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ABSTRACT: Despite considerable work, the neural basis of perceptual learning remains uncertain. For visual learning, although some studies suggested that changes in early sensory representations are responsible, other studies point to decision-level reweighting of perceptual readout. These competing possibilities have not been examined in other sensory systems, investigating which could help resolve the issue. Here we report a study of human tactile microspatial learning in which participants achieved >six-fold decline in acuity threshold after multiple training sessions. Functional magnetic resonance imaging was performed during performance of the tactile microspatial task and a control, tactile temporal task. Effective connectivity between relevant brain regions was estimated using multivariate, autoregressive models of hidden neuronal variables obtained by deconvolution of the hemodynamic response. Training-specific increases in task-selective activation assessed using the task × session interaction and associated changes in effective connectivity primarily involved subcortical and anterior neocortical regions implicated in motor and/or decision processes, rather than somatosensory cortical regions. A control group of participants tested twice, without intervening training, exhibited neither threshold improvement nor increases in task-selective activation. Our observations argue that neuroplasticity mediating perceptual learning occurs at the stage of perceptual readout by decision networks. This is consonant with the growing shift away from strictly modular conceptualization of the brain toward the idea that complex network interactions underlie even simple tasks. The convergence of our findings on tactile learning with recent studies of visual learning reconciles earlier discrepancies in the literature on perceptual learning.
Journal of Neuroscience 03/2013; 33(12):5387-98. · 7.11 Impact Factor
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ABSTRACT: In this chapter we review the empirical evidence for visual cortical involvement in tactile perception in normally sighted humans. This evidence stems from functional neuroimaging studies in humans, which have shown that visual cortical regions are recruited during a number of tactile tasks and in a manner that is highly task-specific. Studies of the sighted and of individuals with visual deprivation of long (see Fridman, Celnik, & Cohen, Chap. 45, this volume) or short term are mutually complementary. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
10/2012;
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ABSTRACT: Although the target article provides strong evidence against the locationist view, evidence for the constructionist view is inconclusive, because co-activation of brain regions does not necessarily imply connectivity between them. We propose a rigorous approach wherein connectivity between co-activated regions is first modeled using exploratory Granger causality, and then confirmed using dynamic causal modeling or Bayesian modeling.
Behavioral and Brain Sciences 06/2012; 35(3):148-9. · 25.06 Impact Factor
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ABSTRACT: Learning and memory deficits typify patients with mild cognitive impairment (MCI) and are generally attributed to medial temporal lobe dysfunction. Although the hippocampus is perhaps the most commonly studied neuroanatomical structure in these patients, there have been few attempts to identify rehabilitative interventions that facilitate its functioning. Here, we present results from a randomized, controlled, single-blind study in which patients with MCI and healthy elderly controls (HEC) were randomized to either three sessions of mnemonic strategy training (MS) or a matched-exposure control group (XP). All participants underwent pre- and posttraining fMRI scanning as they encoded and retrieved object-location associations. For the current report, fMRI analyses were restricted to the hippocampus, as defined anatomically. Before training, MCI patients showed reduced hippocampal activity during both encoding and retrieval, relative to HEC. Following training, the MCI MS group demonstrated increased activity during both encoding and retrieval. There were significant differences between the MCI MS and MCI XP groups during retrieval, especially within the right hippocampus. Thus, MS facilitated hippocampal functioning in a partially restorative manner. We conclude that cognitive rehabilitation techniques may help mitigate hippocampal dysfunction in MCI patients.
Hippocampus 02/2012; 22(8):1652-8. · 5.18 Impact Factor
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ABSTRACT: Conceptual metaphor theory suggests that knowledge is structured around metaphorical mappings derived from physical experience. Segregated processing of object properties in sensory cortex allows testing of the hypothesis that metaphor processing recruits activity in domain-specific sensory cortex. Using functional magnetic resonance imaging (fMRI) we show that texture-selective somatosensory cortex in the parietal operculum is activated when processing sentences containing textural metaphors, compared to literal sentences matched for meaning. This finding supports the idea that comprehension of metaphors is perceptually grounded.
