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ABSTRACT: Reward can influence visual performance, but the neural basis of this effect remains poorly understood. Here we used functional magnetic resonance imaging to investigate how rewarding feedback affected activity in distinct areas of human visual cortex, separating rewarding feedback events after correct performance from preceding visual events. Participants discriminated oriented gratings in either hemifield, receiving auditory feedback at trial end that signaled financial reward after correct performance. Greater rewards improved performance for all but the most difficult trials. Rewarding feedback increased blood-oxygen-level-dependent (BOLD) signals in striatum and orbitofrontal cortex. It also increased BOLD signals in visual areas beyond retinotopic cortex, but not in primary visual cortex representing the judged stimuli. These modulations were seen at a time point in which no visual stimuli were presented or expected, demonstrating a novel type of activity change in visual cortex that cannot reflect modulation of response to incoming or anticipated visual stimuli. Rewarded trials led on the next trial to improved performance and enhanced visual activity contralateral to the judged stimulus, for retinotopic representations of the judged visual stimuli in V1. Our findings distinguish general effects in nonretinotopic visual cortex when receiving rewarding feedback after correct performance from consequences of reward for spatially specific responses in V1.
Journal of Neurophysiology 09/2010; 104(3):1746-57. · 3.32 Impact Factor
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ABSTRACT: Spatial models of functional magnetic resonance imaging (fMRI) data allow one to estimate the spatial smoothness of general linear model (GLM) parameters and eschew pre-process smoothing of data entailed by conventional mass-univariate analyses. Recently diffusion-based spatial priors [Harrison, L.M., Penny, W., Daunizeau, J., and Friston, K.J. (2008). Diffusion-based spatial priors for functional magnetic resonance images. NeuroImage.] were proposed, which provide a way to formulate an adaptive spatial basis, where the diffusion kernel of a weighted graph-Laplacian (WGL) is used as the prior covariance matrix over GLM parameters. An advantage of these is that they can be used to relax the assumption of isotropy and stationarity implicit in smoothing data with a fixed Gaussian kernel. The limitation of diffusion-based models is purely computational, due to the large number of voxels in a brain volume. One solution is to partition a brain volume into slices, using a spatial model for each slice. This reduces computational burden by approximating the full WGL with a block diagonal form, where each block can be analysed separately. While fMRI data are collected in slices, the functional structures exhibiting spatial coherence and continuity are generally three-dimensional, calling for a more informed partition. We address this using the graph-Laplacian to divide a brain volume into sub-graphs, whose shape can be arbitrary. Their shape depends crucially on edge weights of the graph, which can be based on the Euclidean distance between voxels (isotropic) or on GLM parameters (anisotropic) encoding functional responses. The result is an approximation the full WGL that retains its 3D form and also has potential for parallelism. We applied the method to high-resolution (1 mm(3)) fMRI data and compared models where a volume was divided into either slices or graph-partitions. Models were optimized using Expectation-Maximization and the approximate log-evidence computed to compare these different ways to partition a spatial prior. The high-resolution fMRI data presented here had greatest evidence for the graph partitioned anisotropic model, which was best able to preserve fine functional detail.
NeuroImage 09/2008; 43(4):694-707. · 5.89 Impact Factor
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ABSTRACT: Summary:
Background: Regions in human frontal cortex may have modulatory top-down influences on retinotopic visual cortex, but to date neuroimaging methods have only been able to provide indirect evidence for such functional interactions between remote but interconnected brain regions. Here we combined transcranial magnetic stimulation (TMS) with concurrent functional magnetic resonance imaging (fMRI), plus psychophysics, to show that stimulation of the right human frontal eye-field (FEF) produced a characteristic topographic pattern of activity changes in retinotopic visual areas V1-V4, with functional consequences for visual perception.
