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Showing papers on "Receptive field published in 2004"


Journal ArticleDOI
TL;DR: The results show that feature-based attention exerts a multiplicative modulation upon neuronal responses and that the strength of this modulation depends on the similarity between the attended feature and the cell's preferred feature, in line with the feature-similarity gain model.

656 citations


Journal ArticleDOI
TL;DR: The results provide the first demonstration of perceptual learning modifying basic neuronal response properties at an intermediate level of visual cortex and give insights into the distribution of plasticity across adult visual cortex.
Abstract: Previous studies have shown that perceptual learning can substantially alter the response properties of neurons in the primary somatosensory and auditory cortices. Although psychophysical studies suggest that perceptual learning induces similar changes in primary visual cortex (V1), studies that have measured the response properties of individual neurons have failed to find effects of the size described for the other sensory systems. We have examined the effect of learning on neuronal response properties in a visual area that lies at a later stage of cortical processing, area V4. Adult macaque monkeys were trained extensively on orientation discrimination at a specific retinal location using a narrow range of orientations. During the course of training, the subjects achieved substantial improvement in orientation discrimination that was primarily restricted to the trained location. After training, neurons in V4 with receptive fields overlapping the trained location had stronger responses and narrower orientation tuning curves than neurons with receptive fields in the opposite, untrained hemifield. The changes were most prominent for neurons that preferred orientations close to the trained range of orientations. These results provide the first demonstration of perceptual learning modifying basic neuronal response properties at an intermediate level of visual cortex and give insights into the distribution of plasticity across adult visual cortex.

412 citations


Journal ArticleDOI
TL;DR: This work uses the Bayes' rule Chapman-Kolmogorov paradigm with a linear state equation and point process observation models to derive adaptive filters appropriate for estimation from neural spike trains and suggests a practical approach for constructing filtering algorithms to track neural receptive field dynamics on a millisecond timescale.
Abstract: Neural receptive fields are dynamic in that with experience, neurons change their spiking responses to relevant stimuli. To understand how neural systems adapt their representations of biological information, analyses of receptive field plasticity from experimental measurements are crucial. Adaptive signal processing, the well-established engineering discipline for characterizing the temporal evolution of system parameters, suggests a framework for studying the plasticity of receptive fields. We use the Bayes' rule Chapman-Kolmogorov paradigm with a linear state equation and point process observation models to derive adaptive filters appropriate for estimation from neural spike trains. We derive point process filter analogues of the Kalman filter, recursive least squares, and steepest-descent algorithms and describe the properties of these new filters. We illustrate our algorithms in two simulated data examples. The first is a study of slow and rapid evolution of spatial receptive fields in hippocampal neurons. The second is an adaptive decoding study in which a signal is decoded from ensemble neural spiking activity as the receptive fields of the neurons in the ensemble evolve. Our results provide a paradigm for adaptive estimation for point process observations and suggest a practical approach for constructing filtering algorithms to track neural receptive field dynamics on a millisecond timescale.

379 citations


Journal ArticleDOI
TL;DR: A model of invariant visual object recognition in the brain that incorporates feedback biasing effects of top-down attentional mechanisms on a hierarchically organized set of visual cortical areas with convergent forward connectivity, reciprocal feedback connections, and local intra-area competition is described.

331 citations


Journal ArticleDOI
TL;DR: A large difference in predictive power suggests that natural spatiotemporal stimulus statistics activate nonlinear response properties in a different manner than the grating stimulus, which contributes to the modulatory effects of nonlinear temporal summation during natural vision.
Abstract: Studies of the primary visual cortex (V1) have produced models that account for neuronal responses to synthetic stimuli such as sinusoidal gratings. Little is known about how these models generalize to activity during natural vision. We recorded neural responses in area V1 of awake macaques to a stimulus with natural spatiotemporal statistics and to a dynamic grating sequence stimulus. We fit nonlinear receptive field models using each of these data sets and compared how well they predicted time-varying responses to a novel natural visual stimulus. On average, the model fit using the natural stimulus predicted natural visual responses more than twice as accurately as the model fit to the synthetic stimulus. The natural vision model produced better predictions in >75% of the neurons studied. This large difference in predictive power suggests that natural spatiotemporal stimulus statistics activate nonlinear response properties in a different manner than the grating stimulus. To characterize this modulation, we compared the temporal and spatial response properties of the model fits. During natural stimulation, temporal responses often showed a stronger late inhibitory component, indicating an effect of nonlinear temporal summation during natural vision. In addition, spatial tuning underwent complex shifts, primarily in the inhibitory, rather than excitatory, elements of the response profile. These differences in late and spatially tuned inhibition accounted fully for the difference in predictive power between the two models. Both the spatial and temporal statistics of the natural stimulus contributed to the modulatory effects.

