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ABSTRACT: To optimize perception, neurons in the visual system adapt to the current environment. What determines the durability of this plasticity? Longer exposures to an environment produce longer-lasting effects, which could be due to either (i) a single mechanism controlling adaptation that gains strength over time, or (ii) long-term mechanisms that become active after long-term exposure. Using recently developed technology, we tested adaptation durations an order of magnitude greater that those tested previously, and used a "deadaptation" procedure to reveal effects of a unique long-term mechanism in the longest adaptation periods. After 4 h of contrast adaptation, human observers were exposed to natural images for 15 min, which completely cancelled perceptual aftereffects of adaptation. Strikingly, during continued testing this deadaptation faded, and the original adaptation effects reappeared. This pattern strongly suggests that adaptation was maintained in a distinct long-term mechanism, whereas deadaptation affected a short-term mechanism.
Proceedings of the National Academy of Sciences 03/2012; 109(15):5898-903. · 9.68 Impact Factor
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ABSTRACT: Eccentric viewing in macular disease has been described for half a century. However, a clear definition of eccentric viewing and preferred retinal locus (PRL) does not exist. Here, we determine how the PRL in macular disease is defined by researchers active in this field and, based on the responses received, propose a standardized definition of the preferred retinal locus.
A literature review of articles describing the PRL or eccentric viewing was performed. The first and senior authors of all identified publications were contacted and were asked to define the preferred retinal locus. Themes of responses were identified using inductive qualitative research techniques.
Frequently recurring themes related to the definition of the PRL included 1) it is a retinal area used for fixation, 2) it is task specific, 3) more than one PRL can be used, 4) it is a well-defined region of retina, and 5) the same PRL is used on repeated testing.
Based on the responses received, a consensus definition of the PRL is proposed. It is suggested that researchers define the PRL carefully in experimental reports and an instrument that images the retina is used to define the location of the PRL.
Retina (Philadelphia, Pa.) 05/2011; 31(10):2109-14. · 2.93 Impact Factor
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ABSTRACT: To what extent does attention modulate neural activity in early visual areas? fMRI measurements of attentional modulation in primary visual cortex (V1) show large effects, while single unit recordings show much smaller ones. This discrepancy suggests that fMRI measures of attention may be inflated, perhaps by activity related to other processes. To test whether effects measured with fMRI actually reflect attentional enhancement, we used a rapid acquisition protocol to determine their timing. Subjects were presented with two stimuli on either side of fixation and were cued to attend one and ignore the other. Attended stimuli showed a greater magnitude of response in V1, but this increase was delayed, by roughly one second in time, relative to both unattended responses and response increases due to boosting stimulus contrast. These results suggest that fMRI measurements of attention may primarily depend upon other processes that take a relatively long time to feed back to V1. Our results demonstrate the importance of using the fine timing information available in the fMRI response.
NeuroImage 04/2011; 57(3):1177-83. · 5.89 Impact Factor
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ABSTRACT: There is mounting evidence that visual perception abnormalities in schizophrenia are partly explained by a dysfunction of the lateral occipital complex (LO). We previously demonstrated that schizophrenia patients had broader topography and reduced magnitude of activity of LO. However, the functional connectivity of LO with other brain regions during visual perception has not been directly investigated in schizophrenia.
Eighteen patients with schizophrenia and eighteen matched controls performed a backward masking task during functional magnetic resonance imaging (fMRI). Stimulus onset asynchronies were manipulated to change the level of target visibility. To examine connectivity with LO function we conducted psychophysiological interactions (PPI) analyses using: 1) a region of interest (ROI) approach and 2) a whole brain analysis. ROIs were defined based on a contrast of trials on which a target was presented versus null trials in which no stimuli were presented.
Eleven ROIs were identified. Both groups showed similar strength of coupling between LO and the 11 ROIs when visibility was not taken into account. Healthy controls showed clear changes in coupling between LO and prefrontal and parietal regions as a function of target visibility (higher coupling with more visible targets). In comparison, patients showed reduced dynamic coupling with LO in the right superior frontal gyrus (significant after correcting for multiple comparisons) and a trend for reduced coupling in the left precuneus and left inferior frontal regions. Whole brain analysis identified additional regions that showed dynamic coupling with LO in healthy controls, but not in patients.
