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Neurons in the primary visual cortex respond to oriented stimuli placed in the center of their receptive field, yet their response is modulated by stimuli outside the receptive field (the surround). Classically, this surround modulation is assumed to be strongest if the orientation of the surround stimulus aligns with the neuron's preferred orientation, irrespective of the actual center stimulus. This neuron-dependent surround modulation has been used to explain a wide range of psychophysical phenomena, such as biased tilt perception and saliency of stimuli with contrasting orientation. However, several neurophysiological studies have shown that for most neurons surround modulation is instead center dependent: it is strongest if the surround orientation aligns with the center stimulus. As the impact of such center-dependent modulation on the population level is unknown, we examine this using computational models. We find that with neuron-dependent modulation the biases in orientation coding, commonly used to explain the tilt illusion, are larger than psychophysically reported, but disappear with center-dependent modulation. Therefore we suggest that a mixture of the two modulation types is necessary to quantitatively explain the psychophysically observed biases. Next, we find that under center-dependent modulation average population responses are more sensitive to orientation differences between stimuli, which in theory could improve saliency detection. However, this effect depends on the specific saliency model. Overall, our results thus show that center-dependent modulation reduces coding bias, while possibly increasing the sensitivity to salient features. [ABSTRACT FROM AUTHOR]

Activity of cortical inhibitory interneurons is rapidly reduced in response to monocular deprivation during the critical period for ocular dominance plasticity and in response to salient events encountered during learning. In the case of primary sensory cortex, a decrease in mean evoked firing rate of parvalbumin-positive (PV) inhibitory neurons is causally linked to a reorganization of excitatory networks following sensory perturbation. Converging evidence indicates that it is deprivation, and not an imbalance between open- and closed-eye inputs, that triggers rapid plasticity in PV neurons. However, this has not been directly tested in vivo. Using two-photon guided cell-attached recording, we examined the impact of closing both eyes for 24 h on PV neuron response properties in mouse primary visual cortex. We found that binocular deprivation induces a 30% reduction in stimulus-evoked mean firing rate and that this reduction is specific to critical period-aged mice. The number of PV neurons showing detectable tuning to orientation increased after 24 h of deprivation, and this effect was also specific to critical period-aged mice. In contrast to evoked mean firing rate and orientation tuning, measurements of trial-to-trial variability revealed that stimulus-driven decreases in variability are significantly dampened by deprivation during both the critical period and the postcritical period. These data establish that open-eye inputs are not required to drive deprivation-induced weakening of PV neuron evoked activity and that other aspects of in vivo PV neuron activity are malleable throughout life. [ABSTRACT FROM AUTHOR]

Lymph drainage and propulsion are sustained by an extrinsic mechanism, based on mechanical forces acting from the surrounding tissues against the wall of lymphatic vessels, and by an intrinsic mechanism attributable to active spontaneous contractions of the lymphatic vessel muscle. Despite being heterogeneous, the mechanisms underlying the generation of spontaneous contractions share a common biochemical nature and are thus modulated by temperature. In this study, we challenged excised tissues from rat diaphragm and hindpaw, endowed with spontaneously contracting lymphatic vessels, to temperatures from 24°C (hindpaw) or 33°C (diaphragmatic vessels) to 40°C while measuring lymphatic contraction frequency (fc) and amplitude. Both vessel populations displayed a sigmoidal relationship between fc and temperature, each centered around the average temperature of surrounding tissue (36.7 diaphragmatic and 32.1 hindpaw lymphatics). Although the slope factor of the sigmoidal fit to the fc change of hindpaw vessels was 2.3°C·cycles-1·min-1, a value within the normal range displayed by simple biochemical reactions, the slope factor of the diaphragmatic lymphatics was 0.62°C·cycles-1·min-1, suggesting the added involvement of temperature-sensing mechanisms. Lymph flow calculated as a function of temperature confirmed the relationship observed on fc data alone and showed that none of the two lymphatic vessel populations would be able to adapt to the optimal working temperature of the other tissue district. This poses a novel question whether lymphatic vessels might not adapt their function to accommodate the change if exposed to a surrounding temperature, which is different from their normal condition. [ABSTRACT FROM AUTHOR]

Contrast sensitivity is fundamental to natural visual processing and an important tool for characterizing both visual function and clinical disorders. We simultaneously measured contrast sensitivity and neural contrast response functions and compared measurements in common laboratory conditions with naturalistic conditions. In typical experiments, a subject holds fixation and a stimulus is flashed on, whereas in natural vision, saccades bring stimuli into view. Motivated by our previous V1 findings, we tested the hypothesis that perceptual contrast sensitivity is lower in natural vision and that this effect is associated with corresponding changes in V1 activity. We found that contrast sensitivity and V1 activity are correlated and that the relationship is similar in laboratory and naturalistic paradigms. However, in the more natural situation, contrast sensitivity is reduced up to 25% compared with that in a standard fixation paradigm, particularly at lower spatial frequencies, and this effect correlates with significant reductions in V1 responses. Our data suggest that these reductions in natural vision result from fast adaptation on one fixation that lowers the response on a subsequent fixation. This is the first demonstration of rapid, natural-image adaptation that carries across saccades, a process that appears to constantly influence visual sensitivity in natural vision. [ABSTRACT FROM AUTHOR]

Whisker trimming causes substantial reorganization of neuronal response properties in barrel cortex. However, little is known about experience-dependent rerouting of sensory processing following sensory deprivation. To address this, we performed in vivo intracellular recordings from layers 2/3 (L2/3), layer 4 (L4), layer 5 regular-spiking (L5RS), and L5 intrinsically bursting (L5IB) neurons and measured their multiwhisker receptive field at the level of spiking activity, membrane potential, and synaptic conductance before and after sensory deprivation. We used Chernoff information to quantify the "sensory information" contained in the firing patterns of cells in response to spared and deprived whisker stimulation. In the control condition, information for flanking-row and same-row whiskers decreased in the order L4, L2/3, L5IB, L5RS. However, after whisker-row deprivation, spared flanking-row whisker information was reordered to L4, L5RS, L5IB, L2/3. Sensory information from the trimmed whiskers was reduced and delayed in L2/3 and L5IB neurons, whereas sensory information from spared whiskers was increased and advanced in L4 and L5RS neurons. Sensory information from spared whiskers was increased in L5IB neurons without a latency change. L5RS cells exhibited the largest changes in sensory information content through an atypical plasticity combining a significant decrease in spontaneous activity and an increase in a short-latency excitatory conductance. [ABSTRACT FROM AUTHOR]