330 citations


Journal ArticleDOI
TL;DR: It is suggested that the extraction of information of angles embedded within contour stimuli may start in area V2, and Angle-selective V2 neurons are unlikely to be specific angle detectors, because the magnitude of their responses to the optimal angle was indistinguishable from that from the optimal half-lines.
Abstract: Angles and junctions embedded within contours are important features to represent the shape of objects. To study the neuronal basis to extract these features, we conducted extracellular recordings while two macaque monkeys performed a fixation task. Angle stimuli were the combination of two straight half-lines larger than the size of the classical receptive fields (CRFs). Each line was drawn from the center to outside the CRFs in 1 of 12 directions, so that the stimuli passed through the CRFs and formed angles at the center of the CRFs. Of 114 neurons recorded from the superficial layer of area V2, 91 neurons showed selective responses to these angle stimuli. Of these, 41 neurons (36.0%) showed selective responses to wide angles between 60° and 150° that were distinct from responses to straight lines or sharp angles (30°). Responses were highly selective to a particular angle in approximately one-fourth of neurons. When we tested the selectivity of the same neurons to individual half-lines, the preferred direction was more or less consistent with one or two components of the optimal angle stimuli. These results suggest that the selectivity of the neurons depends on both the combination of two components and the responses to individual components. Angle-selective V2 neurons are unlikely to be specific angle detectors, because the magnitude of their responses to the optimal angle was indistinguishable from that to the optimal half-lines. We suggest that the extraction of information of angles embedded within contour stimuli may start in area V2.

288 citations


Journal ArticleDOI
TL;DR: In vivo whole-cell methods in the rat auditory cortex are used to record subthreshold membrane potential fluctuations elicited by complex acoustic stimuli, including animal vocalizations, and find that the STRF has a rich dynamical structure, including excitatory regions positioned in general accord with the prediction of the classical tuning curve.
Abstract: How do cortical neurons represent the acoustic environment? This question is often addressed by probing with simple stimuli such as clicks or tone pips. Such stimuli have the advantage of yielding easily interpreted answers, but have the disadvantage that they may fail to uncover complex or higher-order neuronal response properties. Here, we adopt an alternative approach, probing neuronal responses with complex acoustic stimuli, including animal vocalizations. We used in vivo whole-cell methods in the rat auditory cortex to record subthreshold membrane potential fluctuations elicited by these stimuli. Most neurons responded robustly and reliably to the complex stimuli in our ensemble. Using regularization techniques, we estimated the linear component, the spectrotemporal receptive field (STRF), of the transformation from the sound (as represented by its time-varying spectrogram) to the membrane potential of the neuron. We find that the STRF has a rich dynamical structure, including excitatory regions positioned in general accord with the prediction of the classical tuning curve. However, whereas the STRF successfully predicts the responses to some of the natural stimuli, it surprisingly fails completely to predict the responses to others; on average, only 11% of the response power could be predicted by the STRF. Therefore, most of the response of the neuron cannot be predicted by the linear component, although the response is deterministically related to the stimulus. Analysis of the systematic errors of the STRF model shows that this failure cannot be attributed to simple nonlinearities such as adaptation to mean intensity, rectification, or saturation. Rather, the highly nonlinear response properties of auditory cortical neurons must be attributable to nonlinear interactions between sound frequencies and time-varying properties of the neural encoder.

285 citations


Journal ArticleDOI
TL;DR: It is shown that visual learning can selectively increase the overall response to trained stimuli in primary visual cortex (V1) and correlated moderately well with the magnitude of changes in detection performance.