Brain and Language 02/2012; 120(3):416-21. · 3.12 Impact Factor
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ABSTRACT: Segregation of information flow along a dorsally directed pathway for processing object location and a ventrally directed pathway for processing object identity is well established in the visual and auditory systems, but is less clear in the somatosensory system. We hypothesized that segregation of location vs. identity information in touch would be evident if texture is the relevant property for stimulus identity, given the salience of texture for touch. Here, we used functional magnetic resonance imaging (fMRI) to investigate whether the pathways for haptic and visual processing of location and texture are segregated, and the extent of bisensory convergence. Haptic texture-selectivity was found in the parietal operculum and posterior visual cortex bilaterally, and in parts of left inferior frontal cortex. There was bisensory texture-selectivity at some of these sites in posterior visual and left inferior frontal cortex. Connectivity analyses demonstrated, in each modality, flow of information from unisensory non-selective areas to modality-specific texture-selective areas and further to bisensory texture-selective areas. Location-selectivity was mostly bisensory, occurring in dorsal areas, including the frontal eye fields and multiple regions around the intraparietal sulcus bilaterally. Many of these regions received input from unisensory areas in both modalities. Together with earlier studies, the activation and connectivity analyses of the present study establish that somatosensory processing flows into segregated pathways for location and object identity information. The location-selective somatosensory pathway converges with its visual counterpart in dorsal frontoparietal cortex, while the texture-selective somatosensory pathway runs through the parietal operculum before converging with its visual counterpart in visual and frontal cortex. Both segregation of sensory processing according to object property and multisensory convergence appear to be universal organizing principles.
NeuroImage 07/2011; 57(2):462-75. · 5.89 Impact Factor
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ABSTRACT: In this chapter, the authors use the computation, anatomy, and physiology (CAP) principles to consider the impact of common clinical problems on action. They focus on 3 major syndromes: paresis, apraxia, and ataxia. They also review mechanisms that could account for spontaneous recovery, using what is known about the best-studied clinical dysfunction--paresis--and also ataxia. Together, this and the previous chapter lay the groundwork for the third chapter in this series, which reviews the relevant rehabilitative interventions.
Neurorehabilitation and neural repair 06/2011; 25(5 Suppl):21S-32S. · 4.49 Impact Factor
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ABSTRACT: Remembering the location of objects in the environment is both important in everyday life and difficult for patients with amnestic mild cognitive impairment (aMCI), a clinical precursor to Alzheimer's disease. To test the hypothesis that memory impairment for object location in aMCI reflects hippocampal dysfunction, we used an event-related functional magnetic resonance imaging paradigm to compare patients with aMCI and healthy elderly controls (HEC) as they encoded 90 ecologically relevant object-location associations (OLAs). Two additional OLAs, repeated a total of 45 times, served as control stimuli. Memory for these OLAs was assessed following a 1-h delay. The groups were well matched on demographics and brain volumetrics. Behaviorally, HEC remembered significantly more OLAs than did aMCI patients. Activity differences were assessed by contrasting activation for successfully encoded Novel stimuli vs. Repeated stimuli. The HEC demonstrated activity within object-related (ventral visual stream), spatial location-related (dorsal visual stream), and feature binding-related cortical regions (hippocampus and other memory-related regions) as well as in frontal cortex and associated subcortical structures. Activity in most of these regions correlated with memory test performance. Although the aMCI patients demonstrated a similar activation pattern, the HEC showed significantly greater activity within each of these regions. Memory test performance in aMCI patients, in contrast to the HEC, was correlated with activity in regions involved in sensorimotor processing. We conclude that aMCI patients demonstrate widespread cerebral dysfunction, not limited to the hippocampus, and rely on encoding-related mechanisms that differ substantially from healthy individuals.
Neuropsychologia 04/2011; 49(9):2349-61. · 3.64 Impact Factor
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ABSTRACT: A recent study showed that people evaluate products more positively when they are physically associated with art images than similar non-art images. Neuroimaging studies of visual art have investigated artistic style and esthetic preference but not brain responses attributable specifically to the artistic status of images. Here we tested the hypothesis that the artistic status of images engages reward circuitry, using event-related functional magnetic resonance imaging (fMRI) during viewing of art and non-art images matched for content. Subjects made animacy judgments in response to each image. Relative to non-art images, art images activated, on both subject- and item-wise analyses, reward-related regions: the ventral striatum, hypothalamus and orbitofrontal cortex. Neither response times nor ratings of familiarity or esthetic preference for art images correlated significantly with activity that was selective for art images, suggesting that these variables were not responsible for the art-selective activations. Investigation of effective connectivity, using time-varying, wavelet-based, correlation-purged Granger causality analyses, further showed that the ventral striatum was driven by visual cortical regions when viewing art images but not non-art images, and was not driven by regions that correlated with esthetic preference for either art or non-art images. These findings are consistent with our hypothesis, leading us to propose that the appeal of visual art involves activation of reward circuitry based on artistic status alone and independently of its hedonic value.
