Interactions In Visual Cortex Research Articles

Invited Session I: 30 years of normalization in the visual system: Contrast normalization accounts for binocular interactions in healthy vision and amblyopia.

We investigated the interaction between dichoptic stimuli using steady-state visual evoked potentials and frequency-domain analysis. The stimulus in each eye flickered with a unique temporal frequency, which allowed us to "frequency-tag" the responses to each eye's input (self terms) as well as the responses to the combination of inputs from the two eyes (intermodulation terms). We measured two forms of binocular interaction: one associated with the suppressive effect of one eye's stimulus on the other, and the other associated with a direct measure of interocular interaction between the two eyes' inputs. Fits of a contrast gain control model to the data demonstrated that a common gain control mechanism is consistent with both forms of binocular interaction. We then used the contrast normalization framework to investigate the disruptions to binocular interaction in amblyopia. Although anisometropic amblyopes showed a similar pattern of responses to normal-vision observers, strabismic amblyopes exhibited both reduced responses to the amblyopic eye stimulus in the presence of a mask in the other eye, as well as substantially reduced intermodulation responses indicating reduced interocular interactions in visual cortex. A contrast normalization model that simultaneously fit self- and IM-term responses showed that the excitatory contribution to binocular interaction is significantly reduced in strabismic amblyopia.

Neural correlates of visual acuity for fine text

Human sensitivity to visual input often scales with the magnitude of evoked responses in the brain. Here, we demonstrate an exception. We record electroencephalography (EEG) while people attempt to resolve fine print – similar to people attempting to read eye charts (the world’s most popular means of testing vision). We find that the ability to resolve fine print is associated with smaller evoked responses recorded by large clusters of occipital-parietal sensors ∼150 ms after people see words. Moreover, we find that a better ability to resolve fine print is associated with enhanced alpha-band oscillatory brain activity immediately prior to word presentations. These investigations were inspired by psychophysical data, which suggested the ability to resolve fine print can be enhanced by pre-adaptation to flicker, which should encourage a reduced neural response to inputs. We included this manipulation in this study, and our results are broadly consistent with this conjecture. As alpha-band activity has been linked to inhibitory interactions in visual cortex, we regard our data as evidence that smaller neural responses to fine print can be promoted by inhibitory processes that target unhelpful blur-related signals, which thereby sharpen subsequent visual experiences.

Excitatory Contribution to Binocular Interactions in Human Visual Cortex Is Reduced in Strabismic Amblyopia.

Binocular summation in strabismic amblyopia is typically reported as being absent or greatly reduced in behavioral studies and is thought to be because of a preferential loss of excitatory interactions between the eyes. Here, we studied how excitatory and suppressive interactions contribute to binocular contrast interactions along the visual cortical hierarchy of humans with strabismic and anisometropic amblyopia in both sexes, using source-imaged steady-state visual evoked potentials (SSVEP) over a wide range of relative contrast between the two eyes. Dichoptic parallel grating stimuli modulated at unique temporal frequencies in each eye allowed us to quantify spectral response components associated with monocular inputs (self-terms) and the response components because of interaction of the inputs of the two eyes [intermodulation (IM) terms]. Although anisometropic amblyopes revealed a similar pattern of responses to normal-vision observers, strabismic amblyopes exhibited substantially reduced IM responses across cortical regions of interest (V1, V3a, hV4, hMT+ and lateral occipital cortex), indicating reduced interocular interactions in visual cortex. A contrast gain control model that simultaneously fits self- and IM-term responses within each cortical area revealed different patterns of binocular interactions between individuals with normal and disrupted binocularity. Our model fits show that in strabismic amblyopia, the excitatory contribution to binocular interactions is significantly reduced in both V1 and extra-striate cortex, whereas suppressive contributions remain intact. Our results provide robust electrophysiological evidence supporting the view that disruption of binocular interactions in strabismus or amblyopia is because of preferential loss of excitatory interactions between the eyes.SIGNIFICANCE STATEMENT We studied how excitatory and suppressive interactions contribute to binocular contrast interactions along the visual cortical hierarchy of humans with normal and amblyopic vision, using source-imaged SSVEP and frequency-domain analysis of dichoptic stimuli over a wide range of relative contrast between the two eyes. A dichoptic contrast gain control model was used to characterize these interactions in amblyopia and provided a quantitative comparison to normal vision. Our model fits revealed different patterns of binocular interactions between normal and amblyopic vision. Strabismic amblyopia significantly reduced excitatory contributions to binocular interactions, whereas suppressive contributions remained intact. Our results provide robust evidence supporting the view that the preferential loss of excitatory interactions disrupts binocular interactions in strabismic amblyopia.

