stimuli were recorded extracellularly from neurons while microiontophoretically applying gamma-aminobutyric acid (GABA) and the GABAA receptor antagonist bicuculline methiodide (BMI). Neurons could respond with a directional preference exhibiting stronger responses to one direction of motion or a shift of receptive field (RF) borders depending on direction of motion. BMI influenced the motion direction sensitivity of 53% of neurons. In 21% of neurons the motion-direction sensitivity was decreased by BMI by decreasing either directional preference or RF shift. In neurons with a directional preference, BMI increased the spike number for the preferred direction by a similar amount as for the nonpreferred direction. Thus, inhibition was not direction specific. BMI increased motion-direction sensitivity by either increasing directional preference or magnitude of RF shifts in 22% of neurons. Ten percent of neurons changed their response from a RF shift to a directional preference under BMI. In these neurons, the observed effects could often be better explained by adaptation of excitation rather than inhibition. The results suggest, that adaptation of excitation, as well as cortex specific GABAergic inhibition, contribute to motion-direction sensitivity in the auditory cortex of the rufous horseshoe bat.

Eye Movements Modulate Visual Receptive Fields of V4 Neurons

The receptive field, defined as the spatiotemporal selectivity of neurons to sensory stimuli, is central to our understanding of the neuronal mechanisms of perception. However, despite the fact that eye movements are critical during normal vision, the influence of eye movements on the structure of receptive fields has never been characterized. Here, we map the receptive fields of macaque area V4 neurons during saccadic eye movements and find that receptive fields are remarkably dynamic. Specifically, before the initiation of a saccadic eye movement, receptive fields shrink and shift towards the saccade target. These spatiotemporal dynamics may enhance information processing of relevant stimuli during the scanning of a visual scene, thereby assisting the selection of saccade targets and accelerating the analysis of the visual scene during free viewing.

Can Hebbian volume learning explain discontinuities in cortical maps?

It has recently been shown that orientation and retinotopic position, both of which are mapped in primary visual cortex, can show correlated jumps (Das & Gilbert, 1997). This is not consistent with maps generated by Kohonen's algorithm (Kohonen, 1982), where changes in mapped variables tend to be anticorrelated. We show that it is possible to obtain correlated jumps by introducing a Hebbian component (Hebb, 1949) into Kohonen's algorithm. This correspondents to a volume learning mechanism where synaptic facilitation depends not only on the spread of a signal from a maximally active neuron but also requires postsynaptic activity at a synapse. The maps generated by this algorithm show discontinuities across which both orientation and retinotopic position change rapidly, but these regions, which include the orientation singularities, are also aligned with the edges of ocular dominance columns, and this is not a realistic feature of cortical maps. We conclude that cortical maps are better modeled by standard, non-Hebbian volume learning, perhaps coupled with some other mechanism (e.g., that of Ernst, Pawelzik, Tsodyks, & Sejnowski, 1999) to produce receptive field shifts.