Previous neurophysiological studies performed in macaque monkeys have revealed complex somatosensory responses in the secondary somatosensory area (SII), such as large receptive fields (RFs), as well as bilateral ones. However, systematic analyses of neurons with large RFs have not been performed. In the present study, we recorded single-unit activities in SII of awake macaque monkeys to investigate systematically large RFs by dividing the whole body into four body regions (head, trunk, forelimb, and hindlimb). Recorded neurons were classified into two types, according to whether the RFs were confined to one body region: single (n = 817) and combined (n = 282) body-region types. These two types were distinct in terms of the percentage of bilateral RFs: 55% in the single-region type and 90% in the combined type, demonstrating that two types of RF enlargement occur simultaneously in the combined type, namely, RF convergence from different body regions and RF convergence from both hemibodies. Among the combined-type RFs, two tendencies of RF convergence were found: 1) the distal parts of the limbs (i.e., hand and foot) and the mouth are interconnected, and 2) the trunk RFs extend continuously toward the distal parts of the limb and head to cover the entire body surface. Our distribution analysis on unfolded maps clarified that neurons having RFs with these two tendencies were distributed within specific subregions in SII. [ABSTRACT FROM AUTHOR]

The neuronal representation of luminance contrast has not been thoroughly studied in birds. Here we present a detailed quantitative analysis of the contrast response of 120 individual neurons recorded from the visual wulst of awake burrowing owls (Athene cunicularia). Stimuli were sine-wave gratings presented within the cell classical receptive field and optimized in terms of eye preference, direction of drift, and spatiotemporal frequency. As contrast intensity was increased from zero to near 100%, most cells exhibited a monotonic response profile with a compressive, at times saturating, nonlinearity at higher contrasts. However, contrast response functions were found to have a highly variable shape across cells. With the view to capture a systematic trend in the data, we assessed the performance of four plausible models (linear, power, logarithmic, and hyperbolic ratio) using classical goodness-of-fit measures and more rigorous statistical tools for multimodel inferences based on the Akaike information criterion. From this analysis, we conclude that a high degree of model uncertainty is present in our data, meaning that no single descriptor is able on its own to capture the heterogeneous nature of single-unit contrast responses in the wulst. We further show that the generalizability of the hyperbolic ratio model established, for example, in the primary visual cortex of cats and monkeys is not tenable in the owl wulst mainly because most neurons in this area have a much wider dynamic range that starts at low contrast. The challenge for future research will be to understand the functional implications of these findings. [ABSTRACT FROM AUTHOR]

Neurons in area MT/V5 of the macaque visual cortex encode visual motion. Some cells are selective for the motion of oriented features (component direction-selective, CDS); others respond to the true direction of complex patterns (pattern-direction selective, PDS). There is a continuum of selectivity in MT, with CDS cells at one extreme and PDS cells at the other; we compute a pattern index that captures this variation. It is unknown how a neuron's pattern index is related to its other tuning characteristics. We therefore analyzed the responses of 792 MT cells recorded in the course of other experiments from opiate-anesthetized macaque monkeys, as a function of the direction, spatial frequency, drift rate, size, and contrast of sinusoidal gratings and of the direction and speed of random-dot textures. We also compared MT responses to those of 718 V1 cells. As expected, MT cells with higher pattern index tended to have stronger direction selectivity and broader direction tuning to gratings, and they responded better to plaids than to gratings. Strongly PDS cells also tended to have smaller receptive fields and stronger surround suppression. Interestingly, they also responded preferentially to higher drift rates and higher speeds of moving dots. The spatial frequency preferences of PDS cells depended strongly on their preferred temporal frequencies, whereas these preferences were independent in component-selective cells. Pattern direction selectivity is statistically associated with many response properties of MT cells but not strongly associated with any particular property. Pattern-selective signals are thus available in association with most other signals exported by MT. [ABSTRACT FROM AUTHOR]

Attending to a stimulus modulates the responses of sensory neurons that represent features of that stimulus, a phenomenon named "feature attention." For example, attending to a stimulus containing upward motion enhances the responses of upward-preferring direction-selective neurons in the middle temporal area (MT) and suppresses the responses of downward-preferring neurons, even when the attended stimulus is outside of the spatial receptive fields of the recorded neurons (Treue S, Martinez-Trujillo JC. Nature 399: 575-579, 1999). This modulation renders the representation of sensory information across a neuronal population more selective for the features present in the attended stimulus (Martinez-Trujillo JC, Treue S. Curr Biol 14: 744-751, 2004). We hypothesized that if feature attention modulates neurons according to their tuning preferences, it should also be sensitive to their tuning strength, which is the magnitude of the difference in responses to preferred and null stimuli. We measured how the effects of feature attention on MT neurons in rhesus monkeys (Macaca mulatta) depended on the relationship between features--in our case, direction of motion and binocular disparity--of the attended stimulus and a neuron's tuning for those features. We found that, as for direction, attention to stimuli containing binocular disparity cues modulated the responses of MT neurons and that the magnitude of the modulation depended on both a neuron's tuning preferences and its tuning strength. Our results suggest that modulation by feature attention may depend not just on which features a neuron represents but also on how well the neuron represents those features. [ABSTRACT FROM AUTHOR]

Some neurons in early visual cortex are highly selective for the position of oriented edges in their receptive fields (simple cells), whereas others are largely position insensitive (complex cells). These characteristics are reflected in their sensitivity to the spatial phase of moving sine-wave gratings: simple cell responses oscillate at the fundamental frequency of the stimulus, whereas this is less so for complex cells. In primates, when assessed at high stimulus contrast, simple cells and complex cells are roughly equally represented in the first visual cortical area, V1, whereas in the second visual area, V2, the majority of cells are complex. Recent evidence has shown that phase sensitivity of complex cells is contrast dependent. This has led to speculation that reduced contrast may lead to changes in the spatial structure of receptive fields, perhaps due to changes in how feedforward and recurrent signals interact. Given the substantial interconnections between V1 and V2 and recent evidence for the emergence of unique functional capacity in V2, we assess the relationship between contrast and phase sensitivity in the two brain regions. We show that a substantial proportion of complex cells in macaque V1 exhibit significant increases in phase sensitivity at low contrast, whereas this is rarely observed in V2. Our results support a degree of hierarchical processing from V1 to V2 with the differences possibly relating to the fact that V1 combines both subcortical and cortical input, whereas V2 receives input purely from cortical circuits. [ABSTRACT FROM AUTHOR]