Desflurane Anesthesia Alters Cortical Layer-specific Hierarchical Interactions in Rat Cerebral Cortex.

Neurocognitive investigations suggest that conscious sensory perception depends on recurrent neuronal interactions among sensory, parietal, and frontal cortical regions, which are suppressed by general anesthetics. The purpose of this work was to investigate if local interactions in sensory cortex are also altered by anesthetics. The authors hypothesized that desflurane would reduce recurrent neuronal interactions in cortical layer-specific manner consistent with the anatomical disposition of feedforward and feedback pathways. Single-unit neuronal activity was measured in freely moving adult male rats (268 units; 10 animals) using microelectrode arrays chronically implanted in primary and secondary visual cortex. Layer-specific directional interactions were estimated by mutual information and transfer entropy of multineuron spike patterns within and between cortical layers three and five. The effect of incrementally increasing and decreasing steady-state concentrations of desflurane (0 to 8% to 0%) was tested for statistically significant quadratic trend across the successive anesthetic states. Desflurane produced robust, state-dependent reduction (P = 0.001) of neuronal interactions between primary and secondary visual areas and between layers three and five, as indicated by mutual information (37 and 41% decrease at 8% desflurane from wakeful baseline at [mean ± SD] 0.52 ± 0.51 and 0.53 ± 0.51 a.u., respectively) and transfer entropy (77 and 78% decrease at 8% desflurane from wakeful baseline at 1.86 ± 1.56 a.u. and 1.87 ± 1.67 a.u., respectively). In addition, a preferential suppression of feedback between secondary and primary visual cortex was suggested by the reduction of directional index of transfer entropy overall (P = 0.001; 89% decrease at 8% desflurane from 0.11 ± 0.18 a.u. at baseline) and specifically, in layer five (P = 0.001; 108% decrease at 8% desflurane from 0.12 ± 0.19 a.u. at baseline). Desflurane anesthesia reduces neuronal interactions in visual cortex with a preferential effect on feedback. The findings suggest that neuronal disconnection occurs locally, among hierarchical sensory regions, which may contribute to global functional disconnection underlying anesthetic-induced unconsciousness.

Frequency and phase-specific direct interaction in visual cortex between visually evoked and tACS induced neuronal signals

Frequency and phase-specific direct interaction in visual cortex between visually evoked and tACS induced neuronal signals

Asymmetric Dichoptic Masking in Visual Cortex of Amblyopic Macaque Monkeys.

In amblyopia, abnormal visual experience leads to an extreme form of eye dominance, in which vision through the nondominant eye is degraded. A key aspect of this disorder is perceptual suppression: the image seen by the stronger eye often dominates during binocular viewing, blocking the image of the weaker eye from reaching awareness. Interocular suppression is the focus of ongoing work aimed at understanding and treating amblyopia, yet its physiological basis remains unknown. We measured binocular interactions in visual cortex of anesthetized amblyopic monkeys (female Macaca nemestrina), using 96-channel "Utah" arrays to record from populations of neurons in V1 and V2. In an experiment reported recently (Hallum et al., 2017), we found that reduced excitatory input from the amblyopic eye (AE) revealed a form of balanced binocular suppression that is unaltered in amblyopia. Here, we report on the modulation of the gain of excitatory signals from the AE by signals from its dominant fellow eye (FE). Using a dichoptic masking technique, we found that AE responses to grating stimuli were attenuated by the presentation of a noise mask to the FE, as in a normal control animal. Responses to FE stimuli, by contrast, could not be masked from the AE. We conclude that a weakened ability of the amblyopic eye to modulate cortical response gain creates an imbalance of suppression that favors the dominant eye.SIGNIFICANCE STATEMENT In amblyopia, vision in one eye is impaired as a result of abnormal early visual experience. Behavioral observations in humans with amblyopia suggest that much of their visual loss is due to active suppression of their amblyopic eye. Here we describe experiments in which we studied binocular interactions in macaques with experimentally induced amblyopia. In normal monkeys, the gain of neuronal response to stimulation of one eye is modulated by contrast in the other eye, but in monkeys with amblyopia the balance of gain modulation is altered so that the weaker, amblyopic eye has little effect while the stronger fellow eye has a strong effect. This asymmetric suppression may be a key component of the perceptual losses in amblyopia.