NeuroImage 03/2011; 55(1):420-33. · 5.89 Impact Factor
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ABSTRACT: Visual imagery comprises object and spatial dimensions. Both types of imagery encode shape but a key difference is that object imagers are more likely to encode surface properties than spatial imagers. Since visual and haptic object representations share many characteristics, we investigated whether haptic and multisensory representations also share an object-spatial continuum. Experiment 1 involved two tasks in both visual and haptic within-modal conditions, one requiring discrimination of shape across changes in texture, the other discrimination of texture across changes in shape. In both modalities, spatial imagers could ignore changes in texture but not shape, whereas object imagers could ignore changes in shape but not texture. Experiment 2 re-analyzed a cross-modal version of the shape discrimination task from an earlier study. We found that spatial imagers could discriminate shape across changes in texture but object imagers could not and that the more one preferred object imagery, the more texture changes impaired discrimination. These findings are the first evidence that object and spatial dimensions of imagery can be observed in haptic and multisensory representations.
Experimental Brain Research 03/2011; 213(2-3):267-73. · 2.39 Impact Factor
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ABSTRACT: Behavioral studies show that the unisensory representations underlying within-modal visual and haptic object recognition are strikingly similar in terms of view- and size-sensitivity, and integration of structural and surface properties. However, the basis for these attributes differs in each modality, indicating that while these representations are functionally similar, they are not identical. Imaging studies reveal bisensory, visuo-haptic object selectivity, notably in the lateral occipital complex and the intraparietal sulcus, that suggests a shared representation of objects. Such a multisensory representation could underlie visuo-haptic cross-modal object recognition. In this chapter, we compare visual and haptic within-modal object recognition and trace a progression from functionally similar but separate unisensory representations to a shared multisensory representation underlying cross-modal object recognition as well as view-independence, regardless of modality. We outline, and provide evidence for, a model of multisensory object recognition in which representations are flexibly accessible via top-down or bottom-up processing, the choice of route being influenced by object familiarity and individual preference along the object-spatial continuum of mental imagery.
Progress in brain research 01/2011; 191:165-76. · 3.04 Impact Factor
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ABSTRACT: Mild cognitive impairment (MCI) is often a precursor to Alzheimer disease. Little research has examined the efficacy of cognitive rehabilitation in patients with MCI, and the relevant neural mechanisms have not been explored. The authors previously showed the behavioral efficacy of cognitive rehabilitation using mnemonic strategies for face-name associations in patients with MCI. Here, the authors used functional magnetic resonance imaging (fMRI) to test whether there were training-specific changes in activation and connectivity within memory-related areas.
A total of 6 patients with amnestic, multidomain MCI underwent pretraining and posttraining fMRI scans, during which they encoded 90 novel face-name pairs and completed a 4-choice recognition memory test immediately after scanning. Patients were taught mnemonic strategies for half the face-name pairs during 3 intervening training sessions.
Training-specific effects comprised significantly increased activation within a widespread cerebral cortical network involving medial frontal, parietal, and occipital regions; the left frontal operculum and angular gyrus; and regions in the left lateral temporal cortex. Increased activation common to trained and untrained stimuli was found in a separate network involving inferior frontal, lateral parietal, and occipital cortical regions. Effective connectivity analysis using multivariate, correlation-purged Granger causality analysis revealed generally increased connectivity after training, particularly involving the middle temporal gyrus and foci in the occipital cortex and the precuneus.
The authors' findings suggest that the effectiveness of explicit-memory training in patients with MCI is associated with training-specific increases in activation and connectivity in a distributed neural system that includes areas involved in explicit memory.