Performance is often improved when targets are presented in space near the hands rather than far from the hands. Performance in hand-near space may be improved because participants can use proprioception from the nearby limb and hand to provide a narrower and more resolute frame of reference. An equally compelling alternative is that targets appearing near the hand fall within the receptive fields of visual-tactile bimodal cells, recruiting them to assist in the visual representation of targets that appear near but not far from the hand. We distinguished between these two alternatives by capitalizing on research showing that vision and proprioception have differential effects on the precision of target representation (van Beers RJ, Sittig AC, Denier van der Gon JJ. Exp Brain Res 122: 367-377, 1998). Participants performed an in-tocenter reaching task to an array of central target locations with their right hand, while their left hand rested near (beneath) or far from the target array. Reaching end-point accuracy, variability, time, and speed were assessed. We predicted that if proprioception contributes to the representation of hand-near targets, then error variability in depth will be smaller in the hand-near condition than in the hand-far condition. By contrast, if vision contributes to the representation of hand-near targets, then error variability along the lateral dimension will be smaller in the hand-near than in the hand-far condition. Our results showed that the placement of the hand near the targets reduced end-point error variability along the lateral dimension only. The results suggest that hand-near targets are represented with greater visual resolution than far targets. [ABSTRACT FROM AUTHOR]

Somatosensory processing in the anesthetized macaque monkey was examined by reversibly deactivating posterior parietal areas 5L and 7b and motor/premotor cortex (M1/ PM) with microfluidic thermal regulators developed by our laboratories. We examined changes in receptive field size and configuration for neurons in areas 1 and 2 that occurred during and after cooling deactivation. Together the deactivated fields and areas 1 and 2 form part of a network for reaching and grasping in human and nonhuman primates. Cooling area 7b had a dramatic effect on receptive field size for neurons in areas 1 and 2, while cooling area 5 had moderate effects and cooling M1/PM had little effect. Specifically, cooling discrete locations in 7b resulted in expansions of the receptive fields for neurons in areas 1 and 2 that were greater in magnitude and occurred in a higher proportion of sites than similar changes evoked by cooling the other fields. At some sites, the neural receptive field returned to the precooling configuration within 5-22 min of rewarming, but at other sites changes in receptive fields persisted. These results indicate that there are profound top-down influences on sensory processing of early cortical areas in the somatosensory cortex. [ABSTRACT FROM AUTHOR]

We simultaneously compared the sensitivity of single primary afferent neurons supplying the glabrous skin of the hand and the psychophysical amplitude discrimination thresholds in human subjects for a set of vibrotactile stimuli delivered to the receptive field. All recorded afferents had a dynamic range narrower than the range of amplitudes across which the subjects could discriminate. However, when the vibration amplitude was chosen to be within the steepest part of the afferent's stimulus-response function the response of single afferents, defined as the spike count over the vibration duration (500 ms), was often more sensitive in discriminating vibration amplitude than the perceptual judgment of the participants. We quantified how the neuronal performance depended on the integration window: for short windows the neuronal performance was inferior to the performance of the subject. The neuronal performance progressively improved with increasing spike count duration and reached a level significantly above that of the subjects when the integration window was 250 ms or longer. The superiority in performance of individual neurons over observers could reflect a nonoptimal integration window or be due to the presence of noise between the sensory periphery and the cortical decision stage. Additionally, it could indicate that the range of perceptual sensitivity comes at the cost of discrimination through pooling across neurons with different response functions. [ABSTRACT FROM AUTHOR]

The midlevel visual cortical area V4 in the primate is thought to be critical for the neural representation of visual shape. Several studies agree that V4 neurons respond to contour features, e.g., convexities and concavities along a shape boundary, that are more complex than the oriented segments encoded by neurons in the primary visual cortex. Here we compare two distinct approaches to modeling V4 shape selectivity: one based on a spectral receptive field (SRF) map in the orientation and spatial frequency domain and the other based on a map in an object-centered angular position and contour curvature space. We test the ability of these two characterizations to account for the responses of V4 neurons to a set of parametrically designed two-dimensional shapes recorded previously in the awake macaque. We report two lines of evidence suggesting that the SRF model does not capture the contour sensitivity of V4 neurons. First, the SRF model discards spatial phase information, which is inconsistent with the neuronal data. Second, the amount of variance explained by the SRF model was significantly less than that explained by the contour curvature model. Notably, cells best fit by the curvature model were poorly fit by the SRF model, the latter being appropriate for a subset of V4 neurons that appear to be orientation tuned. These limitations of the SRF model suggest that a full understanding of midlevel shape representation requires more complicated models that preserve phase information and perhaps deal with object segmentation. [ABSTRACT FROM AUTHOR]

Multiple visual stimuli are common in natural scenes, yet it remains unclear how multiple stimuli interact to influence neuronal responses. We investigated this question by manipulating relative signal strengths of two stimuli moving simultaneously within the receptive fields (RFs) of neurons in the extrastriate middle temporal (MT) cortex. Visual stimuli were overlapping random-dot patterns moving in two directions separated by 90°. We first varied the motion coherence of each random-dot pattern and characterized, across the direction tuning curve, the relationship between neuronal responses elicited by bidirectional stimuli and by the constituent motion components. The tuning curve for bidirectional stimuli showed response normalization and can be accounted for by a weighted sum of the responses to the motion components. Allowing nonlinear, multiplicative interaction between the two component responses significantly improved the data fit for some neurons, and the interaction mainly had a suppressive effect on the neuronal response. The weighting of the component responses was not fixed but dependent on relative signal strengths. When two stimulus components moved at different coherence levels, the response weight for the higher-coherence component was significantly greater than that for the lower-coherence component. We also varied relative luminance levels of two coherently moving stimuli and found that MT response weight for the higher-luminance component was also greater. These results suggest that competition between multiple stimuli within a neuron's RF depends on relative signal strengths of the stimuli and that multiplicative nonlinearity may play an important role in shaping the response tuning for multiple stimuli [ABSTRACT FROM AUTHOR]

