{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T20:37:07Z","timestamp":1791319027138,"version":"4.3.1"},"reference-count":30,"publisher":"American Association for the Advancement of Science (AAAS)","issue":"5482","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Science"],"published-print":{"date-parts":[[2000,8,18]]},"abstract":"<jats:p>A sudden touch on one hand can improve vision near that hand, revealing crossmodal links in spatial attention. It is often assumed that such links involve only multimodal neural structures, but unimodal brain areas may also be affected. We tested the effect of simultaneous visuo-tactile stimulation on the activity of the human visual cortex. Tactile stimulation enhanced activity in the visual cortex, but only when it was on the same side as a visual target. Analysis of effective connectivity between brain areas suggests that touch influences unimodal visual cortex via back-projections from multimodal parietal areas. This provides a neural explanation for crossmodal links in spatial attention.<\/jats:p>","DOI":"10.1126\/science.289.5482.1206","type":"journal-article","created":{"date-parts":[[2002,7,27]],"date-time":"2002-07-27T05:37:46Z","timestamp":1027748266000},"page":"1206-1208","source":"Crossref","is-referenced-by-count":473,"title":["Modulation of Human Visual Cortex by Crossmodal Spatial Attention"],"prefix":"10.1126","volume":"289","author":[{"given":"Emiliano","family":"Macaluso","sequence":"first","affiliation":[{"name":"Institute of Cognitive Neuroscience, University College London, UK."},{"name":"Wellcome Department of Cognitive Neurology, Institute of Neurology, London, UK."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chris D.","family":"Frith","sequence":"additional","affiliation":[{"name":"Wellcome Department of Cognitive Neurology, Institute of Neurology, London, UK."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jon","family":"Driver","sequence":"additional","affiliation":[{"name":"Institute of Cognitive Neuroscience, University College London, UK."}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"221","reference":[{"key":"e_1_3_1_2_2","unstructured":"S. A. Hillyard G. R. Mangun M. G. Woldorff S. J. Luck in The Cognitive Neurosciences M. S. Gazzaniga Ed. (MIT Press Cambridge MA 1995) pp. 665\u2013681."},{"key":"e_1_3_1_3_2","doi-asserted-by":"crossref","unstructured":"H. E. Pashler The Psychology of Attention (MIT Press Cambridge MA 1998).","DOI":"10.7551\/mitpress\/5677.001.0001"},{"key":"e_1_3_1_4_2","doi-asserted-by":"crossref","first-page":"1231","DOI":"10.1016\/0028-3932(89)90035-3","volume":"27","author":"Butter C. M.","year":"1989","unstructured":"Butter C. M., Buchtel H. A., Santucci R., Neuropsychologia 27, 1231 (1989).","journal-title":"Neuropsychologia"},{"key":"e_1_3_1_5_2","doi-asserted-by":"crossref","first-page":"544","DOI":"10.3758\/BF03206045","volume":"60","author":"Spence C.","year":"1998","unstructured":"Spence C., Nicholls M. E., Gillespie N., Driver J., Percept. Psychophys. 60, 544 (1998).","journal-title":"Percept. Psychophys."},{"key":"e_1_3_1_6_2","doi-asserted-by":"crossref","first-page":"461","DOI":"10.1016\/0028-3932(89)90051-1","volume":"27","author":"Farah M. J.","year":"1989","unstructured":"Farah M. J., Wong A. B., Monheit M. A., Morrow L. A., Neuropsychologia 27, 461 (1989).","journal-title":"Neuropsychologia"},{"key":"e_1_3_1_7_2","unstructured":"M. I. Posner in Cognitive Neuropsychology and Neurolinguistics: Advances in Models of Cognitive Functions and Impairment A. Caramazza Ed. (Erlbaum Hillsdale NJ 1990) pp. 187\u2013210."},{"key":"e_1_3_1_8_2","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1038\/41921","volume":"388","author":"Di Pellegrino G.","year":"1997","unstructured":"Di Pellegrino G., L\u00e0davas E., Farn\u00e9 A., Nature 388, 730 (1997).","journal-title":"Nature"},{"key":"e_1_3_1_9_2","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1016\/S1364-6613(98)01188-7","volume":"2","author":"Driver J.","year":"1998","unstructured":"Driver J., Spence C., Trends Cogn. Sci. 2, 254 (1998).","journal-title":"Trends Cogn. Sci."},{"key":"e_1_3_1_10_2","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1038\/11219","volume":"2","author":"Hahnloser R.","year":"1999","unstructured":"Hahnloser R., Douglas R. J., Mahowald M., Hepp K., Nature Neurosci. 2, 746 (1999).","journal-title":"Nature Neurosci."},{"key":"e_1_3_1_11_2","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1007\/BF00247570","volume":"69","author":"Haenny P. E.","year":"1988","unstructured":"Haenny P. E., Maunsell J. H. R., Schiller P. H., Exp. Brain Res. 69, 245 (1988).","journal-title":"Exp. Brain Res."