276 citations


Journal ArticleDOI
TL;DR: Recording and spike sorting techniques are reported that enable us to record from all or nearly all of the ganglion cells in a patch of the retina with a dense multi-electrode array.
Abstract: To understand a neural circuit completely requires simultaneous recording from most of the neurons in that circuit. Here we report recording and spike sorting techniques that enable us to record from all or nearly all of the ganglion cells in a patch of the retina. With a dense multi-electrode array, each ganglion cell produces a unique pattern of activity on many electrodes when it fires an action potential. Signals from all of the electrodes are combined with an iterative spike sorting algorithm to resolve ambiguities arising from overlapping spike waveforms. We verify that we are recording from a large fraction of ganglion cells over the array by labeling the ganglion cells with a retrogradely transported dye and by comparing the number of labeled and recorded cells. Using these methods, we show that about 60 receptive fields of ganglion cells cover each point in visual space in the salamander, consistent with anatomical findings.

245 citations


Journal ArticleDOI
TL;DR: It is found that the modulation ratio, when derived from the subthreshold membrane potential instead of from spike rate, is unimodally distributed, but highly skewed.
Abstract: The existence of two classes of cells, simple and complex, discovered by Hubel and Wiesel in 1962, is one of the fundamental features of cat primary visual cortex. A quantitative measure used to distinguish simple and complex cells is the ratio between modulated and unmodulated components of spike responses to drifting gratings, an index that forms a bimodal distribution. We have found that the modulation ratio, when derived from the subthreshold membrane potential instead of from spike rate, is unimodally distributed, but highly skewed. The distribution of the modulation ratio as derived from spike rate can, in turn, be predicted quantitatively by the nonlinear properties of spike threshold applied to the skewed distribution of the subthreshold modulation ratio. Threshold also increases the spatial segregation of ON and OFF regions of the receptive field, a defining attribute of simple cells. The distinction between simple and complex cells is therefore enhanced by threshold, much like the selectivity for stimulus features such as orientation and direction. In this case, however, a continuous distribution in the spatial organization of synaptic inputs is transformed into two distinct classes of cells.

230 citations


Journal ArticleDOI
TL;DR: It is suggested that LGN bursts are an important part of the neural code, providing a nonlinear amplification of stimulus features that are typical of the natural environment.
Abstract: The role of the lateral geniculate nucleus (LGN) of the thalamus in visual encoding remains an open question. Here, we characterize the function of tonic and burst spikes in cat LGN X-cells in signaling features of natural stimuli. A significant increase in bursting was observed during natural stimulation (relative to white noise stimulation) and was linked to the strong correlation structure of the natural scene movies. Burst responses were triggered by specific stimulus events consisting of a prolonged inhibitory stimulus, followed by an excitatory stimulus, such as the movement of an object into the receptive field. LGN responses to natural scene movies were predicted using an integrate-and-fire (IF) framework and compared with experimentally observed responses. The standard IF model successfully predicted LGN responses to natural scene movies during tonic firing, indicating a linear relationship between stimulus and response. However, the IF model typically underpredicted the LGN response during periods of bursting, indicating a nonlinear amplification of the stimulus in the actual response. The addition of a burst mechanism to the IF model was necessary to accurately predict the entire LGN response. These results suggest that LGN bursts are an important part of the neural code, providing a nonlinear amplification of stimulus features that are typical of the natural environment.

Journal ArticleDOI
TL;DR: It is proposed that thalamocortical and intracortical pathways preferentially contribute to responses evoked by CF and non-CF stimuli, respectively, and that intracortsical projections linking frequency representations determine the breadth of receptive fields in primary ACx.
Abstract: To examine the basis of frequency receptive fields in auditory cortex (ACx), we have recorded intracellular (whole cell) and extracellular (local field potential, LFP) responses to tones in anesthe...

