Activity In Early Visual Cortex Research Articles

Neural Correlates of the Formation and Retention of Cocaine-Induced Stimulus–Reward Associations

Cocaine can elicit drug-seeking behavior for drug-predicting stimuli, even after a single stimulus-cocaine pairing. Although orbitofrontal cortex is thought to be important during encoding and maintenance of stimulus-reward value, we still lack a comprehensive model of the neural circuitry underlying this cognitive process. We studied the conditioned effects of cocaine with monkey functional magnetic resonance imaging and classical conditioning by pairing a visual shape (conditioning stimulus [CS+]) with a noncontingent cocaine infusion; a control stimulus was never paired. We correlated the behavioral preference of the monkey for the CS+, as measured offline, with the activity induced by the CS+ relative to the control stimulus as function of time. We observed that during formation of stimulus-cocaine associations strong CS+-induced functional magnetic resonance imaging activations emerged in frontal cortex that correlated significantly with behavioral CS+ preference. Afterward, CS+ preference correlated only with activity in early visual cortex. Control experiments suggest that these findings cannot be explained by increased familiarity for the CS+. Our findings suggest a complex interaction between frontal and occipital cortex during cocaine conditioning. Frontal cortex is important for establishing novel representations of stimulus valence when cocaine is used as reinforcer, whereas early visual cortex is involved in retaining these cocaine-stimulus associations.

The Amplitude and Timing of the BOLD Signal Reflects the Relationship between Local Field Potential Power at Different Frequencies

There is growing evidence that several components of the mass neural activity contributing to the local field potential (LFP) can be partly separated by decomposing the LFP into nonoverlapping frequency bands. Although the blood oxygen level-dependent (BOLD) signal has been found to correlate preferentially with specific frequency bands of the LFP, it is still unclear whether the BOLD signal relates to the activity expressed by each LFP band independently of the others or if, instead, it also reflects specific relationships among different bands. We investigated these issues by recording, simultaneously and with high spatiotemporal resolution, BOLD signal and LFP during spontaneous activity in early visual cortices of anesthetized monkeys (Macaca mulatta). We used information theory to characterize the statistical dependency between BOLD and LFP. We found that the alpha (8-12 Hz), beta (18-30 Hz), and gamma (40-100 Hz) LFP bands were informative about the BOLD signal. In agreement with previous studies, gamma was the most informative band. Both increases and decreases in BOLD signal reliably followed increases and decreases in gamma power. However, both alpha and beta power signals carried information about BOLD that was largely complementary to that carried by gamma power. In particular, the relationship between alpha and gamma power was reflected in the amplitude of the BOLD signal, while the relationship between beta and gamma bands was reflected in the latency of BOLD with respect to significant changes in gamma power. These results lay the basis for identifying contributions of different neural pathways to cortical processing using fMRI.

Cross-Modal Multivariate Pattern Analysis

Multivariate pattern analysis (MVPA) is an increasingly popular method of analyzing functional magnetic resonance imaging (fMRI) data1-4. Typically, the method is used to identify a subject's perceptual experience from neural activity in certain regions of the brain. For instance, it has been employed to predict the orientation of visual gratings a subject perceives from activity in early visual cortices5 or, analogously, the content of speech from activity in early auditory cortices6.Here, we present an extension of the classical MVPA paradigm, according to which perceptual stimuli are not predicted within, but across sensory systems. Specifically, the method we describe addresses the question of whether stimuli that evoke memory associations in modalities other than the one through which they are presented induce content-specific activity patterns in the sensory cortices of those other modalities. For instance, seeing a muted video clip of a glass vase shattering on the ground automatically triggers in most observers an auditory image of the associated sound; is the experience of this image in the "mind's ear" correlated with a specific neural activity pattern in early auditory cortices? Furthermore, is this activity pattern distinct from the pattern that could be observed if the subject were, instead, watching a video clip of a howling dog?In two previous studies7,8, we were able to predict sound- and touch-implying video clips based on neural activity in early auditory and somatosensory cortices, respectively. Our results are in line with a neuroarchitectural framework proposed by Damasio9,10, according to which the experience of mental images that are based on memories - such as hearing the shattering sound of a vase in the "mind's ear" upon seeing the corresponding video clip - is supported by the re-construction of content-specific neural activity patterns in early sensory cortices.