Intrinsic sensory neurons (ISNs) of the enteric nervous system respond to stimuli such as muscle tension, muscle length, distortion of the mucosa, and the chemical content in the lumen. ISNs form recurrent networks that probably drive many intestinal motor patterns and reflexes. ISNs express a large number of voltage- and calcium-gated ion channels, some of which are modified by inflammation or repeated physiological stimuli, but how interactions between different ionic currents in ISNs produce both normal and pathological behaviors in the intestine remains unclear. We constructed a model of ISNs including voltagegated sodium and potassium channels, N-type calcium channels, big conductance calcium-dependent potassium (BK) channels, calciumdependent nonspecific cation channels (NSCa), intermediate conductance calcium-dependent potassium (IK) channels, hyperpolarizationactivated cation (Ih) channels, and internal calcium dynamics. The model was based on data from the literature and our electrophysiological studies. The model reproduced responses to short or long depolarizing current pulses and responses to long hyperpolarizing current pulses. Sensitivity analysis showed that Ih, IK, NSCa, and BK have the largest influence on the number of action potentials observed during prolonged depolarizations. The model also predicts that changes to the voltage of activation for Ih have a large influence on excitability, but changes to the time constant of activation for Ih have a minor effect. Our model identifies how interactions between different iconic currents influence the excitability of ISNs and highlights an important role for Ih in enteric neuroplasticity resulting from disease. [ABSTRACT FROM AUTHOR]

Activities of nearby cortical cells show temporal correlation on many timescales. In particular, previous studies of primary visual cortex (V1) indicate precise correlation on a timescale of milliseconds and loose correlation on a timescale of tens of milliseconds. To characterize cortical organization of these correlations, we investigated their spatial extent, laminar organization, and dependence on receptive field (RF) similarities. By simultaneously recording neuronal activity across layers within a horizontal distance of 1.2 mm, we found that loose correlation was widely observed for neuronal pairs horizontally or vertically separated over the whole distance range regardless of the layers. The incidence of loose correlation tended to be lower in layer 4 than in other layers. Loose correlation also accompanied a consistent delay in firing that was monotonically related to the vertical, but not horizontal, distance between the paired neurons. In contrast, the spatial range in which precise correlation was observed was more limited, with its incidence dropping sharply within 0.4 mm in both vertical and horizontal directions for all layers. With these spatial ranges, precise correlation was typically observed for pairs of neurons in the same layers, while loose correlation was often present even for pairs of neurons in widely separated layers. Furthermore, precise correlation was predominantly seen for pairs with similar RF properties, whereas loose correlation was seen even in pairs showing dissimilar properties. Our results show that neuronal correlations in V1 show markedly different structures for horizontal and vertical dimensions depending on correlation timescales. [ABSTRACT FROM AUTHOR]

The ventral intraparietal area (VIP) processes multisensory visual, vestibular, tactile, and auditory signals in diverse reference frames. We recently reported that visual heading signals in VIP are represented in an approximately eye-centered reference frame when measured using large-field optic flow stimuli. No VIP neuron was found to have head-centered visual heading tuning, and only a small proportion of cells had reference frames that were intermediate between eye- and head-centered. In contrast, previous studies using moving bar stimuli have reported that visual receptive fields (RFs) in VIP are head-centered for a substantial proportion of neurons. To examine whether these differences in previous findings might be due to the neuronal property examined (heading tuning vs. RF measurements) or the type of visual stimulus used (full-field optic flow vs. a single moving bar), we have quantitatively mapped visual RFs of VIP neurons using a large-field, multipatch, random-dot motion stimulus. By varying eye position relative to the head, we tested whether visual RFs in VIP are represented in head- or eye-centered reference frames. We found that the vast majority of VIP neurons have eye-centered RFs with only a single neuron classified as head-centered and a small minority classified as intermediate between eye- and head-centered. Our findings suggest that the spatial reference frames of visual responses in VIP may depend on the visual stimulation conditions used to measure RFs and might also be influenced by how attention is allocated during stimulus presentation. [ABSTRACT FROM AUTHOR]

The stereo correspondence problem poses a challenge to visual neurons because localized receptive fields potentially cause false responses. Neurons in the primary visual cortex (V1) partially resolve this problem by combining excitatory and suppressive responses to encode binocular disparity. We explored the time course of this combination in awake, monkey V1 neurons using subspace mapping of receptive fields. The stimulus was a binocular noise pattern constructed from discrete spatial frequency components. We forward correlated the firing of the V1 neuron with the occurrence of binocular presentations of each spatial frequency component. The forward correlation yielded a complete set of response time courses to every combination of spatial frequency and interocular phase difference. Some combinations produced suppressive responses. Typically, if an interocular phase difference for a given spatial frequency produced strong excitation, we saw suppression in response to the opposite interocular phase difference at lower spatial frequencies. The suppression was delayed relative to the excitation, with a median difference in latency of 7 ms. We found that the suppressive mechanism explains a well-known mismatch of monocular and binocular signals. The suppressive components increased power at low spatial frequencies in disparity tuning, whereas they reduced the monocular response to low spatial frequencies. This long-recognized mismatch of binocular and monocular signals reflects a suppressive mechanism that helps reduce the response to false matches. [ABSTRACT FROM AUTHOR]

We explore the visual world by making rapid eye movements (saccades) to focus on objects and locations of interest. Despite abrupt retinal image shifts, we see the world as stable. Remapping contributes to visual stability by updating the internal image with every saccade. Neurons in macaque lateral intraparietal cortex (LIP) and other brain areas update information about salient locations around the time of a saccade. The depth of information transfer remains to be thoroughly investigated. Area LIP, as part of the dorsal visual stream, is regarded as a spatially selective area, yet there is evidence that LIP neurons also encode object features. We sought to determine whether LIP remaps shape information. This knowledge is important for understanding what information is retained from each glance. We identified 82 remapping neurons. First, we presented shapes within the receptive field and tested for shape selectivity in a fixation task. Among the remapping neurons, 28 neurons (34%) were selective for shape. Second, we presented the same shapes in the future location of the receptive field around the time of the saccade and tested for shape selectivity during remapping. Thirty-one (38%) neurons were selective for shape. Of 11 neurons that were shape selective in both tasks, 5 showed significant correlation between shape selectivity in the two tasks. Across the population, there was a weak but significant correlation between responses to shape in the two tasks. Our results provide neurophysiological evidence that remapped responses in area LIP can encode shape information as well as spatial information. [ABSTRACT FROM AUTHOR]