},{"key":"e_1_3_1_12_2","unstructured":"Data acquisition. For each participant fMRI data were acquired in a single session lasting 28 min. Each of the four possible event types was repeated 50 times in randomly intermingled sequence. The interstimulus interval was 8.22 \u00b1 4.11 s (uniform distribution sampled at 85-ms intervals). Functional images were acquired with a 2T Magnetom VISION MRI scanner (Siemens Erlangen Germany). BOLD (blood oxygenation-level dependent) contrast was obtained with echo-planar T2* weighted imaging (EPI). For each participant we acquired 400 volumes in a transverse orientation with 48 slices and an effective repetition time (TR) of 4.11 s. The voxel size was 3 mm by 3 mm by 3 mm."},{"key":"e_1_3_1_13_2","unstructured":"Participants. Six neurologically healthy volunteers participated (mean age 27 years old; range 21 to 34 years old; all right-handed males). After receiving an explanation of the procedures participants gave written informed consent. The study was approved by the Joint Ethics Committee of The Institute of Neurology and The National Hospital for Neurology & Neurosurgery London."},{"key":"e_1_3_1_14_2","unstructured":"Stimuli and task. The visual stimulation consisted of light-emitting diodes (LEDs) flickering (at 10 Hz) for 300 ms in one visual field or the other. Two arrays of three LEDs each (arranged to form an equilateral triangle with sides of 2 cm) were placed 10 cm to the left or right of the central fixation point at 6 degrees of visual angle. The right array was attached to the right index finger where any concurrent tactile stimulation was delivered. This was a vibration (60 Hz) lasting 300 ms generated by a pulsed current passed through a coil in the magnetic resonance scanner's magnetic field. Participants viewed the LEDs and the fixation point via a mirror placed on top of the whole-head radio-frequency coil. The inside of the magnet was illuminated so participants could also see their right hand placed vertically on a plastic support with the right LEDs placed on the index finger. Through the mirror the right hand still appeared in the right visual hemifield next to the right LEDs. We took care to place the right light very near the right hand in this way as previous evidence suggests that visuo-tactile links depend on the proximity of visual stimuli to the current location of the hand in external space (7 8 20). Future work could vary this proximity systematically in our imaging paradigm. During scanning participants were asked to respond every time they saw the LEDs flickering by a button press with the (unseen) left index finger (25). Participants were told to ignore any tactile stimulation. The only other task requirement was to maintain central fixation throughout the experiment."},{"key":"e_1_3_1_15_2","unstructured":"fMRI analysis. Data were analyzed with SPM99b (www.fil.ion.ucl.ac.uk). To account for the delay between data acquisition in the top and bottom part of the brain acquisition timing was corrected using the top slice as reference (26) for each subject. The 400 volumes were realigned with the first as reference. To allow inter-subject analysis images were normalized to the Montreal Neurological Institute (MNI) standard space using the mean of the 400 functional images. All images were smoothed using an isotropic Gaussian kernel (full width at half maximum = 10 mm). Data were best fitted (least-squares fit) at every voxel using a linear combination of the effects of interest (multiple regression). These were the timing of each of the four event types convolved with the SPM99 standard hemodynamic response function. Each of these regressors represents the expected BOLD signal for each trial type given the slow hemodynamic response produced by the underlying neuronal activity. Because of the many voxels tested we used the Theory of Gaussian Fields to assign P values corrected for multiple comparisons. For the main effect of side of the visual stimulation and presence of the tactile stimulation correction at the cluster level was used (cluster size set by thresholding the SPM maps at P -uncorrected = 0.001). For the interaction between side and presence of the tactile stimulation the statistical threshold was initially set at P -uncorrected = 0.05. The search volume was restricted to areas showing an effect of side of the visual stimulation hence corrected P values were then assigned using a small volume correction (27). The search volume was confined to the cluster in the left lingual gyrus that showed a main effect of right versus left visual stimulation (447 voxels)."},{"key":"e_1_3_1_16_2","unstructured":"The activation clusters for the side of visual stimulation extended from the fourth occipital gyrus into the posterior lingual gyrus and posterior fusiform gyrus. Both clusters included several secondary maxima likely to include the borders of retinotopic visual areas V2v VP\/V3v and V4v (28) . Our peripheral and low-contrast visual stimulation did not produce any robust activation in contralateral striate cortex (V1)."