Results: FEF TMS led to activity increases for retinotopic representations of the peripheral visual field, but to activity decreases for the central field, in areas V1-V4. These frontal influences on visual cortex occurred in a top-down manner, independently of visual input. TMS of a control site (vertex) did not elicit such visual modulations, and saccades, blinks, or pupil dilation could not account for our results. Finally, the effects of FEF TMS on activity in retinotopic visual cortex led to a behavioral prediction that we confirmed psychophysically by showing that TMS of the frontal site (again compared with vertex) enhanced perceived contrast for peripheral relative to central visual stimuli.
Conclusions: Our results provide causal evidence that circuits originating in the human FEF can modulate activity in retinotopic visual cortex, in a manner that differentiates the central and peripheral visual field, with functional consequences for perception. More generally, our study illustrates how the new approach of concurrent TMS-fMRI can now reveal causal interactions between remote but interconnected areas of the human brain.
Ruff, C.C. and Blankenburg, F. and Bjoertomt, O. and Bestmann, S. and Freeman, E. and Haynes, J.-D. and Rees, H.G. and Josephs, O. and Deichmann, R. and Driver, J. (2006) Concurrent TMS-fMRI and psychophysics reveal frontal influences on human retinotopic visual cortex. Current Biology, 16 (15). pp. 1479-1488. ISSN 09609822.
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Pleger, B. and Blankenburg, F. and Bestmann, S. and Ruff, C.C. and Wiech, K. and Stephan, K.E. and Friston, K.J. and Dolan, R.J. (2006) Repetitive transcranial magnetic stimulation-induced changes in sensorimotor coupling parallel improvements of somatosensation in humans. Journal of Neuroscience, 26 (7). pp.1945 - 1952. ISSN 02706474.
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ABSTRACT: It has often been proposed that regions of the human parietal and/or frontal lobe may modulate activity in visual cortex, for example during selective attention or saccade preparation. However, direct evidence for such causal claims is largely missing in human studies, and it remains unclear to what degree the putative roles of parietal and frontal regions in modulating visual cortex may differ. Here we used concurrent TMS-fMRI to show that stimulating right human intra-parietal sulcus (IPS, at a site previously implicated in attention) elicits a pattern of activity changes in visual cortex that strongly depends on current visual context. Increased intensity of IPS TMS affected the BOLD signal in V5/MT+ only when moving stimuli were present to drive this visual region, whereas TMS-elicited BOLD signal changes were observed in areas V1-V4 only during the absence of visual input. These influences of IPS-TMS upon remote visual cortex differed significantly from corresponding effects of frontal (eye field) TMS, in terms of how they related to current visual input and their spatial topography for retinotopic areas V1-V4. Our results show directly that parietal and frontal regions can indeed have distinct patterns of causal influence upon functional activity in human visual cortex.
Ruff, C.C. and Bestmann, S. and Blankenburg, F. and Bjoertomt, O. and Josephs, O. and Weiskopf, N. and Deichmann, R. and Driver, J. (2008) Distinct causal influences of parietal and frontal brain regions on activity in human retinotopic visual cortex. Cerebral Cortex, 18 (4). pp. 817-827. ISSN 10473211.
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ABSTRACT: We used functional magnetic resonance imaging (fMRI) to study neural correlates of a robust somatosensory illusion that can dissociate tactile perception from physical stimulation. Repeated rapid stimulation at the wrist, then near the elbow, can create the illusion of touches at intervening locations along the arm, as if a rabbit hopped along it. We examined brain activity in humans using fMRI, with improved spatial resolution, during this version of the classic cutaneous rabbit illusion. As compared with control stimulation at the same skin sites (but in a different order that did not induce the illusion), illusory sequences activated contralateral primary somatosensory cortex, at a somatotopic location corresponding to the filled-in illusory perception on the forearm. Moreover, the amplitude of this somatosensory activation was comparable to that for veridical stimulation including the intervening position on the arm. The illusion additionally activated areas of premotor and prefrontal cortex. These results provide direct evidence that illusory somatosensory percepts can affect primary somatosensory cortex in a manner that corresponds somatotopically to the illusory percept.