The neural response to a sensory stimulus tends to be more strongly reduced when the stimulus is preceded by the same, rather than a different, stimulus. This stimulus-specific adaptation (SSA) is ubiquitous across the senses. In hearing, SSA has been suggested to play a role in change detection as indexed by the mismatch negativity. This study sought to test whether SSA, measured in human auditory cortex, is caused by neural fatigue (reduction in neural responsiveness) or by sharpening of neural tuning to the adapting stimulus. For that, we measured event-related cortical potentials to pairs of pure tones with varying frequency separation and stimulus onset asynchrony (SOA). This enabled us to examine the relationship between the degree of specificity of adaptation as a function of frequency separation and the rate of decay of adaptation with increasing SOA. Using simulations of tonotopic neuron populations, we demonstrate that the fatigue model predicts independence of adaptation specificity and decay rate, whereas the sharpening model predicts interdependence. The data showed independence and thus supported the fatigue model. In a second experiment, we measured adaptation specificity after multiple presentations of the adapting stimulus. The multiple adapters produced more adaptation overall, but the effect was more specific to the adapting frequency. Within the context of the fatigue model, the observed increase in adaptation specificity could be explained by assuming a 2.5-fold increase in neural frequency selectivity. We discuss possible bottom-up and top-down mechanisms of this effect. [ABSTRACT FROM AUTHOR]

Recently, it has been recognized that a single airway sensory unit may contain multiple receptive fields and that each field houses at least one encoder. Since some units respond to both lung inflation and deflation, we hypothesized that these units contain heterogeneous encoders for sensing inflation and deflation, respectively. Single unit activities were recorded from the cervical vagus nerve in anesthetized, open chest, and mechanically ventilated rabbits. Fifty-two airway sensory units with multiple receptive fields that responded to both lung inflation and deflation were identified. Among them, 13 units had separate receptive fields for inflation and deflation, where one of the fields could be blocked by local injection of 2% lidocaine (10 μl). In 8 of the 13 units, the deflation response was blocked without affecting the unit's response to inflation, whereas in the remaining five units, the inflation response was blocked without affecting the deflation response. Our results support the hypothesis that a single mechanosensory unit may contain heterogeneous encoders that can respond to either inflation or deflation. [ABSTRACT FROM AUTHOR]

It is widely claimed that interactions among simultaneously presented visual stimuli are suppressive and that these interactions primarily occur when stimuli fall within the same receptive field (Desimone and Duncan 1995). Here, we show evidence for a novel form of interaction between simultaneously presented but distant stimuli that does not fit either pattern. To examine interactions between simultaneously presented stimuli, we measure the response to a single stimulus as a function of whether or not other stimuli are also presented simultaneously, and we further ask how the response to a given stimulus is affected by whether the simultaneously present stimuli are identical or different from each other. Our method reveals a new phenomenon of "redundancy gain:" the visual response to a stimulus is higher when accompanied by identical stimuli than when that stimulus is presented alone, even though the stimuli are displayed in separate visual quadrants. This pattern is observed throughout the visual hierarchy, including V1 and V2, and we show that it is distinct from the wellknown simultaneous suppression effect (Kastner et al. 1998). We propose that the redundancy gain in early retinotopic cortex results from feedback from higher visual areas and may underlie perceptual averaging and other ensemble coding phenomena observed behaviorally. [ABSTRACT FROM AUTHOR]

Responses of many neurons in primary visual cortex (V1) are suppressed by stimuli exceeding the classical receptive field (RF), an important property that might underlie the computation of visual saliency. Traditionally, it has proven difficult to disentangle the underlying neural circuits, including feedforward, horizontal intracortical, and feedback connectivity. Since circuit-level analysis is particularly feasible in the mouse, we asked whether neural signatures of spatial integration in mouse V1 are similar to those of higher-order mammals and investigated the role of parvalbumin-expressing (PV+) inhibitory interneurons. Analogous to what is known from primates and carnivores, we demonstrate that, in awake mice, surround suppression is present in the majority of V1 neurons and is strongest in superficial cortical layers. Anesthesia with isoflurane-urethane, however, profoundly affects spatial integration: it reduces the laminar dependency, decreases overall suppression strength, and alters the temporal dynamics of responses. We show that these effects of brain state can be parsimoniously explained by assuming that anesthesia affects contrast normalization. Hence, the full impact of suppressive influences in mouse V1 cannot be studied under anesthesia with isoflurane-urethane. To assess the neural circuits of spatial integration, we targeted PV+ interneurons using optogenetics. Optogenetic depolarization of PV+ interneurons was associated with increased RF size and decreased suppression in the recorded population, similar to effects of lowering stimulus contrast, suggesting that PV+ interneurons contribute to spatial integration by affecting overall stimulus drive. We conclude that the mouse is a promising model for circuit-level mechanisms of spatial integration, which relies on the combined activity of different types of inhibitory interneurons. [ABSTRACT FROM AUTHOR]

The archer fish is well known for its extreme visual behavior in shooting water jets at prey hanging on vegetation above water. This fish is a promising model in the study of visual system function because it can be trained to respond to artificial targets and thus to provide valuable psychophysical data. Although much behavioral data have indeed been collected over the past two decades, little is known about the functional organization of the main visual area supporting this visual behavior, namely, the fish optic tectum. In this article we focus on a fundamental aspect of this functional organization and provide a detailed analysis of receptive field properties of cells in the archer fish optic tectum. Using extracellular measurements to record activities of single cells, we first measure their retinotectal mapping. We then determine their receptive field properties such as size, selectivity for stimulus direction and orientation, tuning for spatial frequency, and tuning for temporal frequency. Finally, on the basis of all these measurements, we demonstrate that optic tectum cells can be classified into three categories: orientation-tuned cells, direction-tuned cells, and direction-agnostic cells. Our results provide an essential basis for future investigations of information processing in the archer fish visual system. [ABSTRACT FROM AUTHOR]

A key property of neurons in primary visual cortex (V1) is the distinction between simple and complex cells. Recent reports in cat visual cortex indicate the categorization of simple and complex can change depending on stimulus conditions. We investigated the stability of the simple/complex classification with changes in drive produced by either contrast or modulation by the extraclassical receptive field (eCRF). These two conditions were reported to increase the proportion of simple cells in cat cortex. The ratio of the modulation depth of the response (F1) to the elevation of response (F0) to a drifting grating (F1/F0 ratio) was used as the measure of simple/complex. The majority of V1 complex cells remained classified as complex with decreasing contrast. Near contrast threshold, an equal proportion of simple and complex cells changed their classification. The F1/F0 ratio was stable between optimal and large stimulus areas even for those neurons that showed strong eCRF suppression. There was no discernible overall effect of surrounding spatial context on the F1/F0 ratio. Simple/complex cell classification is relatively stable across a range of stimulus drives, produced by either contrast or eCRF suppression. [ABSTRACT FROM AUTHOR]