Neurorehabilitation and neural repair 10/2010; 25(3):210-22. · 4.49 Impact Factor
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ABSTRACT: Effective connectivity in brain networks can be studied using Granger causality analysis, which is based on temporal precedence, while functional connectivity is usually derived using zero-lag correlation. Due to the smoothing of the neuronal activity by the hemodynamic response inherent in the functional magnetic resonance imaging (fMRI) acquisition process, Granger causality, as normally computed from fMRI data, may be contaminated by zero-lag correlation. Simulations performed in this paper showed that the zero-lag correlation does "leak" into estimates of time-lagged causality. To eliminate this leak, we introduce a method in which the zero-lag influences are explicitly modeled in the vector autoregressive model but omitted while calculating Granger causality. The effectiveness of this method is demonstrated using fMRI data obtained from healthy humans performing a verbal working memory task.
IEEE transactions on bio-medical engineering 06/2010; 57(6):1446-56. · 2.15 Impact Factor
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ABSTRACT: Because many visuospatial memory tests do not reliably detect right medial temporal lobe (MTL) dysfunction, we developed a novel object recognition test using complex three-dimensional stimuli. To influence encoding strategy, half the stimuli were multicolored (color towers) and accompanied by verbally based instructions, and half were gray (gray towers) and accompanied by visuospatially based instructions. In Experiment 1, healthy subjects completed the test while performing verbal or visuospatial interference tasks or without interference. In Experiment 2, patients with unilateral amygdalohippocampectomies for intractable epilepsy completed the test without interference. Results suggest that color tower recognition was partially dependent on verbal processing and sensitive to MTL lesions in general. Recognition of gray towers was reliant on visuospatial processing, and the decay in accuracy after a delay was sensitive and specific to right MTL lesions. These findings suggest that test stimuli such as three-dimensional objects can be useful in assessing right MTL dysfunction.
Epilepsy & Behavior 05/2010; 18(1-2):54-60. · 2.34 Impact Factor
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ABSTRACT: Object recognition studies have almost exclusively involved vision, focusing on shape rather than surface properties such as color. Visual object representations are thought to integrate shape and color information because changing the color of studied objects impairs their subsequent recognition. However, little is known about integration of surface properties into visuohaptic multisensory representations. Here, participants studied objects with distinct patterns of surface properties (color in Experiment 1, texture in Experiments 2 and 3) and had to discriminate between object shapes when color or texture schemes were altered in within-modal (visual and haptic) and cross-modal (visual study followed by haptic test and vice versa) conditions. In Experiment 1, color changes impaired within-modal visual recognition but had no effect on cross-modal recognition, suggesting that the multisensory representation was not influenced by modality-specific surface properties. In Experiment 2, texture changes impaired recognition in all conditions, suggesting that both unisensory and multisensory representations integrated modality-independent surface properties. However, the cross-modal impairment might have reflected either the texture change or a failure to form the multisensory representation. Experiment 3 attempted to distinguish between these possibilities by combining changes in texture with changes in orientation, taking advantage of the known view-independence of the multisensory representation, but the results were not conclusive owing to the overwhelming effect of texture change. The simplest account is that the multisensory representation integrates shape and modality-independent surface properties. However, more work is required to investigate this and the conditions under which multisensory integration of structural and surface properties occurs.
European Journal of Neuroscience 05/2010; 31(10):1882-8. · 3.63 Impact Factor
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ABSTRACT: This review focusses on cross-modal plasticity resulting from visual deprivation. This is viewed against the background of task-specific visual cortical recruitment that is routine during tactile tasks in the sighted and that may depend in part on visual imagery. Superior tactile perceptual performance in the blind may be practice-related, although there are unresolved questions regarding the effects of Braille-reading experience and the age of onset of blindness. While visual cortical areas are clearly more involved in tactile microspatial processing in the blind than in the sighted, it still remains unclear how to reconcile these tactile processes with the growing literature implicating visual cortical activity in a wide range of cognitive tasks in the blind, including those involving language, or with studies of short-term, reversible visual deprivation in the normally sighted that reveal plastic changes even over periods of hours or days.