Journal ArticleDOI
TL;DR: In vivo whole‐cell recordings from L5 neurones of the somatosensory (barrel) cortex of urethane‐anaesthetized rats conclude that in L5 a whisker deflection is represented in two ways: focally by L5A pyramids and more diffusely byL5B pyramids as a result of combining different inputs from lemniscal and paralemniscal pathways.
Abstract: Layer 5 (L5) pyramidal neurones constitute a major sub- and intracortical output of the somatosensory cortex. This layer 5 is segregated into layers 5A and 5B which receive and distribute relatively independent afferent and efferent pathways. We performed in vivo whole-cell recordings from L5 neurones of the somatosensory (barrel) cortex of urethane-anaesthetized rats (aged 27-31 days). By delivering 6 deg single whisker deflections, whisker pad receptive fields were mapped for 16 L5A and 11 L5B neurones located below the layer 4 whisker-barrels. Average resting membrane potentials were -75.6 +/- 1.1 mV, and spontaneous action potential (AP) rates were 0.54 +/- 0.14 APs s(-1). Principal whisker (PW) evoked responses were similar in L5A and L5B neurones, with an average 5.0 +/- 0.6 mV postsynaptic potential (PSP) and 0.12 +/- 0.03 APs per stimulus. The layer 5A sub- and suprathreshold receptive fields (RFs) were more confined to the principle whisker than those of layer 5B. The basal dendritic arbors of layer 5A and 5B cells were located below both layer 4 barrels and septa, and the cell bodies were biased towards the barrel walls. Responses in both L5A and L5B developed slowly, with onset latencies of 10.1 +/- 0.5 ms and peak latencies of 33.9 +/- 3.3 ms. Contralateral multi-whisker stimulation evoked PSPs similar in amplitude to those of PW deflections; whereas, ipsilateral stimulation evoked smaller and longer latency PSPs. We conclude that in L5 a whisker deflection is represented in two ways: focally by L5A pyramids and more diffusely by L5B pyramids as a result of combining different inputs from lemniscal and paralemniscal pathways. The relevant output evoked by a whisker deflection could be the ensemble activity in the anatomically defined cortical modules associated with a single or a few barrel-columns.

Journal ArticleDOI
10 Jun 2004-Neuron
TL;DR: Observing the development of circuits in the rat barrel cortex using laser-scanning photostimulation analysis of synaptic connectivity, reconstructions of axonal arbors, and in vivo whole-cell recording revealed already spatially well-tuned receptive fields.

Journal ArticleDOI
TL;DR: It is demonstrated that either a simplified model of synaptic depression and facilitation, and/or a cortical network of thalamic excitation and cortical inhibition can account for major trends in the observed findings.
Abstract: Although single units in primary auditory cortex (A1) exhibit accurate timing in their phasic response to the onset of sound (precision of a few milliseconds), paradoxically, they are unable to sustain synchronized responses to repeated stimuli at rates much beyond 20 Hz. To explore the relationship between these two aspects of cortical response, we designed a broadband stimulus with a slowly modulated spectrotemporal envelope riding on top of a rapidly modulated waveform (or fine structure). Using this stimulus, we quantified the ability of cortical cells to encode independently and simultaneously the stimulus envelope and fine structure. Specifically, by reverse-correlating unit responses with these two stimulus dimensions, we measured the spectrotemporal response fields (STRFs) associated with the processing of the envelope, the fine structure, and the complete stimulus. A1 cells respond well to the slow spectrotemporal envelopes and produce a wide variety of STRFs. In over 70% of cases, A1 units also track the fine-structure modulations precisely, throughout the stimulus, and for frequencies up to several hundred Hertz. Such a dual response, however, is contingent on the cell being driven by both fast and slow modulations, in that the response to the slowly modulated envelope gates the expression of the fine structure. We also demonstrate that either a simplified model of synaptic depression and facilitation, and/or a cortical network of thalamic excitation and cortical inhibition can account for major trends in the observed findings. Finally, we discuss the potential functional significance and perceptual relevance of these coexistent, complementary dynamic response modes.

Journal ArticleDOI
06 May 2004-Nature
TL;DR: It is reported that the transfer of adult rats reared in standard laboratory cages to a naturalistic habitat modifies the functional and morphological organization of the facial whisker representation in the somatosensory ‘barrel’ cortex.
Abstract: Much of what is known about the functional organization and plasticity of adult sensory cortex is derived from animals housed in standard laboratory cages. Here we report that the transfer of adult rats reared in standard laboratory cages to a naturalistic habitat modifies the functional and morphological organization of the facial whisker representation in the somatosensory 'barrel' cortex. Cortical whisker representations, visualized with repeated intrinsic signal optical imaging in the same animals, contracted by 46% after four to six weeks of exposure to the naturalistic habitat. Acute, multi-site extracellular recordings demonstrated suppressed evoked neuronal responses and smaller, sharper constituent receptive fields in the upper cortical layers (II/III), but not in the thalamic recipient layer (IV), of rats with naturalistic experience. Morphological plasticity of the layer IV barrel field was observed, but on a substantially smaller scale than the functional plasticity. Thus, transferring animals to an environment that promotes the expression of natural, innate behaviours induces a large-scale functional refinement of cortical sensory maps.