Phase-Dependent Interactions in Visual Cortex to Combinations of First- and Second-Order Stimuli.

Many visual cortex neurons exhibit orientation-selective responses to boundaries defined by differences either in luminance or in texture contrast. Previous studies have examined responses to either type of boundary in isolation, but here we measured systematically responses of cortical neurons to the spatial relationship between superimposed periodic luminance-modulated (LM) and contrast-modulated (CM) stimuli with contrasts adjusted to give equated responses. We demonstrate that neuronal responses to these compound stimuli are highly dependent on the relative phase between the LM and CM components. Diversity in the degree of such phase dependence could help to disambiguate different kinds of boundaries in natural scenes, for example, those arising from surface reflectance changes or from illumination gradients such as shading or shadows.

Abnormal inhibition-excitation imbalance in migraine.

People with migraine show increased surround suppression of perceived contrast, a perceptual analogue of centre-surround antagonistic interactions in visual cortex. A proposed mechanism is that cortical 'hyperexcitability' or 'hyperresponsivity', a prominent theory in the migraine literature, drives abnormal excitatory-inhibitory balance to give increased local inhibition. The purpose of this cross-sectional study was to determine whether cortical hyperresponsivity and excitatory-inhibitory imbalance manifests in the visual cortical response of migraine sufferers. Interictal steady-state visual evoked potentials (VEPs) in response to 0 to 97% contrast were recorded in 30 migraine participants (15 without aura, 15 with aura) and 21 non-headache controls. Monotonicity indices were calculated to determine response saturation or supersaturation. Contrast gain was modelled with a modified saturating hyperbolic function to allow for variation in excitation and inhibition. A greater proportion of migraine participants (43%) than controls (14%) exhibited significant VEP supersaturation at high contrast, based on monotonicity index (chi-square, p = 0.028). Supersaturation was also evident by the trend for greater suppressive exponent values in migraine compared to control individuals (Mann-Whitney rank sum, p = 0.075). Supersaturation in migraine is consistent with excess excitation (hyperresponsivity) driving increased network inhibition and provides support for excitatory-inhibitory imbalance as a pathophysiological disturbance in migraine.

Object grouping based on real-world regularities facilitates perception by reducing competitive interactions in visual cortex.

In virtually every real-life situation humans are confronted with complex and cluttered visual environments that contain a multitude of objects. Because of the limited capacity of the visual system, objects compete for neural representation and cognitive processing resources. Previous work has shown that such attentional competition is partly object based, such that competition among elements is reduced when these elements perceptually group into an object based on low-level cues. Here, using functional MRI (fMRI) and behavioral measures, we show that the attentional benefit of grouping extends to higher-level grouping based on the relative position of objects as experienced in the real world. An fMRI study designed to measure competitive interactions among objects in human visual cortex revealed reduced neural competition between objects when these were presented in commonly experienced configurations, such as a lamp above a table, relative to the same objects presented in other configurations. In behavioral visual search studies, we then related this reduced neural competition to improved target detection when distracter objects were shown in regular configurations. Control studies showed that low-level grouping could not account for these results. We interpret these findings as reflecting the grouping of objects based on higher-level spatial-relational knowledge acquired through a lifetime of seeing objects in specific configurations. This interobject grouping effectively reduces the number of objects that compete for representation and thereby contributes to the efficiency of real-world perception.