Classification of Real and Imagined Sounds in Early Visual Cortex

Early visual cortex has been thought to be mainly involved in the detection of low-level visual features. Here we show that complex natural sounds can be decoded from early visual cortex activity, in the absence of visual stimulation and both when sounds are actually displayed and when they are merely imagined. Blindfolded subjects listened to three complex natural sounds (bird singing, people talking, traffic noise; Exp. 1) or received word cues (“forest”, “people”, “traffic”; Exp 2) to imagine the associated scene. fMRI BOLD activation patterns from retinotopically defined early visual areas were fed into a multivariate pattern classification algorithm (a linear support vector machine). Actual sounds were discriminated above chance in V2 and V3 and imagined sounds were decoded in V1. Also cross-classification, ie, training the classifier to real sounds and testing it to imagined sounds and vice versa, was successful. Two further experiments showed that an orthogonal working memory task does not interfer...

Decoding natural sounds in early visual cortex

Despite early visual cortex being one of the most intensely studied systems in the brain, the majority of processing variance is still unexplained. Here we show that information from hearing is contained in the activity patterns of early visual cortex, in a content-specific manner, in the absence of visual stimulation and despite an orthogonal working memory task. In Exp. 1, subject listened to 3 types of natural sounds while being blindfolded. We successfully decoded the sounds from retinotopically defined early visual cortex BOLD activity (mainly in V2 and V3) using multivariate pattern analysis (a linear support vector machine). To constrain mental imagery, subjects performed an orthogonal working memory task in Exp. 2, again in the absence of visual stimulation. Subjects memorised word lists of animals and objects during natural sound stimulation and performed a delayed match to sample task. Natural sounds were still successfully decoded from early visual cortex activity, despite mental imagery being restricted. These results show that contextual information from the auditory modality is contained in the activity patterns of early visual cortex, possibly to facilitate visual predictions.

Active Ignoring in Early Visual Cortex

Selective attention is critical for controlling the input to mental processes. Attentional mechanisms act not only to select relevant stimuli but also to exclude irrelevant stimuli. There is evidence that we can actively ignore irrelevant information. We measured neural activity relating to successfully ignoring distracters (using preview search) and found increases in both the precuneus and primary visual cortex during preparation to ignore distracters. We also found reductions in activity in fronto-parietal regions while previewing distracters and a reduction in activity in early visual cortex during search when a subset of items was successfully excluded from search, both associated with precuneus activity. These results are consistent with the proposal that actively excluding distractions has two components: an initial stage where distracters are encoded, and a subsequent stage where further processing of these items is inhibited. Our findings suggest that it is the precuneus that controls this process and can modulate activity in visual cortex as early as V1.

Top–Down Modulation of Human Early Visual Cortex after Stimulus Offset Supports Successful Postcued Report

Modulations of sensory processing in early visual areas are thought to play an important role in conscious perception. To date, most empirical studies focused on effects occurring before or during visual presentation. By contrast, several emerging theories postulate that sensory processing and conscious visual perception may also crucially depend on late top-down influences, potentially arising after a visual display. To provide a direct test of this, we performed an fMRI study using a postcued report procedure. The ability to report a target at a specific spatial location in a visual display can be enhanced behaviorally by symbolic auditory postcues presented shortly after that display. Here we showed that such auditory postcues can enhance target-specific signals in early human visual cortex (V1 and V2). For postcues presented 200 msec after stimulus termination, this target-specific enhancement in visual cortex was specifically associated with correct conscious report. The strength of this modulation predicted individual levels of performance in behavior. By contrast, although later postcues presented 1000 msec after stimulus termination had some impact on activity in early visual cortex, this modulation no longer related to conscious report. These results demonstrate that within a critical time window of a few hundred milliseconds after a visual stimulus has disappeared, successful conscious report of that stimulus still relates to the strength of top-down modulation in early visual cortex. We suggest that, within this critical time window, sensory representation of a visual stimulus is still under construction and so can still be flexibly influenced by top-down modulatory processes.