Local field potentials (LFPs) recorded in the auditory cortex of mammals are known to reveal weakly selective and often multimodal spectrotemporal receptive fields in contrast to spiking activity. This may in part reflect the wider “listening sphere” of LFPs relative to spikes due to the greater current spread at low than high frequencies. We recorded LFPs and spikes from auditory cortex of guinea pigs using 16-channel electrode arrays. LFPs were processed by a component analysis technique that produces optimally tuned linear combinations of electrode signals. Linear combinations of LFPs were found to have sharply tuned responses, closer to spike-related tuning. The existence of a sharply tuned component implies that a cortical neuron (or group of neurons) capable of forming a linear combination of its inputs has access to that information. Linear combinations of signals from electrode arrays reveal information latent in the subspace spanned by multichannel LFP recordings and are justified by the fact that the observations themselves are linear combinations of neural sources. [ABSTRACT FROM AUTHOR]

Many migrating insects rely on the plane of sky polarization as a cue to detect spatial directions. Desert locusts (Schistocerca gregaria), like other insects, perceive polarized light through specialized photoreceptors in a dorsal eye region. Desert locusts occur in two phases: a gregarious swarming phase, which migrates during the day, and a solitarious nocturnal phase. Neurons in a small brain area, the anterior optic tubercle (AOTu), are critically involved in processing polarized light in the locust brain. While polarization-sensitive intertubercle cells [lobula-tubercle neuron 1 (LoTu1) and tubercle-tubercle neuron 1 (TuTu1)] interconnect the AOTu of both hemispheres, tubercle-lateral accessory lobe tract (TuLAL1) neurons transmit sky compass signals to a polarization compass in the central brain. To better understand the neural network underlying polarized light processing in the AOTu and to investigate possible adaptations of the polarization vision system to a diurnal versus nocturnal lifestyle, we analyzed receptive field properties, intensity-response relationships, and daytime dependence of responses of AOTu neurons in gregarious and solitarious locusts. Surprisingly, no differences in the physiology of these neurons were found between the two locust phases. Instead, clear differences were observed between the different types of AOTu neurons. Whereas TuTu1 and TuLAL1 neurons encoded E-vector orientation independent of light intensity and would thus be operational in bright daylight, LoTu1 neurons were inhibited by high light intensity and provided strong polarization signaling only under dim light conditions. The presence of high- and low-intensity polarization channels might, therefore, allow solitarious and gregarious locusts to use the same polarization coding system despite their different activity cycles. [ABSTRACT FROM AUTHOR]

Maturation of the nervous system results in changes in both central and peripheral processing. To better understand responses to injury in the young, developmental differences in the acute response to incision were investigated in both tactile and nociceptive myelinated peripheral mechanosensitive afferent neurons in vivo. Neuronal intrasomal recordings were performed in juvenile and infant rats in 34 L5 dorsal root ganglia, and each neuron was phenotypically defined. Neurons had a mechanosensitive receptive field in the glabrous skin on the plantar surface of the hind paw, which was characterized at baseline and for up to 45 min after incision. Fundamental maturational differences in the effect of incision were clear: in high-threshold nociceptive mechanoreceptors, the mechanical threshold decreased immediately and the receptive field size increased rapidly in juvenile rats but not in infant rats. Additionally, a divergence in changes in the instantaneous response frequency of tactile afferents occurred between the two ages. These differences may help explain maturational differences in responses to peripheral injury and suggest that differences in central nervous system responses may be partially mitigated by spatially confined and frequency-dependent differences resulting from tactile and nociceptive mechanosensitive input. [ABSTRACT FROM AUTHOR]

In the primary auditory cortex, spectrotemporal receptive fields (STRFs) are composed of multiple independent components that capture the processing of disparate stimulus aspects by any given neuron. The origin of these multidimensional stimulus filters in the central auditory system is unknown. To determine whether multicomponent STRFs emerge prior to the forebrain, we recorded from single neurons in the main obligatory station of the auditory midbrain, the inferior colliculus. By comparing results of different spike-triggered techniques, we found that the neural responses in the inferior colliculus can be accounted for by a single stimulus filter. This was observed for all temporal response patterns, from strongly phasic to tonic. Our results reveal that spectrotemporal stimulus encoding undergoes a fundamental transformation along the auditory neuraxis, with the emergence of multidimensional receptive fields beyond the auditory midbrain. [ABSTRACT FROM AUTHOR]

Orientation selectivity and its development are basic features of visual cortex. The original model of orientation selectivity proposes that elongated simple cell receptive fields are constructed from convergent input of an array of lateral geniculate nucleus neurons. However, orientation selectivity of simple cells in the visual cortex is generally greater than the linear contributions based on projections from spatial receptive field profiles. This implies that additional selectivity may arise from intracortical mechanisms. The hierarchical processing idea implies mainly linear connections, whereas cortical contributions are generally considered to be nonlinear. We have explored development of orientation selectivity in visual cortex with a focus on linear and nonlinear factors in a population of anesthetized 4-wk postnatal kittens and adult cats. Linear contributions are estimated from receptive field maps by which orientation tuning curves are generated and bandwidth is quantified. Nonlinear components are estimated as the magnitude of the power function relationship between responses measured from drifting sinusoidal gratings and those predicted from the spatial receptive field. Measured bandwidths for kittens are slightly larger than those in adults, whereas predicted bandwidths are substantially broader. These results suggest that relatively strong nonlinearities in early postnatal stages are substantially involved in the development of orientation tuning in visual cortex. [ABSTRACT FROM AUTHOR]

Neurons in the early visual cortex are generally highly sensitive to stimuli presented to the two eyes. However, the majority of studies on spatial and temporal aspects of neural responses were based on monocular measurements. To study neurons under more natural, i.e., binocular, conditions, we presented sinusoidal gratings of a variety of spatial frequencies (SF) dichoptically in rapid sequential flashes and analyzed the data using a binocular reverse correlation technique for neurons in cat area 17. The resulting set of data represents a frequency-domain binocular receptive field from which detailed selectivities, both monocular and binocular, could be obtained. Consistent with previous studies, the responses could generally be explained by linear summation of inputs from the two eyes. Suppressive responses were also observed and were delayed typically by 5–15 ms relative to excitatory responses. However, we have found more diverse nature of suppressive responses than those reported previously. The optimal suppressive frequency could be either higher or lower than that of the excitatory responses. The bandwidth of SF tuning of the suppressive responses was usually broader than that of the excitatory responses. Cells with lower optimal SFs for suppression tended to show high optimal SFs and sharp tuning curves. The dynamic shift of optimal SF from low to high SF was accompanied by suppression with earlier onset and higher peak SF or later onset and lower peak SF than excitation. These results suggest that the suppression plays an essential role in generating the temporal dynamics of SF selectivity. [ABSTRACT FROM AUTHOR]