Mixing of Chromatic and Luminance Retinal Signals in Primate Area V1

Vision emerges from activation of chromatic and achromatic retinal channels whose interaction in visual cortex is still poorly understood. To investigate this interaction, we recorded neuronal activity from retinal ganglion cells and V1 cortical cells in macaques and measured their visual responses to grating stimuli that had either luminance contrast (luminance grating), chromatic contrast (chromatic grating), or a combination of the two (compound grating). As with parvocellular or koniocellular retinal ganglion cells, some V1 cells responded mostly to the chromatic contrast of the compound grating. As with magnocellular retinal ganglion cells, other V1 cells responded mostly to the luminance contrast and generated a frequency-doubled response to equiluminant chromatic gratings. Unlike magnocellular and parvocellular retinal ganglion cells, V1 cells formed a unimodal distribution for luminance/color preference with a 2- to 4-fold bias toward luminance. V1 cells associated with positive local field potentials in deep layers showed the strongest combined responses to color and luminance and, as a population, V1 cells encoded a diverse combination of luminance/color edges that matched edge distributions of natural scenes. Taken together, these results suggest that the primary visual cortex combines magnocellular and parvocellular retinal inputs to increase cortical receptive field diversity and to optimize visual processing of our natural environment.

Network model of top-down influences on local gain and contextual interactions in visual cortex

The visual system uses continuity as a cue for grouping oriented line segments that define object boundaries in complex visual scenes. Many studies support the idea that long-range intrinsic horizontal connections in early visual cortex contribute to this grouping. Top-down influences in primary visual cortex (V1) play an important role in the processes of contour integration and perceptual saliency, with contour-related responses being task dependent. This suggests an interaction between recurrent inputs to V1 and intrinsic connections within V1 that enables V1 neurons to respond differently under different conditions. We created a network model that simulates parametrically the control of local gain by hypothetical top-down modification of local recurrence. These local gain changes, as a consequence of network dynamics in our model, enable modulation of contextual interactions in a task-dependent manner. Our model displays contour-related facilitation of neuronal responses and differential foreground vs. background responses over the neuronal ensemble, accounting for the perceptual pop-out of salient contours. It quantitatively reproduces the results of single-unit recording experiments in V1, highlighting salient contours and replicating the time course of contextual influences. We show by means of phase-plane analysis that the model operates stably even in the presence of large inputs. Our model shows how a simple form of top-down modulation of the effective connectivity of intrinsic cortical connections among biophysically realistic neurons can account for some of the response changes seen in perceptual learning and task switching.

Top-Down Modulation of Lateral Interactions in Visual Cortex

The primary visual cortex (V1) changes its computation according to the perceptual task being performed. We propose that this cognitive modulation results from gating of V1 intrinsic connections. To test this idea, using behavioral paradigms that engage top-down modulation of V1 contextual interactions, we recorded from chronically implanted electrode arrays in macaques. We observed task-dependent changes in interactions between V1 sites measured both by correlation between spike trains and by coherence between local field potentials (LFP-LFP coherence). The direction of the changes in aggregate activity, as measured by LFPs, depended on perceptual strategy: perceptual grouping increased LFP coherence between sites crucial for the task, whereas perceptual segregation lowered the LFP coherence. Using spiking activity as a measure, we found that the behaviorally driven changes in correlation structure between neurons dramatically increased the stimulus-related information that they convey; this additional increase in encoded information at the level of neuronal ensembles equals that obtained from task-driven reconfigurations of neural tuning curves. The improvements in information encoding were strongest for stimuli with greatest discrimination difficulty.

The BCM theory of synapse modification at 30: interaction of theory with experiment

Thirty years have passed since the publication of Elie Bienenstock, Leon Cooper and Paul Munro's 'Theory for the development of neuron selectivity: orientation specificity and binocular interaction in visual cortex', known as the BCM theory of synaptic plasticity. This theory has guided experimentalists to discover some fundamental properties of synaptic plasticity and has provided a mathematical structure that bridges molecular mechanisms and systems-level consequences of learning and memory storage.