A neural basis for real-world visual search in human occipitotemporal cortex

Mammals are highly skilled in rapidly detecting objects in cluttered natural environments, a skill necessary for survival. What are the neural mechanisms mediating detection of objects in natural scenes? Here, we use human brain imaging to address the role of top-down preparatory processes in the detection of familiar object categories in real-world environments. Brain activity was measured while participants were preparing to detect highly variable depictions of people or cars in natural scenes that were new to the participants. The preparation to detect objects of the target category, in the absence of visual input, evoked activity patterns in visual cortex that resembled the response to actual exemplars of the target category. Importantly, the selectivity of multivoxel preparatory activity patterns in object-selective cortex (OSC) predicted target detection performance. By contrast, preparatory activity in early visual cortex (V1) was negatively related to search performance. Additional behavioral results suggested that the dissociation between OSC and V1 reflected the use of different search strategies, linking OSC preparatory activity to relatively abstract search preparation and V1 to more specific imagery-like preparation. Finally, whole-brain searchlight analyses revealed that, in addition to OSC, response patterns in medial prefrontal cortex distinguished the target categories based on the search cues alone, suggesting that this region may constitute a top-down source of preparatory activity observed in visual cortex. These results indicate that in naturalistic situations, when the precise visual characteristics of target objects are not known in advance, preparatory activity at higher levels of the visual hierarchy selectively mediates visual search.

The Functional Neuroanatomy of Object Agnosia: A Case Study

Cortical reorganization of visual and object representations following neural injury was examined using fMRI and behavioral investigations. We probed the visual responsivity of the ventral visual cortex of an agnosic patient who was impaired at object recognition following a lesion to the right lateral fusiform gyrus. In both hemispheres, retinotopic mapping revealed typical topographic organization and visual activation of early visual cortex. However, visual responses, object-related, and -selective responses were reduced in regions immediately surrounding the lesion in the right hemisphere, and also, surprisingly, in corresponding locations in the structurally intact left hemisphere. In contrast, hV4 of the right hemisphere showed expanded response properties. These findings indicate that the right lateral fusiform gyrus is critically involved in object recognition and that an impairment to this region has widespread consequences for remote parts of cortex. Finally, functional neural plasticity is possible even when a cortical lesion is sustained in adulthood.

Reduced hippocampal activity during encoding in cognitively normal adults carrying the APOE ɛ4 allele

Apolipoprotein (APOE) ɛ4-related differences in memory performance have been detected before age 65. The hippocampus and the surrounding medial temporal lobe (MTL) structures are the first site affected by Alzheimer's disease (AD) and the MTL is the seat of episodic memory, including visuo-spatial memory. While reports of APOE ɛ4-related differences in these brain structures are not consistent in either cross-sectional or longitudinal structural and functional magnetic resonance imaging (fMRI) studies, there is increasing evidence that brain activity at baseline (defined as activity during fixation or rest) may differ in APOE ɛ4 carriers compared to non-carriers. In this fMRI study, cognitively normal APOE ɛ4 carriers and non-carriers engaged in a perspective-dependent spatial learning task (Shelton & Gabrieli, 2002) previously shown to activate MTL structures in older participants (Borghesani et al., 2008). A low-level, visually engaging dot-control task was used for comparison, in addition to fixation. APOE ɛ4 carriers showed less activation than non-carriers in the hippocampus proper during encoding. Specifically, when spatial encoding was contrasted against the dot-control task, encoding-related activation was significantly lower in carriers than non-carriers. By contrast, no ɛ4-related differences in the hippocampus were found when spatial encoding was compared with fixation. Lower activation, however, was not global since encoding-related activation in early visual cortex (left lingual gyrus) was not different between APOE ɛ4 carriers and non-carriers. The present data document APOE ɛ4-related differences in the hippocampus proper during encoding and underscore the role of low-level control contrasts for complex encoding tasks. These results have implications for fMRI studies that investigate the default-mode network (DMN) in middle-aged to older APOE ɛ4 carriers to help evaluate AD risk in this otherwise cognitively normal population.

A chronometric exploration of high-resolution ‘sensitive TMS masking’ effects on subjective and objective measures of vision

Transcranial magnetic stimulation (TMS) can induce masking by interfering with ongoing neural activity in early visual cortex. Previous work has explored the chronometry of occipital involvement in vision by using single pulses of TMS with high temporal resolution. However, conventionally TMS intensities have been high and the only measure used to evaluate masking was objective in nature. Recent studies have begun to incorporate subjective measures of vision, alongside objective ones. The current study goes beyond previous work in two regards. First, we explored both objective vision (an orientation discrimination task) and subjective vision (a stimulus visibility rating on a four-point scale), across a wide range of time windows with high temporal resolution. Second, we used a very sensitive TMS-masking paradigm: stimulation was at relatively low TMS intensities, with a figure-8 coil, and the small stimulus was difficult to discriminate already at baseline level. We hypothesized that this should increase the effective temporal resolution of our paradigm. Perhaps for this reason, we are able to report a rather interesting masking curve. Within the classical-masking time window, previously reported to encompass broad SOAs anywhere between 60 and 120 ms, we report not one, but at least two dips in objective performance, with no masking in-between. The subjective measure of vision did not mirror this pattern. These preliminary data from our exploratory design suggest that, with sensitive TMS masking, we might be able to reveal visual processes in early visual cortex previously unreported.