Blankenburg, F. and Ruff, C.C. and Deichmann, R. and Rees, G. and Driver, J. (2006) The cutaneous 'rabbit' illusion affects human primary sensory cortex somatopically. PLoS Biology, 4 (3). 0460-0466. ISSN 15449173.
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ABSTRACT: Cognitive processes, such as spatial attention, are thought to rely on extended networks in the human brain. Both clinical data from lesioned patients and fMRI data acquired when healthy subjects perform particular cognitive tasks typically implicate a wide expanse of potentially contributing areas, rather than just a single brain area. Conversely, evidence from more targeted interventions, such as transcranial magnetic stimulation (TMS) or invasive microstimulation of the brain, or selective study of patients with highly focal brain damage, can sometimes indicate that a single brain area may make a key contribution to a particular cognitive process. But this in turn raises questions about how such a brain area may interface with other interconnected areas within a more extended network to support cognitive processes. Here, we provide a brief overview of new approaches that seek to characterise the causal role of particular brain areas within networks of several interacting areas, by measuring the effects of manipulations for a targeted area on function in remote interconnected areas. In human participants, these approaches include concurrent TMS-fMRI and TMS-EEG, as well as combination of the focal lesion method in selected patients with fMRI and/or EEG measures of the functional impact from the lesion on interconnected intact brain areas. Such approaches shed new light on how frontal cortex and parietal cortex modulate sensory areas in the service of attention and cognition, for the normal and damaged human brain.
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ABSTRACT: Combining transcranial magnetic stimulation (TMS) with concurrent functional magnetic resonance imaging (fMRI) allows study of how local brain stimulation may causally affect activity in remote brain regions. Here, we applied bursts of high- or low-intensity TMS over right posterior parietal cortex, during a task requiring sustained covert visuospatial attention to either the left or right hemifield, or in a neutral control condition, while recording blood oxygenation-level-dependent signal with a posterior MR surface coil. As expected, the active attention conditions activated components of the well-described "attention network," as compared with the neutral baseline. Also as expected, when comparing left minus right attention, or vice versa, contralateral occipital visual cortex was activated. The critical new finding was that the impact of high- minus low-intensity parietal TMS upon these visual regions depended on the currently attended side. High- minus low-intensity parietal TMS increased the difference between contralateral versus ipsilateral attention in right extrastriate visual cortex. A related albeit less pronounced pattern was found for left extrastriate visual cortex. Our results confirm that right human parietal cortex can exert attention-dependent influences on occipital visual cortex and provide a proof of concept for the use of concurrent TMS-fMRI in studying how remote influences can vary in a purely top-down manner with attentional demands.
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ABSTRACT: Interplay between the cerebral hemispheres is vital for coordinating perception and behavior. One influential account holds that the hemispheres engage in rivalry, each inhibiting the other. In the somatosensory domain, a seminal paper claimed to demonstrate such interhemispheric rivalry, reporting improved tactile detection sensitivity on the right hand after transcranial magnetic stimulation (TMS) to the right parietal lobe (Seyal, Ro, & Rafal, 1995). Such improvement in tactile detection ipsilateral to TMS could follow from interhemispheric rivalry, if one assumes that TMS disrupted cortical processing under the coil and thereby released the other hemisphere from inhibition. Here we extended the study by Seyal et al. (1995) to determine the effects of right parietal TMS on tactile processing for either hand, rather than only the ipsilateral hand. We performed two experiments applying TMS in the context of median-nerve stimulation; one experiment required somatosensory detection, the second somatosensory intensity discrimination. We found different TMS effects on detection versus discrimination, but neither set of results followed the prediction from hemispheric rivalry that enhanced performance for one hand should invariably be associated with impaired performance for the other hand, and vice-versa. Our results argue against a strict rivalry interpretation, instead suggesting that parietal TMS can provide a pedestal-like increment in somatosensory response.