Generalized spin models for coupled cortical feature maps obtained by coarse graining correlation based synaptic learning rules

We derive generalized spin models for the development of feedforward cortical architecture from a Hebbian synaptic learning rule in a two layer neural network with nonlinear weight constraints. Our model takes into account the effects of lateral interactions in visual cortex combining local excitation and long range effective inhibition. Our approach allows the principled derivation of developmental rules for low-dimensional feature maps, starting from high-dimensional synaptic learning rules. We incorporate the effects of smooth nonlinear constraints on net synaptic weight projected from units in the thalamic layer (the fan-out) and on the net synaptic weight received by units in the cortical layer (the fan-in). These constraints naturally couple together multiple feature maps such as orientation preference and retinotopic organization. We give a detailed illustration of the method applied to the development of the orientation preference map as a special case, in addition to deriving a model for joint pattern formation in cortical maps of orientation preference, retinotopic location, and receptive field width. We show that the combination of Hebbian learning and center-surround cortical interaction naturally leads to an orientation map development model that is closely related to the XY magnetic lattice model from statistical physics. The results presented here provide justification for phenomenological models studied in Cowan and Friedman (Advances in neural information processing systems 3, 1991), Thomas and Cowan (Phys Rev Lett 92(18):e188101, 2004) and provide a developmental model realizing the synaptic weight constraints previously assumed in Thomas and Cowan (Math Med Biol 23(2):119-138, 2006).

Attention does more than modulate suppressive interactions: attending to multiple items

Directing attention to a visual item enhances its representations, making it more likely to guide behavior (Corbetta et al. 1991). Attention is thought to produce this enhancement by biasing suppressive interactions among multiple items in visual cortex in favor of the attended item (e.g., Desimone and Duncan 1995; Reynolds and Heeger 2009). We ask whether target enhancement and modulation of suppressive interactions are in fact inextricably linked or whether they can be decoupled. In particular, we ask whether simultaneously directing attention to multiple items may be one means of dissociating the influence of attention-related enhancement from the effects of inter-item suppression. When multiple items are attended, suppressive interactions in visual cortex limit the effectiveness with which attention may act on their representations, presumably because "biasing" the interactions in favor of a single item is no longer possible (Scalf and Beck 2010). In this experiment, we directly investigate whether applying attention to multiple competing stimulus items has any influence on either their evoked signal or their suppressive interactions. Both BOLD signal evoked by the items in V4 and behavioral responses to those items were significantly compromised by simultaneous presentation relative to simultaneous presentation, indicating that when the items appeared at the same time, they interacted in a mutually suppressive manner that compromised their ability to guide behavior. Attention significantly enhanced signal in V4. The attentional status of the items, however, had no influence on the suppressive effects of simultaneous presentation. To our knowledge, these data are the first to explicitly decouple the effects of top-down attention from those of inter-item suppression.

Recurrent Processing in V1/V2 Contributes to Categorization of Natural Scenes

Humans are able to categorize complex natural scenes very rapidly and effortlessly, which has led to an assumption that such ultra-rapid categorization is driven by feedforward activation of ventral brain areas. However, recent accounts of visual perception stress the role of recurrent interactions that start rapidly after the activation of V1. To study whether or not recurrent processes play a causal role in categorization, we applied fMRI-guided transcranial magnetic stimulation on early visual cortex (V1/V2) and lateral occipital cortex (LO) while the participants categorized natural images as containing animals or not. The results showed that V1/V2 contributed to categorization speed and to subjective perception during a long activity period before and after the contribution of LO had started. This pattern of results suggests that recurrent interactions in visual cortex between areas along the ventral stream and striate cortex play a causal role in categorization and perception of natural scenes.

Interareal directed interactions and their modulation by selective attention assessed with high density electrocorticography in monkey