Journal ArticleDOI
TL;DR: The hypothesis that the polysensory zone in the precentral gyrus contributes to the control of defensive movements in monkeys is supported, and the view that the pre central gyrus can control movement at the level of complex sensorimotor tasks is supported.
Abstract: The precentral gyrus of monkeys contains a polysensory zone in which the neurons respond to tactile, visual, and sometimes auditory stimuli. The tactile receptive fields of the polysensory neurons are usually on the face, arms, or upper torso, and the visual and auditory receptive fields are usually confined to the space near the tactile receptive fields, within about 30 cm of the body. Electrical stimulation of this polysensory zone, even in anesthetized animals, evokes a specific set of movements. The movements resemble those typically used to defend the body from objects that are near, approaching, or touching the skin. In the present study, to determine whether the stimulation-evoked movements represent a normal set of defensive movements, we tested whether they include a distinctive, nonsaccadic, centering movement of the eyes that occurs during defensive reactions. We report that this centering movement of the eyes is evoked by stimulation of sites in the polysensory zone. We also recorded the activity of neurons in the polysensory zone while the monkey made defensive reactions to an air puff on the face. The neurons became active during the defensive movement, and the magnitude of this activity was correlated with the magnitude of the defensive reaction. These results support the hypothesis that the polysensory zone in the precentral gyrus contributes to the control of defensive movements. More generally, the results support the view that the precentral gyrus can control movement at the level of complex sensorimotor tasks.

Journal ArticleDOI
19 Feb 2004-Neuron
TL;DR: These findings demonstrate that somatosensory cortex can selectively regulate thalamic spatial response tuning by engaging topographically specific excitatory and inhibitory mechanisms in the thalamus.

Journal ArticleDOI
TL;DR: It is demonstrated here that neurons in the inferior colliculus (ICC) of the cat show adaptation to dynamic auditory stimuli that differ in the variance of their modulation depth distribution and suggested that adaptation to variance is a general property of sensory systems that allows them to effectively deal with a nonstationary environment.
Abstract: Short-term adaptation and recovery from adaptation have a strong impact on the processing of dynamic stimuli Adaptive effects on neuronal activity have been studied most commonly for changes in first-order statistics of stimuli such as stepwise increments or decrements in stimulus amplitude However, changes in higher moment statistics, such as the variance of the amplitude distribution in visual stimuli, also can invoke pronounced adaptation behavior We demonstrate here that neurons in the inferior colliculus (ICC) of the cat show adaptation to dynamic auditory stimuli that differ in the variance of their modulation depth distribution In addition, it is shown that neurons show adaptation to other higher moment statistics (eg, kurtosis) of the modulation envelope The time course of adaptation is specific for the altered stimulus property and the direction of parameter change The use of dynamic stimuli allows an estimate of the effects of the adaptation on the temporal response properties of the neurons We demonstrate that temporal receptive fields of neurons undergo change during the course of adaptation We show that adaptation to variance in the ICC has many similarities to that in the retina and suggest that adaptation to variance is a general property of sensory systems that allows them to effectively deal with a nonstationary environment

Journal ArticleDOI
TL;DR: A brief historical account of the evolution of receptive field mapping in visual cortex along with the associated conceptual advancements is provided, and speculation on the shape novel theories of the cortex may take as a result of these measurements is speculated.
Abstract: Nearly 40 years ago, in the pages of this journal, Hubel and Wiesel provided the first description of receptive fields in the primary visual cortex of higher mammals. They defined two classes of cortical cells, ‘simple’ and ‘complex’, based on neural responses to simple visual stimuli. The notion of a hierarchy of receptive fields, where increasingly intricate receptive fields are constructed from more elementary ones, was introduced. Since those early days we have witnessed the birth of quantitative methods to map receptive fields and mathematical descriptions of simple and complex cell function. Insights gained from these models, along with new theoretical concepts, are refining our understanding of receptive field structure and the underlying cortical circuitry. Here, I provide a brief historical account of the evolution of receptive field mapping in visual cortex along with the associated conceptual advancements, and speculate on the shape novel theories of the cortex may take as a result these measurements.