The increased coupling between LO and higher-level parietal and prefrontal regions during visual awareness in healthy controls likely reflects visual reentrant processing. The lack of modulation of coupling between LO and key prefrontal and parietal regions found in schizophrenia may partly reflect abnormalities in LO tuning. The altered LO coupling may contribute to visual perception abnormalities in schizophrenia.
NeuroImage 12/2010; 55(3):1219-26. · 5.89 Impact Factor
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ABSTRACT: Training improves performance on most visual tasks. Such perceptual learning can modify how information is read out from, and represented in, later visual areas, but effects on early visual cortex are controversial. In particular, it remains unknown whether learning can reshape neural response properties in early visual areas independent from feedback arising in later cortical areas. Here, we tested whether learning can modify feedforward signals in early visual cortex as measured by the human electroencephalogram. Fourteen subjects were trained for >24 d to detect a diagonal grating pattern in one quadrant of the visual field. Training improved performance, reducing the contrast needed for reliable detection, and also reliably increased the amplitude of the earliest component of the visual evoked potential, the C1. Control orientations and locations showed smaller effects of training. Because the C1 arises rapidly and has a source in early visual cortex, our results suggest that learning can increase early visual area response through local receptive field changes without feedback from later areas.
Journal of Neuroscience 11/2010; 30(45):15080-4. · 7.11 Impact Factor
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ABSTRACT: Visual masking paradigms assess the early part of visual information processing, which may reflect vulnerability measures for schizophrenia. We examined the neural substrates of visual backward performance in unaffected sibling of schizophrenia patients using functional magnetic resonance imaging (fMRI).
Twenty-one unaffected siblings of schizophrenia patients and 19 healthy controls performed a backward masking task and three functional localizer tasks to identify three visual processing regions of interest (ROI): lateral occipital complex (LO), the motion-sensitive area, and retinotopic areas. In the masking task, we systematically manipulated stimulus onset asynchronies (SOAs). We analyzed fMRI data in two complementary ways: 1) an ROI approach for three visual areas, and 2) a whole-brain analysis.
The groups did not differ in behavioral performance. For ROI analysis, both groups increased activation as SOAs increased in LO. Groups did not differ in activation levels of the three ROIs. For whole-brain analysis, controls increased activation as a function of SOAs, compared with siblings in several regions (i.e., anterior cingulate cortex, posterior cingulate cortex, inferior prefrontal cortex, inferior parietal lobule).
The study found: 1) area LO showed sensitivity to the masking effect in both groups; 2) siblings did not differ from controls in activation of LO; and 3) groups differed significantly in several brain regions outside visual processing areas that have been related to attentional or re-entrant processes. These findings suggest that LO dysfunction may be a disease indicator rather than a risk indicator for schizophrenia.
Biological psychiatry 07/2010; 68(1):78-85. · 8.93 Impact Factor
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ABSTRACT: What happens to neurons in visual cortex when they are deprived of their preferred stimuli? Long-term deprivation during development, spanning weeks, reduces the number of neurons selective for the deprived orientation [1-4]. In contrast, short-term deprivation in adults, for periods of seconds, can increase neural sensitivity relative to a stimulated baseline [5]. Effects over intermediate timescales remain largely unexplored, however. Here we introduce a new method for manipulating the visual environment of adult humans and report effects of four hours of orientation-specific deprivation. Subjects wore a head-mounted video camera that fed into a laptop computer that drove a head-mounted display. Software filtered the video stream in real time, allowing subjects to interact with the world while being deprived of visual input at a specified orientation. Four hours in this environment increased sensitivity to the deprived orientation, which likely reflected an increase in responsiveness of neurons in early visual cortex. Our results help distinguish between two theories of neural adaptation: the response increase optimized the responses of individual neurons, rather than increasing the efficiency of the population code. Our method should be able to produce a wide range of environmental manipulations useful for studying many topics in perception.