Event Abstract Back to Event Interareal directed interactions and their modulation by selective attention assessed with high density electrocorticography in monkey Andre Bastos1, 2*, Pascal Fries2, 3, Robert Oostenveld2, Jan-Mathijs Schoffelen2 and Conrado A. Bosman2 1 University of California at Davis, United States 2 Donders Institute, Radboud University Nijmegen, Netherlands 3 Ernst Strüngmann Institute in Cooperation with Max Planck Society, Germany Behaviorally relevant stimuli are processed in visual cortex at the expense of irrelevant ones. The selection of relevant stimuli and the allocation of selective attention require parietal and frontal regions. Yet, it is not fully understood how these areas convey their selection to visual areas. The selection needs to flexibly implement appropriate patterns of interactions in visual cortex. Such flexible interactions might be subserved by flexible patterns of neuronal synchronization. To test this possibility, we trained two monkeys to perform a selective attention task. During fixation, two gratings were presented in the two hemifields. On a given trial, one of them was the behaviorally relevant target. We recorded local field potentials with a subdural electrocorticography (ECoG) grid that contained 252 electrodes, spaced at 2-3 mm, covering large expanses of the left hemisphere from V1 through V4, parietal and central cortex up to frontal eye field (FEF). Coherence and Granger Causality analyses revealed networks with spatially and spectrally distinct synchronization patterns. Areas V1 and V4 were synchronized in the gamma-frequency band (50-90 Hz) and this interaction was almost unidirectional from V1 to V4. Attention to a contralateral stimulus enhanced this feedforward gamma-band influence. Parietal cortex was synchronized with both V4 and with FEF in the beta frequency band. Between parietal cortex and V4, this interaction was predominantly from parietal to V4. For many V4-parietal electrode pairs, selective attention modulated and often enhanced the interaction. We propose that these two spatially and spectrally segregated networks interact to coordinate executive signals from the parietal and prefrontal cortices with perceptual processing in visual cortex Keywords: electrocorticography, neuronal oscillations Conference: XI International Conference on Cognitive Neuroscience (ICON XI), Palma, Mallorca, Spain, 25 Sep - 29 Sep, 2011. Presentation Type: Symposium: Oral Presentation Topic: Symposium 3: The role of neuronal oscillations in computation and communication in multi-scale brain networks Citation: Bastos A, Fries P, Oostenveld R, Schoffelen J and Bosman CA (2011). Interareal directed interactions and their modulation by selective attention assessed with high density electrocorticography in monkey. Conference Abstract: XI International Conference on Cognitive Neuroscience (ICON XI). doi: 10.3389/conf.fnhum.2011.207.00023 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 09 Nov 2011; Published Online: 15 Nov 2011. * Correspondence: Dr. Andre Bastos, University of California at Davis, Davis, United States, andrembastos@gmail.com Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Andre Bastos Pascal Fries Robert Oostenveld Jan-Mathijs Schoffelen Conrado A Bosman Google Andre Bastos Pascal Fries Robert Oostenveld Jan-Mathijs Schoffelen Conrado A Bosman Google Scholar Andre Bastos Pascal Fries Robert Oostenveld Jan-Mathijs Schoffelen Conrado A Bosman PubMed Andre Bastos Pascal Fries Robert Oostenveld Jan-Mathijs Schoffelen Conrado A Bosman Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Functional Magnetic Resonance Imaging of Binocular Interactions in Visual Cortex in Strabismus

To observe changes in primary visual cortical activation associated with perceptual suppression in individuals with strabismus, using functional magnetic resonance imaging (fMRI). In Phase 1, pilot data were collected from 1 control and 8 strabismic participants, including 5 with amblyopia. In Phase 2, results were collected from 7 participants with strabismus (2 recalled and 5 new), including 2 with amblyopia. fMRI compared primary visual cortex activation in two conditions: visual stimuli presented to both eyes, to evoke perceptual suppression of one eye in individuals with strabismus, and visual stimulation presented only to one eye. Visual cortical activity modulations positively correlated with perceptual suppression were seen in 3 of 5 non-amblyopic Phase 2 study participants, but not in amblyopic subjects. Cortical activity modulations correlated with modulation of strabismic suppression are detectable in non-amblyopic individuals using fMRI, suggesting a neural basis for strabismic suppression in primary visual cortex.

Perceptual load-induced selection as a consequence of spatial interactions in visual cortex

Perceptual load-induced selection as a consequence of spatial interactions in visual cortex

Competition in Visual Cortex Impedes Attention to Multiple Items

Traditional explanations of our limited attentional capacity focus on our ability to direct attention to multiple items. We ask whether this difficulty in simultaneously attending to multiple items stems from an inability to effectively represent multiple attended items. Although attending to one of a set of neighboring stimuli can isolate it from competitive interactions in visual cortex, no such isolation should occur if multiple competing items are attended. Indeed, we find that attention is ineffective at enhancing blood oxygen level-dependent signal in visual cortical area V4 when it is directed to three stimuli simultaneously, but only when those three stimuli compete in visual cortex. This suggests that competition may prevent attention from acting as effectively on representations of multiple items as it does on representations of a single item. In contrast to traditional explanations that posit limits in the sources of attentional control, we show that mechanisms at the sites of stimulus representation may also impose limits on our ability to attend to multiple items simultaneously.