Active ignoring in early visual cortex

Selective attention is critical for controlling the input to mental processes. Attentional mechanisms act not only to select relevant stimuli but also to exclude irrelevant stimuli. There is evidence that we can actively ignore irrelevant information. We measured neural activity relating to successfully ignoring distracters (using preview search) and found increases in both the precuneus and primary visual cortex during preparation to ignore distracters. We also found reductions in activity in fronto-parietal regions while previewing distracters and a reduction in activity in early visual cortex during search when a subset of items was successfully excluded from search, both associated with precuneus activity. These results are consistent with the proposal that actively excluding distractions has two components: an initial stage where distracters are encoded, and a subsequent stage where further processing of these items is inhibited. Our findings suggest that it is the precuneus that controls this process and can modulate activity in visual cortex as early as V1.

Transcranial magnetic stimulation-induced ‘visual echoes’ are generated in early visual cortex

Transcranial magnetic stimulation (TMS) of the early visual areas can trigger perception of a flash of light, a so-called phosphene. Here we show that a very brief presentation of a stimulus can modulate features of a subsequent TMS-induced phosphene, to a level that participants mistake phosphenes for real stimuli, inducing ‘visual echoes’ of a previously seen stimulus. These ‘echoes’ are modulated by visual context at the moment of magnetic stimulation, showing that they are generated in early visual areas, and that the brain processes these ‘echoes’ as if they are factually presented stimuli. This shows that TMS can re-activate weak visual representations in early visual areas. Based on the pattern of contextual modulation of visual echoes, we theorize that perception of these echoes is not a passive reactivation of residual activity in early visual cortex, but an active interpretation of the combined activity of TMS-induced neural noise and cortical state.

Baseline fMRI pattern activity in early visual cortex predicts the initial dominant percept in subsequent binocular rivalry

Baseline fMRI pattern activity in early visual cortex predicts the initial dominant percept in subsequent binocular rivalry

V1 responses to different types of luminance histogram contrast

It has been suggested that there exist neural mechanisms which are sensitive to luminance histogram skew, and that these mechanisms play a role in estimating surface properties (Motoyoshi, Nishida, Sharan & Adelson, 2007), or texture discrimination (Chubb, Landy, Econopouly, 2004). However, to the best of our knowledge, no systematic measurements of neural activity in early visual cortex in response to stimuli with skewed luminance histogram have been made to corroborate the existence of the postulated mechanisms. Using fMRI, we measured BOLD signal in early visual cortex in response to noise images with either positive, negative or zero skewness of the luminance histogram. Stimuli were circular (radius = 6.35 deg visual angle) random noise images whose histogram mean (M=128) and standard deviation (SD=43) were equated. Histogram contrast was manipulated by adjusting the third moment (skewness) of the luminance histogram to take on one of three values: positive (+1.4), negative (−1.4), or zero (0). Images were presented at a rate of 10Hz, in blocks of 24s separated by 24s presentation of a static image with uniform mean luminance (M=128, SD=0). The order of presentation during each scan (TR=2s) was: blank, positive skew, blank, negative skew, blank, zero skew. This sequence was repeated twice during each scan; there were a total of six scans per session. Observers fixated at the center of each stimulus, and performed a demanding fixation task during each scan. The BOLD signal was analyzed within an independently determined ROI corresponding to an annulus located within the stimulus area (inner radius= 2 deg, outer radius 4 deg). On average V1 responses to noise images with positively and negatively skewed luminance histogram were significantly larger than those to images with zero skew. Furthermore, the V1 BOLD signal in response to images with negative contrast was larger than that to those with positive luminance histogram contrast.