Journal ArticleDOI
TL;DR: In this article, the effect of top-down inputs on the firing rate and coherence of neurons in the visual cortex has been investigated in the presence of mod- ulatory inputs.
Abstract: Recordings from area V4 of monkeys have revealed that when the focus of attention is on a visual stimulus within the receptive field of a cortical neuron, two distinct changes can occur: The firing rate of the neuron can change and there can be an increase in the coherence between spikes and the local field potential (LFP) in the gamma-frequency range (30-50 Hz). The hypothesis explored here is that these observed effects of attention could be a consequence of changes in the synchrony of local interneuron networks. We performed computer simulations of a Hodgkin-Huxley type neuron driven by a constant depolarizing current, I, representing visual stimulation and a mod- ulatory inhibitory input representing the effects of attention via local interneuron networks. We observed that the neurons firing rate and the coherence of its output spike train with the synaptic inputs was modulated by the degree of synchrony of the inhibitory inputs. When inhibitory synchrony increased, the coherence of spiking model neurons with the synaptic input increased, but the firing rate either increased or remained the same. The mean number of synchronous inhibitory inputs was a key determinant of the shape of the firing rate versus current (f-I) curves. For a large number of inhibitory inputs (� 50), the f-I curve saturated for large I and an increase in input synchrony resulted in a shift of sensitivity—the model neuron responded to weaker inputs I. For a small number (� 10), the f-I curves were non-saturating and an increase in input synchrony led to an increase in the gain of the response—the firing rate in response to the same input was multiplied by an approximately constant factor. The firing rate modulation with inhibitory synchrony was highest when the input network oscillated in the gamma frequency range. Thus, the observed changes in firing rate and coherence of neurons in the visual cortex could be controlled by top-down inputs that regulated the coherence in the activity of a local inhibitory network discharging at gamma frequencies. � 2005 Elsevier Ltd. All rights reserved.

Journal ArticleDOI
TL;DR: The results suggest that surround suppression is not primarily attributable to the intracortical inhibition, but because of a reduction of thalamocortical inputs, which drive the cortex excitation and inhibition, and a subsequent decrease in the cortical excitatory interactions.
Abstract: In the primary visual cortex (V1), the single-neuron response to a grating stimulus placed in the classical receptive field (CRF) is suppressed by a similar stimulus presented in the CRF surround. To assess the input mechanism underlying the surround suppression, we tested the effects of iontophoretically administered GABAA-receptor antagonist, bicuculline methiodide (BMI), for the 46 V1 neurons in anesthetized cats. First, the stimulus-size tuning curves were studied, with or without BMI administration, for each neuron by changing the size of the grating patch. During the BMI administration, the shape of the normalized size tuning curve did not change considerably. Second, the dependency of surround suppression on the orientation of the surround grating was examined. In the control, the surround suppression showed the clear orientation tuning that peaked at an orientation the same as the optimal orientation of the CRF response. The BMI administration did not change the orientation dependency of surround suppression. We also estimated the relative contribution of excitation and inhibition to the size and orientation tuning of surround suppression. It was concluded that cortical excitation and inhibition were well balanced, having similar tuning profiles for both stimulus size and orientation of the surround grating. Furthermore, surround stimuli used for V1 neurons suppressed the CRF response of neurons in the lateral geniculate nucleus. These results suggest that surround suppression is not primarily attributable to the intracortical inhibition, but because of a reduction of thalamocortical inputs, which drive the cortical excitation and inhibition, and a subsequent decrease in the cortical excitatory interactions.

Journal ArticleDOI
TL;DR: Interestingly, simultaneous dual whole-cell in vivo recordings show that chandelier cells, which rarely fire action potentials spontaneously, fire more robustly than other types of cortical neurons when the overall cortical excitation increases.
Abstract: Chandelier cells form inhibitory axo-axonic synapses on pyramidal neurons with their characteristic candlestick-like axonal terminals. The functional role of chandelier cells is still unclear, although the preferential loss of this cell type at epileptic loci suggests a role in epilepsy. Here we report an examination of whisker- and spontaneous activity-evoked responses in chandelier cells and other fast-spiking nonpyramidal neurons and regular-spiking pyramidal neurons in layer 2/3 of the barrel cortex. Fast-spiking nonpyramidal neurons, including chandelier cells, basket cells, neurogliaform cells, double bouquet cells, net basket cells, bitufted cells, and regular-spiking pyramidal neurons all respond to stimulation of multiple whiskers on the contralateral face. Whisker stimulation, however, evokes small, delayed EPSPs preceded by an earlier IPSP and no action potentials in chandelier cells, different from other nonpyramidal and pyramidal neurons. In addition, chandelier cells display a larger receptive field with lower acuity than other fast-spiking nonpyramidal neurons and pyramidal neurons. Notably, simultaneous dual whole-cell in vivo recordings show that chandelier cells, which rarely fire action potentials spontaneously, fire more robustly than other types of cortical neurons when the overall cortical excitation increases. Thus, chandelier cells may not process fast ascending sensory information but instead may be reserved to prevent excessive excitatory activity in neuronal networks.