A common technique used to study the response selectivity of neurons is to measure the relationship between sensory stimulation and action potential responses. Action potentials, however, are only indirectly related to the synaptic inputs that determine the underlying, subthreshold, response selectivity. We present a method to predict membrane potential, the measurable result of the convergence of synaptic inputs, based on spike rate alone and then test its utility by comparing predictions to actual membrane potential recordings from simple cells in primary visual cortex. Using a noise stimulus, we found that spike rate receptive fields were in precise correspondence with membrane potential receptive fields (R2= 0.74). On average, spike rate alone could predict 44% of membrane potential fluctuations to dynamic noise stimuli, demonstrating the utility of this method to extract estimates of subthreshold responses. We also found that the nonlinear relationship between membrane potential and spike rate could also be extracted from spike rate data alone by comparing predictions from the noise stimulus with the actual spike rate. Our analysis reveals that linear receptive field models extracted from noise stimuli accurately reflect the underlying membrane potential selectivity and thus represent a method to generate estimates of the underlying average membrane potential from spike rate data alone. [ABSTRACT FROM AUTHOR]

We used dynamic dense noise stimuli and local spectral reverse correlation methods to reveal the local sensitivities of neurons in visual area 2 (V2) of macaque monkeys to orientation and spatial frequency within their receptive fields. This minimized the potentially confounding assumptions that are inherent in stimulus selections. The majority of neurons exhibited a relatively high degree of homogeneity for the preferred orientations and spatial frequencies in the spatial matrix of facilitatory subfields. However, about 20% of all neurons showed maximum orientation differences between neighboring subfields that were greater than 25 deg. The neurons preferring horizontal or vertical orientations showed less inhomogeneity in space than the neurons preferring oblique orientations. Over 50% of all units also exhibited suppressive profiles, and those were more heterogeneous than facilitatory profiles. The preferred orientation and spatial frequency of suppressive profiles differed substantially from those of facilitatory profiles, and the neurons with suppressive subfields had greater orientation selectivity than those without suppressive subfields. The peak suppression occurred with longer delays than the peak facilitation. These results suggest that the receptive field profiles of the majority of V2 neurons reflect the orderly convergence of V1 inputs over space, but that a subset of V2 neurons exhibit more complex response profiles having both suppressive and facilitatory subfields. These V2 neurons with heterogeneous subfield profiles could play an important role in the initial processing of complex stimulus features. [ABSTRACT FROM AUTHOR]

The balance of inhibitory to excitatory (I/E) synaptic inputs is thought to control information processing and behavioral output of the central nervous system. We sought to test the effects of the decreased or increased I/E ratio on visual circuit function and visually guided behavior in Xenopus tadpoles. We selectively decreased inhibitory synaptic transmission in optic tectal neurons by knocking down the γ2 subunit of the GABAA receptors (GABAAR) using antisense morpholino oligonucleotides or by expressing a peptide corresponding to an intracellular loop of the γ2 subunit, called ICL, which interferes with anchoring GABAAR at synapses. Recordings of miniature inhibitory postsynaptic currents (mIPSCs) and miniature excitatory PSCs (mEPSCs) showed that these treatments decreased the frequency of mIPSCs compared with control tectal neurons without affecting mEPSC frequency, resulting in an ~50% decrease in the ratio of I/E synaptic input. ICL expression and γ2-subunit knockdown also decreased the ratio of optic nerve-evoked synaptic I/E responses. We recorded visually evoked responses from optic tectal neurons, in which the synaptic I/E ratio was decreased. Decreasing the synaptic I/E ratio in tectal neurons increased the variance of first spike latency in response to full-field visual stimulation, increased recurrent activity in the tectal circuit, enlarged spatial receptive fields, and lengthened the temporal integration window. We used the benzodiazepine, diazepam (DZ), to increase inhibitory synaptic activity. DZ increased optic nerve-evoked inhibitory transmission but did not affect evoked excitatory currents, resulting in an increase in the I/E ratio of ~30%. Increasing the I/E ratio with DZ decreased the variance of first spike latency, decreased spatial receptive field size, and lengthened temporal receptive fields. Sequential recordings of spikes and excitatory and inhibitory synaptic inputs to the same visual stimuli demonstrated that decreasing or increasing the I/E ratio disrupted input/output relations. We assessed the effect of an altered I/E ratio on a visually guided behavior that requires the optic tectum. Increasing and decreasing I/E in tectal neurons blocked the tectally mediated visual avoidance behavior. Because ICL expression, γ2-subunit knockdown, and DZ did not directly affect excitatory synaptic transmission, we interpret the results of our study as evidence that partially decreasing or increasing the ratio of I/E disrupts several measures of visual system information processing and visually guided behavior in an intact vertebrate. [ABSTRACT FROM AUTHOR]

The observation of figure-ground selectivity in neurons of the visual cortex shows that these neurons can be influenced by the image context far beyond the classical receptive field. To clarify the nature of the context integration mechanism, we studied the latencies of neural edge signals, comparing the emergence of context-dependent definition of border ownership with the onset of local edge definition (contrast polarity; stereoscopic depth order). Single-neuron activity was recorded in areas V1 and V2 of Macaca mulatta under behaviorally induced fixation. Whereas local edge definition emerged immediately (<13 ms) after the edge onset response, the contextdependent signal was delayed by about 30 ms. To see if the context influence was mediated by horizontal fibers within cortex, we measured the latencies of border ownership signals for two conditions in which the relevant context information was located at different distances from the receptive field and compared the latency difference with the difference predicted from horizontal signal propagation. The prediction was based on the increase in cortical distance, computed from the mapping of the test stimuli in the cortex, and the known conduction velocities of horizontal fibers. The measured latencies increased with cortical distance, but much less than predicted by the horizontal propagation hypothesis. Probability calculations showed that an explanation of the context influence by horizontal signal propagation alone is highly unlikely, whereas mechanisms involving back projections from other extrastriate areas are plausible. [ABSTRACT FROM AUTHOR]