Dissociation of Neural Mechanisms Underlying Orientation Processing in Humans

Orientation selectivity is a fundamental, emergent property of neurons in early visual cortex, and the discovery of that property has dramatically shaped how we conceptualize visual processing. However, much remains unknown about the neural substrates of this basic building block of perception, and what is known primarily stems from animal physiology studies. To probe the neural concomitants of orientation processing in humans, we employed repetitive transcranial magnetic stimulation (rTMS), which can significantly attenuate neuronal spiking activity, hemodynamic responses, and local field potentials within a focused cortical region. Using rTMS to suppress neural responses evoked by stimuli falling within a local region of the visual field, we were able to dissociate two distinct components of the neural circuitry underlying orientation processing: selectivity and contextual effects. Orientation selectivity gauged by masking was unchanged by rTMS, whereas an otherwise robust orientation repulsion illusion was weakened after rTMS. This dissociation implies that orientation processing in humans relies on distinct mechanisms, only one of which was impacted by rTMS. These results are consistent with models positing that orientation selectivity is governed by patterns of convergence of thalamic afferents onto cortical neurons, with intracortical activity then shaping population responses amongst those cortical neurons.

Specificity of auditory-guided visual perceptual learning suggests crossmodal plasticity in early visual cortex

Sounds modulate visual perception. Blind humans show altered brain activity in early visual cortex. However, it is still unclear whether crossmodal activity in visual cortex results from unspecific top-down feedback, a lack of visual input, or genuinely reflects crossmodal interactions at early sensory levels. We examined how sounds affect visual perceptual learning in sighted adults. Visual motion discrimination was tested prior to and following eight sessions in which observers were exposed to irrelevant moving dots while detecting sounds. After training, visual discrimination improved more strongly for motion directions that were paired with a relevant sound during training than for other directions. Crossmodal learning was limited to visual field locations that overlapped with the sound source and was little affected by attention. The specificity and automatic nature of these learning effects suggest that sounds automatically guide visual plasticity at a relatively early level of processing.

Affective Learning Enhances Visual Detection and Responses in Primary Visual Cortex

The affective significance of a visual item is thought to lead to enhanced visual processing. However, the precise link between enhanced visual perception of emotion-laden items and increased visual responses remains poorly understood. To investigate this link, we acquired functional magnetic resonance imaging (fMRI) data while participants performed a challenging visual detection task. Grating stimuli were physically identical and differed only as a function of their previous exposure history; CS+ stimuli were initially paired with shock, whereas CS- stimuli were not. Behaviorally, subjects were both faster and more accurate during CS+ relative to CS- target detection. These behavioral results were paralleled by increases in fMRI responses across early, retinotopically organized visual cortex, which was mapped in a separate fMRI session. Logistic regression analyses revealed that trial-by-trial fluctuations in fMRI responses were closely linked to trial type, such that fMRI signal strength reliably predicted the probability of a hit trial across retinotopically organized visual cortex, including area V1. For instance, during the CS+ condition, a 0.5% signal change increased the probability of a hit from chance to 67.3-73.5% in V1-V4 (the highest increase was observed in area V1). Furthermore, across participants, differential fMRI responses to hits versus correct rejects were correlated with behavioral performance. Our findings provide a close link between increased activation in early visual cortex and improved behavioral performance as a function of the affective significance of an item.

Perceptual Learning and Dynamic Changes in Primary Visual Cortex

Perceptual learning is the improved performance that follows practice in a perceptual task. In this issue of Neuron, Yotsumoto et al. use fMRI to show that stimuli presented at the location used in training initially evoke greater activation in primary visual cortex than stimuli presented elsewhere, but this difference disappears once learning asymptotes.

Spatial Attention Does Not Strongly Modulate Neuronal Responses in Early Human Visual Cortex

Attention can dramatically enhance behavioral performance based on a visual stimulus, but the degree to which attention modulates activity in early visual cortex is unclear. Whereas single-unit studies of spatial attention in monkeys have repeatedly revealed relatively modest attentional modulations in V1, human functional magnetic resonance imaging studies demonstrate a large attentional enhancement of the blood oxygen level-dependent (BOLD) signal in V1. To explore this discrepancy, we used intracranial electrodes to directly measure the effect of spatial attention on the responses of neurons near the human occipital pole. We found that spatial attention does not robustly modulate stimulus-driven local field potentials in early human visual cortex, but instead produces modest modulations that are consistent with those seen in monkey neurophysiology experiments. This finding suggests that the neuronal activity that underlies visual attention in humans is similar to that found in other primates and that behavioral state may alter the linear relationship between neuronal activity and BOLD.