Current biology: CB 11/2009; 19(22):1956-60. · 10.99 Impact Factor
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ABSTRACT: Studies examining medial temporal lobe (MTL) involvement in memory formation typically assess memory performance after a single, short delay. Thus, the relationship between MTL encoding activity and memory durability over time remains poorly characterized. To explore this relationship, we scanned participants using high-resolution functional imaging of the MTL as they encoded object pairs; using the remember/know paradigm, we then assessed memory performance for studied items both 10 min and 1 week later. Encoding trials were classified as either subsequently recollected across both delays, transiently recollected (i.e., recollected at 10 min but not after 1 week), consistently familiar, or consistently forgotten. Activity in perirhinal cortex (PRC) and a hippocampal subfield comprising the dentate gyrus and CA fields 2 and 3 reflected successful encoding only when items were recollected consistently across both delays. Furthermore, in PRC, encoding activity for items that later were consistently recollected was significantly greater than that for transiently recollected and consistently familiar items. Parahippocampal cortex, in contrast, showed a subsequent memory effect during encoding of items that were recollected after 10 min, regardless of whether they also were recollected after 1 week. These data suggest that MTL subfields contribute uniquely to the formation of memories that endure over time, and highlight a role for PRC in supporting subsequent durable episodic recollection.
Journal of Cognitive Neuroscience 11/2009; 22(11):2652-62. · 5.18 Impact Factor
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ABSTRACT: Perceptual systems can be altered by immersing observers in environments with statistical properties that differ from those naturally encountered. Here we present a novel method for placing observers in naturalistic audio visual environments whose statistics can be manipulated in very targeted ways. We present the results of a case study that used this method. Observers were exposed to an environment where there was a novel statistical relationship between two simple, visual patterns in otherwise natural scenes. Exposure to this altered environment strengthened perceptual interactions between the two patterns.
Vision research 05/2009; 49(14):1757-64. · 2.29 Impact Factor
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ABSTRACT: The human visual pathways that are specialized for object recognition stretch from lateral occipital cortex (LO) to the ventral surface of the temporal lobe, including the fusiform gyrus. Plasticity in these pathways supports the acquisition of visual expertise, but precisely how training affects the different regions remains unclear. We used functional magnetic resonance imaging to measure neural activity in both LO and the fusiform gyrus in radiologists as they detected abnormalities in chest radiographs. Activity in the right fusiform face area (FFA) correlated with visual expertise, measured as behavioral performance during scanning. In contrast, activity in left LO correlated negatively with expertise, and the amount of LO that responded to radiographs was smaller in experts than in novices. Activity in the FFA and LO correlated negatively in experts, whereas in novices, the 2 regions showed no stable relationship. Together, these results suggest that the FFA becomes more engaged and left LO less engaged in interpreting radiographic images over the course of training. Achieving expert visual performance may involve suppressing existing neural representations while simultaneously developing others.
Cerebral Cortex 04/2009; 19(11):2746-54. · 6.54 Impact Factor
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ABSTRACT: Memories for certain events tend to linger in rich, vivid detail, and retrieval of these memories includes a sense of re-experiencing the details of the event. Most events, however, are not retained in any detailed way for more than a few days. According to one theory, the hippocampus plays a specific role in supporting episodic retrieval, that is, the re-experiencing of an event as part of one's personal past. This theory predicts that as episodic memories fade over time and are reduced to feelings of familiarity, activity in the hippocampus should no longer be associated with retrieval. We used high-resolution functional imaging to explore neural activity in medial temporal lobe subregions while participants performed a recognition task at both a short (10-min) and long (1-week) study-test delay. For each recognized item, subjects made "Remember/Know" judgments, allowing us to distinguish between items that were consistently episodic across the two tests and items that were initially episodic, but later became merely familiar. Our results demonstrate that activity in the subiculum is specifically associated with episodic recollection. Overall, recollected items were associated with higher activity in the subiculum than other items. For transiently recollected items, there was a decrease in subicular activity across the 1-week delay as memory faded from recollection to familiarity, whereas consistently recollected items were associated with enhanced subicular activity at both delays. These results provide evidence of a link between subicular activation and recollective experience.
Hippocampus 11/2008; 19(3):265-72. · 5.18 Impact Factor
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ABSTRACT: The neural mechanisms responsible for unifying noncontiguous regions of a visual image into a percept of a single surface remain largely unknown. To investigate these mechanisms, we used a novel stimulus in which local luminance was the only cue for surface segmentation. Subjects viewed an array of small adjoining elements that were randomly assigned as either surface or noise every 100 ms. On each trial, the luminance of surface elements was fixed to a single value and the luminance of noise elements was randomly assigned. As the ratio of surface to noise elements changed, subjects perceived either a surface embedded in noise or noise alone. In three functional magnetic resonance imaging (fMRI) experiments, early visual area V1 responded most strongly during trials with a low surface-to-noise ratio while later areas responded most strongly during trials with a high ratio. Furthermore, even at identical surface-to-noise ratios, responses in area V4 were higher during trials in which the subject perceived a surface than during trials in which the subject did not. Early visual areas did not show this pattern. These results suggest that visual area V4 contains neurons critical for the representation of surfaces.