Journal ArticleDOI
TL;DR: In this paper, a camera carried by a freely moving cat was used to capture the dynamics of image motion as the animal explores an outdoor environment, and the movie sequences were used as input to simulated neurons.
Abstract: Sensory areas should be adapted to the properties of their natural stimuli. What are the underlying rules that match the properties of complex cells in primary visual cortex to their natural stimuli? To address this issue, we sampled movies from a camera carried by a freely moving cat, capturing the dynamics of image motion as the animal explores an outdoor environment. We use these movie sequences as input to simulated neurons. Following the intuition that many meaningful high-level variables, e.g., identities of visible objects, do not change rapidly in natural visual stimuli, we adapt the neurons to exhibit firing rates that are stable over time. We find that simulated neurons, which have optimally stable activity, display many properties that are observed for cortical complex cells. Their response is invariant with respect to stimulus translation and reversal of contrast polarity. Furthermore, spatial frequency selectivity and the aspect ratio of the receptive field quantitatively match the experimentally observed characteristics of complex cells. Hence, the population of complex cells in the primary visual cortex can be described as forming an optimally stable representation of natural stimuli.

Journal ArticleDOI
TL;DR: An attention-driven center-surround distribution of neural activity may enhance the contrast between attended and non-attended objects and relate the different behavior of extrastriate areas to their lower spatial resolution, i.e. larger receptive fields.
Abstract: By enhancing neural activity in respective retinotopic cortical representations attention increases the efficiency with which visual information at a selected location is processed. Behavioral data also suggest that information from the vicinity of the attended region is actively suppressed. In search for a physiological correlate of this 'spotlight's penumbra' we assessed neural responses in retinotopic representations of an attended location and of locations at different distances to it. Relative to passive viewing we found suppressed striate activity for the nearby but not for the far locations. This attention-driven center-surround distribution of neural activity may enhance the contrast between attended and non-attended objects. We relate the different behavior of extrastriate areas to their lower spatial resolution, i.e. larger receptive fields.

Journal ArticleDOI
TL;DR: A study of the functional organization of the SII hand region that combines microelectrode mapping using hand-held stimuli with single-unit recordings using a motorized, computer-controlled tactile oriented bar finds evidence that this region consists of multiple functional fields, with a central field containing neurons that are driven well by cutaneous stimuli.
Abstract: The detailed functional organization of the macaque second somatosensory cortex (SII) is not well understood. Here we report the results of a study of the functional organization of the SII hand region that combines microelectrode mapping using hand-held stimuli with single-unit recordings using a motorized, computer-controlled tactile oriented bar. The data indicate that the SII hand region extends ∼10 mm in the anteroposterior (AP) dimension, primarily within the upper bank of the lateral sulcus. Furthermore, we find evidence that this region consists of multiple functional fields, with a central field containing neurons that are driven well by cutaneous stimuli, flanked by an anterior field and a posterior field that each contain neurons that are driven well by proprioceptive stimuli and less well by cutaneous stimuli. The anterior field extends ∼4-5 mm AP, the central field extends ∼3-4 mm, and the posterior field extends ∼3 mm. Data from the motorized stimulator indicate that neurons in the central field are more responsive to oriented bars, more frequently exhibit orientation tuning, and have larger receptive fields than neurons in the anterior and posterior fields. We speculate that the three putative fields play different functional roles in tactile perception; the anterior and posterior fields process information that involves both proprioceptive and cutaneous input such as sensorimotor integration or stereognosis, whereas the central field processes primarily cutaneous information.