Restorative neurology and neuroscience 01/2010; 28(2):271-81. · 2.51 Impact Factor
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ABSTRACT: In this work, we investigated the effect of the regional variability of the hemodynamic response on the sensitivity of Granger causality (GC) analysis of functional magnetic resonance imaging (fMRI) data to neuronal causal influences. We simulated fMRI data by convolving a standard canonical hemodynamic response function (HRF) with local field potentials (LFPs) acquired from the macaque cortex and manipulated the causal influence and neuronal delays between the LFPs, the hemodynamic delays between the HRFs, the signal-to-noise ratio (SNR), and the sampling period (TR) to assess the effect of each of these factors on the detectability of the neuronal delays from GC analysis of fMRI. In our first bivariate implementation, we assumed the worst-case scenario of the hemodynamic delay being at the empirical upper limit of its normal physiological range and opposing the direction of neuronal delay. We found that, in the absence of HRF confounds, even tens of milliseconds of neuronal delays can be inferred from fMRI. However, in the presence of HRF delays which opposed neuronal delays, the minimum detectable neuronal delay was hundreds of milliseconds. In our second multivariate simulation, we mimicked the real situation more closely by using a multivariate network of four time series and assumed the hemodynamic and neuronal delays to be unknown and drawn from a uniform random distribution. The resulting accuracy of detecting the correct multivariate network from fMRI was well above chance and was up to 90% with faster sampling. Generically, under all conditions, faster sampling and low measurement noise improved the sensitivity of GC analysis of fMRI data to neuronal causality.
NeuroImage 12/2009; 52(3):884-96. · 5.89 Impact Factor
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ABSTRACT: Although visual cortical engagement in haptic shape perception is well established, its relationship with visual imagery remains controversial. We addressed this using functional magnetic resonance imaging during separate visual object imagery and haptic shape perception tasks. Two experiments were conducted. In the first experiment, the haptic shape task employed unfamiliar, meaningless objects, whereas familiar objects were used in the second experiment. The activations evoked by visual object imagery overlapped more extensively, and their magnitudes were more correlated, with those evoked during haptic shape perception of familiar, compared to unfamiliar, objects. In the companion paper (Deshpande et al., this issue), we used task-specific functional and effective connectivity analyses to provide convergent evidence: these analyses showed that the neural networks underlying visual imagery were similar to those underlying haptic shape perception of familiar, but not unfamiliar, objects. We conclude that visual object imagery is more closely linked to haptic shape perception when objects are familiar, compared to when they are unfamiliar.
NeuroImage 11/2009; 49(3):1977-90. · 5.89 Impact Factor
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ABSTRACT: We previously showed that cross-modal recognition of unfamiliar objects is view-independent, in contrast to view-dependence within-modally, in both vision and haptics. Does the view-independent, bisensory representation underlying cross-modal recognition arise from integration of unisensory, view-dependent representations or intermediate, unisensory but view-independent representations? Two psychophysical experiments sought to distinguish between these alternative models. In both experiments, participants began from baseline, within-modal, view-dependence for object recognition in both vision and haptics. The first experiment induced within-modal view-independence by perceptual learning, which was completely and symmetrically transferred cross-modally: visual view-independence acquired through visual learning also resulted in haptic view-independence and vice versa. In the second experiment, both visual and haptic view-dependence were transformed to view-independence by either haptic-visual or visual-haptic cross-modal learning. We conclude that cross-modal view-independence fits with a model in which unisensory view-dependent representations are directly integrated into a bisensory, view-independent representation, rather than via intermediate, unisensory, view-independent representations.
Experimental Brain Research 06/2009; 198(2-3):329-37. · 2.39 Impact Factor
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ABSTRACT: This review surveys the recent literature on visuo-haptic convergence in the perception of object form, with particular reference to the lateral occipital complex (LOC) and the intraparietal sulcus (IPS) and discusses how visual imagery or multisensory representations might underlie this convergence. Drawing on a recent distinction between object- and spatially-based visual imagery, we propose a putative model in which LOtv, a subregion of LOC, contains a modality-independent representation of geometric shape that can be accessed either bottom-up from direct sensory inputs or top-down from frontoparietal regions. We suggest that such access is modulated by object familiarity: spatial imagery may be more important for unfamiliar objects and involve IPS foci in facilitating somatosensory inputs to the LOC; by contrast, object imagery may be more critical for familiar objects, being reflected in prefrontal drive to the LOC.
Brain Topography 04/2009; 21(3-4):269-74. · 3.45 Impact Factor