Journal of Vision 02/2008; 8(7):28.1-9. · 3.38 Impact Factor
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ABSTRACT: Because of well-known nonlinearities in fMRI, responses measured with rapid event-related designs are smaller than responses measured with spaced designs. Surprisingly, no study to date has tested whether rapid designs also change the pattern of responses across different stimulus conditions. Here we report the results of such a test. We measured cortical responses to a flickering checkerboard at different contrasts using rapid and spaced event-related fMRI. The relative magnitude of responses across contrast conditions differed between rapid and spaced designs. Modeling the effect of the rapid design as a scaling of stimulus strength provided a good account of the data. The data were less well fit by a model that scaled the strength of responses. A similar stimulus scaling model has explained effects of neural adaptation, which suggests that adaptation may account for the observed difference between rapid and spaced designs. In a second experiment, we changed the stimulus in ways known to reduce neural adaptation and found much smaller differences between the two designs. Stimulus scaling provides a simple way to account for nonlinearities in event-related fMRI and relate data from rapid designs to data gathered using slower presentation rates.
NeuroImage 02/2007; 34(2):651-60. · 5.89 Impact Factor
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ABSTRACT: Lesions to ventral occipital cortex can produce severe deficits in color vision, a syndrome known as cerebral achromatopsia. Because most studies examine relatively few cases, however, uncertainty remains about precisely which cortical loci, when damaged, produce the syndrome. In addition, the extents of the associated perceptual deficits remain unclear. To address these issues, we performed a meta-analysis of 92 case reports from the literature. The severity of color vision deficits of the cases varied greatly, although nearly all showed some deficit in color discrimination. Almost all cases tested also showed some loss of spatial vision. Lesion overlap analyses revealed a relatively small region of high overlap in ventral occipital cortex. The region of high overlap was located near areas identified by neuroimaging studies as important for color perception. For comparison, we performed a similar analysis of prosopagnosia, a disorder of face perception, and found several regions of high lesion overlap adjacent to the region associated with achromatopsia. Because the behavioral deficits in achromatopsia are often incomplete and never restricted to color vision, the region of high lesion overlap may be one critical stage within a stream of many visual areas that participate nonexclusively in color perception.
Cerebral Cortex 03/2006; 16(2):183-91. · 6.54 Impact Factor
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ABSTRACT: Vision uses specific image features or cues to infer physical properties of the world. Here, we use a novel illusion to show that occlusion, traditionally thought of as a cue to depth, is also a powerful cue to motion. A display of stacking disks that contains only occlusion as a cue to depth generates a vivid sense of movement that is likely computed in early or middle levels of visual processing.
Journal of Vision 02/2006; 6(5):649-52. · 3.38 Impact Factor
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ABSTRACT: The hippocampal formation performs two related but distinct memory functions: encoding of novel information and retrieval of episodes. Little evidence, however, resolves how these two processes are implemented within the same anatomical structure. Here we use high-resolution functional magnetic resonance imaging to show that distinct subregions of the hippocampus are differentially involved in encoding and retrieval. We found that regions early in the hippocampal circuit (dentate gyrus and CA fields 2 and 3) were selectively active during episodic memory formation, whereas a region later in the circuit (the subiculum) was active during the recollection of the learning episode. Different components of the hippocampal circuit likely contribute to different degrees to the two basic memory functions.
Journal of Neuroscience 04/2005; 25(13):3280-6. · 7.11 Impact Factor
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Stephen A Engel
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ABSTRACT: Primary visual cortex contains at least two distinct populations of color-selective cells: neurons in one have circularly symmetric receptive fields and respond best to reddish and greenish light, while neurons in another have oriented receptive fields and a variety of color preferences. The relative prevalence and perceptual roles of the two kinds of neurons remain controversial, however. We used fMRI and a selective adaptation technique to measure responses attributable to these two populations. The technique revealed evidence of adaptation in both populations and indicated that they each produced strong signals in V1 and other human visual areas. The activity of both sets of neurons was also reflected in color appearance measurements made with the same stimuli. Thus, both oriented and unoriented color-selective cells in V1 are important components of the neural pathways that underlie perception of color.