Journal ArticleDOI
TL;DR: Data are consistent with the hypothesis that the surrounds of light-adapted parasol ganglion cells are generated primarily by non-GABAergic horizontal cell feedback in the outer retina, with a small contribution from GABAergic amacrine cells of the inner retina.
Abstract: Although the center-surround receptive field is a fundamental property of retinal ganglion cells, the circuitry that mediates surround inhibition remains controversial. We examined the contribution of horizontal cells and amacrine cells to the surround of parasol ganglion cells of macaque and baboon retina by measuring receptive field structure before and during the application of drugs that have been shown previously to affect surrounds in a range of mammalian and nonmammalian species. Carbenoxolone and cobalt, thought to attenuate feedback from horizontal cells to cones, severely reduced the surround. Tetrodotoxin, which blocks sodium spiking in amacrine cells, and picrotoxin, which blocks the inhibitory action of GABA, only slightly reduced the surround. These data are consistent with the hypothesis that the surrounds of light-adapted parasol ganglion cells are generated primarily by non-GABAergic horizontal cell feedback in the outer retina, with a small contribution from GABAergic amacrine cells of the inner retina.

Journal ArticleDOI
TL;DR: In many species, neurons responding to visual motion at higher processing stages are often specifically tuned to particular flow fields; however, the neural circuitry that leads to this selectivity is not yet understood, and 'vertical system' cells of the blowfly lobula plate are studied.
Abstract: In many species, neurons responding to visual motion at higher processing stages are often specifically tuned to particular flow fields; however, the neural circuitry that leads to this selectivity is not yet understood. Here we have studied this problem in 'vertical system' (VS) cells of the blowfly lobula plate. These neurons possess distinctive local preferred directions in different parts of their receptive field. Dual recordings from pairs of VS cells show that they are electrically coupled. This coupling is responsible for the elongated horizontal extent of their receptive fields. VS cells with a lateral receptive field have additional connections to a VS cell with a frontal receptive field and to the horizontal system, tuning these cells to rotational flow fields. In summary, the receptive field of these cells consists of two components: one that they receive from local motion detectors on their dendrite, and one that they import from other large-field neurons.

Journal ArticleDOI
TL;DR: Cortical mechanisms sensitive to chromatic contrast seem to provide little input to the suppressive surrounds of V1 neurons but substantial input to those of V2 neurons, and the chromatic properties of the CRF in the presence of differently colored surrounds were measured.
Abstract: Stimulation of the suppressive surround of a cortical neuron affects the responsivity and tuning of the classical receptive field (CRF) on several stimulus dimensions. In V1 and V2 of macaques prepared for acute electrophysiological experiments, we explored the chromatic sensitivity of the surround and its influence on the chromatic tuning of the CRF. We studied receptive fields of single neurons with patches of drifting grating of optimal spatial frequency and orientation and variable size, modulated along achromatic or isoluminant color directions. The responses of most neurons declined as the patch was enlarged beyond the optimal size (surround suppression). In V1 the suppression evoked by isoluminant gratings was less than one-half that evoked by achromatic gratings. Consequently, many cells were most sensitive to achromatic modulation when patches just covered the CRF but were most sensitive to isoluminant modulation when patches were enlarged to cover the suppressive surround. Non-oriented neurons that were strongly chromatically opponent generally lacked suppressive surrounds. In V2 most neurons showed equal surround suppression from isoluminant gratings and achromatic gratings. This makes the relative sensitivity of V2 neurons to achromatic and isoluminant gratings mainly independent of the size of the grating. We also measured the chromatic properties of the CRF in the presence of differently colored surrounds. In neither V1 nor V2 did the surround alter the chromatic tuning of the CRF. Cortical mechanisms sensitive to chromatic contrast seem to provide little input to the suppressive surrounds of V1 neurons but substantial input to those of V2 neurons.

Journal ArticleDOI
TL;DR: It is suggested that a moving stimulus provides ‘preactivation’ that allows more rapid processing than for a single flash event.
Abstract: Psychophysical evidence in humans indicates that localization is different for stationary flashed and coherently moving objects. To address how the primary visual cortex represents object position we used a population approach that pools spiking activity of many neurones in cat area 17. In response to flashed stationary squares (0.4 deg) we obtained localized activity distributions in visual field coordinates, which we referred to as profiles across a ‘population receptive field’ (PRF). We here show how motion trajectories can be derived from activity across the PRF and how the representation of moving and flashed stimuli differs in position. We found that motion was represented by peaks of population activity that followed the stimulus with a speed-dependent lag. However, time-to-peak latencies were shorter by ∼16 ms compared to the population responses to stationary flashes. In addition, motion representation showed a directional bias, as latencies were more reduced for peripheral-to-central motion compared to the opposite direction. We suggest that a moving stimulus provides ‘preactivation’ that allows more rapid processing than for a single flash event.