Cortical neurons are organized in columns, distinguishable by their physiological properties and input-output organization. Columns are thought to be the fundamental information-processing modules of the cortex. The barrel cortex of rats and mice is an attractive model system for the study of cortical columns, because each column is defined by a layer 4 (L4) structure called a barrel, which can be clearly visualized. A great deal of information has been collected regarding the connectivity of neurons in barrel cortex, but the nature of the input to a given L4 barrel remains unclear. We measured this input by making comprehensive maps of whisker-evoked activity in L4 of rat barrel cortex using recordings of multiunit activity and current source density analysis of local field potential recordings of animals under light isoflurane anesthesia. We found that a large number of whiskers evoked a detectable response in each barrel (mean of 13 suprathreshold, 18 subthreshold) even after cortical activity was abolished by application of muscimol, a GABAA agonist. We confirmed these findings with intracellular recordings and single-unit extracellular recordings in vivo. This constitutes the first direct confirmation of the hypothesis that subcortical mechanisms mediate a substantial multiwhisker input to a given cortical barrel. [ABSTRACT FROM AUTHOR]

The ganglion cell output of the retina constitutes a bottleneck in sensory processing in that ganglion cells must encode multiple stimulus parameters in their responses. Here we investigate encoding strategies of On-Off directionally selective retinal ganglion cells (On-Off DS RGCs) in rabbits, a class of cells dedicated to representing motion. The exquisite axial discrimination of these cells to preferred vs. null direction motion is well documented: it is invariant with respect to speed, contrast, spatial configuration, spatial frequency, and motion extent. However, these cells have broad direction tuning curves and their responses also vary as a function of other parameters such as speed and contrast. In this study, we examined whether the variation in responses across multiple stimulus parameters is systematic, that is the same for all cells, and separable, such that the response to a stimulus is a product of the effects of each stimulus parameter alone. We extracellularly recorded single On-Off DS RGCs in a superfused eyecup preparation while stimulating them with moving bars. We found that spike count responses of these cells scaled as independent functions of direction, speed, and luminance. Moreover, the speed and luminance functions were common across the whole sample of cells. Based on these findings, we developed a model that accurately predicted responses of On-Off DS RGCs as products of separable functions of direction, speed, and luminance (r = 0.98; P < 0.0001). Such a multiplicatively separable encoding strategy may simplify the decoding of these cells' outputs by the higher visual centers. [ABSTRACT FROM AUTHOR]

Retinitis pigmentosa (RP) is a leading cause of degenerative vision loss, yet its progressive effects on visual signals transmitted from the retina to the brain are not well understood. The transgenic P23H rat is a valuable model of human autosomal dominant RP, exhibiting extensive similarities to the human disease pathology, time course, and electrophysiology. In this study, we examined the physiological effects of degeneration in retinal ganglion cells (RGCs) of P23H rats aged between P37 and P752, and compared them with data from wild-type control animals. The strength and the size of visual receptive fields of RGCs decreased rapidly with age in P23H retinas. Light responses mediated by rod photoreceptors declined earlier (∼ P300) than cone-mediated light responses (∼ P600). Responses of ON and OFF RGCs diminished at a similar rate. However, OFF cells exhibited hyperactivity during degeneration, whereas ON cells showed a decrease in firing rate. The application of synaptic blockers abolished about half of the elevated firing in OFF RGCs, indicating that the remodeled circuitry was not the only source of degeneration-induced hyperactivity. These results advance our understanding of the functional changes associated with retinal degeneration. [ABSTRACT FROM AUTHOR]

Neurons in the hand representation of primary somatosensory cortex (area 3b) are known to have discretely localized receptive fields; and these neurons form modules that can be visualized histologically as distinct digit and palm representations. Despite these indicators of the importance of local processing in area 3b, widespread interactions between stimuli presented to locations across the hand have been reported. We investigated the relationship of neuron firing rate with distance from the site of maximum activation in cortex by recording from a 100-electrode array with electrodes spaced 400 μm apart, implanted into the area 3b hand representation in anesthetized owl monkeys. For each stimulated location on the hand, the electrode site where neurons had the highest peak firing rate was defined as the peak activation site. The lesser firing rates of neurons at all other electrode sites in the grid were compared with the firing rates of neurons at the peak activation site. On average, peak firing rates of neurons decreased rapidly with distance away from the peak activation site. The effect of distance on the variance of firing rates was highly significant (P < 0.0001). However, individual neurons retained high firing rates for distances over 3 mm. The clear decline in firing rate with distance from the most activated location indicates that local processing is emphasized in area 3b, while the distance of neurons with reduced but maintained firing rates ≤3–4 mm from the site of best activation demonstrated widespread activation in primary somatosensory cortex. [ABSTRACT FROM AUTHOR]

People can find objects in a visual scene fast and effortlessly. It is thought that this may be accomplished by creating a map of the outside world that incorporates bottom-up sensory and top-down cognitive inputs—a priority map. Eye movements are made toward the location represented by the highest activity on the priority map. We hypothesized that the lateral intraparietal area (LIP) of posterior parietal cortex acts as such a map. To test this, we performed low current microstimulation on animals trained to perform a foraging task and asked whether we could bias the animals to make a saccade to a particular stimulus, by creating an artificial peak of activity at the location representing that stimulus on the map. We found that microstimulation slightly biased the animals to make saccades to visual stimuli at the stimulated location, without actively generating saccades. The magnitude of this effect was small, but it appeared to be similar for all visual stimuli. We interpret these results to mean that microstimulation slightly biased saccade goal selection to the object represented at the stimulated location in LIP. [ABSTRACT FROM AUTHOR]

In the primary visual cortex (V1), it has been shown that the neuronal response elicited by a grating patch in the receptive field (RF) center can be suppressed or facilitated by an annular grating presented in the RF surround area; the effect depends on the relative orientations of the two gratings. The effect is thought to play a role in figure-ground segregation. Here we have found that response modulation similar to that reported in cortical area V1 can also be found in all major classes of retinal ganglion cells (RGCs), including “concentric" cells. Orientation-specific response modulation of this kind cannot result from interactions of independent RF mechanisms; therefore more complex mechanism, which takes into account the relative orientations of the gratings in the RF center and surround, or sensing the borders between texture regions, has to be present in RFs of RGCs, even of the concentric type. This challenges the consensus notion that their responses to visual stimuli are governed entirely by a RF composed of separate mechanisms: center, antagonistic surround, and modulatory extraclassical surround. Our findings raise the question of whether initial stages of complex analysis of visual input, normally attributed to the visual cortex, can be achieved within the retina. [ABSTRACT FROM AUTHOR]