Neuron 03/2005; 45(4):613-23. · 14.74 Impact Factor
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ABSTRACT: In visual backward masking, the visibility of a briefly presented visual target is disrupted by a mask that is presented shortly thereafter. The goal of the current study was to identify regions in the human cortex that may provide the neural basis of visual masking. We searched for areas whose activity correlated with perception as we systematically varied the strength of masking. A total of 13 subjects performed a backward masking task during functional magnetic resonance imaging. Target and mask were presented at three delay intervals (34, 68, and 102 msec) and behavioral measures confirmed that the targets were more visible at longer masking intervals. Two sets of regions of interest were identified: Distinct regions in the visual cortex (V1/V2, LO, hMT+) were segregated using scans to localize visual processing drawn from the existing literature. Additional cortical regions were selected in a data-driven approach based on their activity during the backward masking task. For each set, we determined the regions whose magnitude of activation increased at longer masking intervals. Nine of the subjects provided valid behavioral performance data on the visual masking task and imaging data from these subjects were used for subsequent analysis. The scans of visual processing areas identified four regions, including: early visual areas (V1 and V2), the motion-sensitive regions in the lateral occipital (LO) lobe (hMT+), and two components (dorsal and ventral) of the object-sensitive region, LO. Of these, the ventral and dorsal LO regions were sensitive to the strength of the mask. For the data-driven approach, six regions were identified on the basis of a difference map in which all masking intervals were contrasted with rest. These included the inferior parietal, anterior cingulate, precentral, insula, thalamic, and occipital areas. The predicted effects of more activity with weaker masking were seen in the thalamus, inferior parietal, and anterior cingulate. This study isolated three types of visual processing areas. The first included regions that subserve key stages of vision (including object and motion processing). The second type responded to the presentation of brief ly presented visual stimuli, regardless of masking interval. The third type (selected from the first two) included regions sensitive to the interval between the target and mask. These latter regions (including ventral LO, inferior parietal, anterior cingulate, and thalamus) may form the neural substrate of backward masking.
Journal of Cognitive Neuroscience 02/2005; 17(1):13-23. · 5.18 Impact Factor
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ABSTRACT: Training can significantly improve performance on even the most basic visual tasks, such as detecting a faint patch of light or determining the orientation of a bar (for reviews, see ). The neural mechanisms of visual learning, however, remain controversial. One simple way to improve behavior is to increase the overall neural response to the trained stimulus by increasing the number or gain of responsive neurons. Learning of this type has been observed in other sensory modalities, where training increases the number of receptive fields that cover the trained stimulus. Here, we show that visual learning can selectively increase the overall response to trained stimuli in primary visual cortex (V1). We used functional magnetic resonance imaging (fMRI) to measure neural signals before and after one month of practice at detecting very low-contrast oriented patterns. Training increased V1 response for practiced orientations relative to control orientations by an average of 39%, and the magnitude of the change in V1 correlated moderately well with the magnitude of changes in detection performance. The elevation of V1 activity by training likely results from an increase in the number of neurons responding to the trained stimulus or an increase in response gain.
Current Biology 05/2004; 14(7):573-8. · 9.65 Impact Factor
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ABSTRACT: The medial temporal lobe (MTL) is critical in forming new memories, but how subregions within the MTL carry out encoding and retrieval processes in humans is unknown. Using new high-resolution functional magnetic resonance imaging (fMRI) acquisition and analysis methods, we identified mnemonic properties of different subregions within the hippocampal circuitry as human subjects learned to associate names with faces. The cornu ammonis (CA) fields 2 and 3 and the dentate gyrus were active relative to baseline only during encoding, and this activity decreased as associations were learned. Activity in the subiculum showed the same temporal decline, but primarily during retrieval. Our results demonstrate that subdivisions within the hippocampus make distinct contributions to new memory formation.
Science 02/2003; 299(5606):577-80. · 31.20 Impact Factor