},{"key":"e_1_3_1_17_2","unstructured":"To confirm the spatial specificity of the crossmodal effect three individuals participated in this follow-up experiment. The visual stimulation and task remained identical but here any tactile stimulation was applied to the left hand near the left light. Again the main effects for side of visual stimulation activated the contralateral lingual gyrus (right hemisphere: x y z = 28 \u201376 \u201310 Z score = 4.5; left hemisphere: x y z = \u201326 \u201372 \u201310 Z score = 4.4). Significant interaction between side of visual stimulation and presence of touch was now found in the right rather than left lingual gyrus ( x y z = 26 \u201378 \u201312; Z score = 2.9; P -corrected = 0.046); again this is contralateral to the stimulated hand. Maximal response was detected here when left visual targets were coupled with left touch. Responses to right visual stimuli in the left lingual gyrus showed a nonsignificant trend for the reverse pattern of suppression by left touch contrasting with the significant amplification by right touch in this area for the main experiment."},{"key":"e_1_3_1_18_2","doi-asserted-by":"crossref","unstructured":"R. C. Coghill et al. J. Neurosci. 14 4095 (1994).","DOI":"10.1523\/JNEUROSCI.14-07-04095.1994"},{"key":"e_1_3_1_19_2","unstructured":"Changes in effective connectivity. To highlight areas mediating the crossmodal effects observed in the lingual gyrus we used an effective connectivity model. Linear regression detected any voxel that changed its contribution (29) to the signal in the left lingual gyrus depending on the trial type. This was done by fitting the full linear model at every voxel (14) but with the addition of two new regressors. One regressor was the activity at the maxima of the cluster showing an interaction between side of the visual stimulation and presence of tactile input (i.e. the cluster in the left lingual gyrus). The second regressor was created by multiplying the activity of the maxima in the left lingual gyrus with the difference of the regressors modeling the effect of spatially congruent versus incongruent bimodal conditions. The latter regressor represents the effect of interest i.e. the interaction between activity in the left lingual gyrus and the spatial congruence of the bimodal stimulation. Significant fit for this new regressor indicates (29) a trial-specific change in the contribution of any reported area to the activity in the left lingual gyrus. We refer to this effect as a change in the \u201ccoupling\u201d between the two areas. We ensured that this revealed only higher coupling during the bimodal spatially coherent versus spatially incoherent stimulation by specifically testing for positive regression slopes. For this analysis the SPM threshold was set at P -uncorrected = 0.001."},{"key":"e_1_3_1_20_2","doi-asserted-by":"crossref","unstructured":"R. E. Passingham in Comparative Neuropsychology D. Milner Ed. (Oxford Univ. Press Oxford 1998) pp. 271\u2013298.","DOI":"10.1093\/acprof:oso\/9780198524113.003.0015"},{"key":"e_1_3_1_21_2","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1016\/S0959-4388(98)80140-2","volume":"8","author":"Graziano M. S.","year":"1998","unstructured":"Graziano M. S., Gross C. G., Curr. Opin. Neurobiol. 8, 195 (1998).","journal-title":"Curr. Opin. 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The tactile events speeded up visual RTs especially for visual targets on the same side as the stimulated hand. In the absence of right touch left lights were responded to faster than right lights due to their compatibility with the left button press (interparticipant means of median RTs were 365 ms and 374 ms respectively; P = 0.04). When right touch was added this difference disappeared (means of 338 ms and 340 ms n.s.) because adding touch produced the greatest benefit for the closest light stimulus as observed in other psychophysical studies (4 8)."},{"key":"e_1_3_1_27_2","first-page":"125","volume":"9","author":"Henson R. N. A.","year":"1999","unstructured":"Henson R. N. A., Buechel C., Josephs O., Friston K., NeuroImage 9, 125 (1999).","journal-title":"NeuroImage"},{"key":"e_1_3_1_28_2","doi-asserted-by":"crossref","unstructured":"K. J. Worsley et al. Hum. Brain Mapp. 4 58 (1996).","DOI":"10.1002\/(SICI)1097-0193(1996)4:1<58::AID-HBM4>3.3.CO;2-L"},{"key":"e_1_3_1_29_2","doi-asserted-by":"crossref","unstructured":"E. A. DeYoe et al. Proc. Nat. Acad. Sci. U.S.A. 93 2382 (1996).","DOI":"10.1073\/pnas.93.6.2382"},{"key":"e_1_3_1_30_2","doi-asserted-by":"crossref","unstructured":"K. J. Friston et al. NeuroImage 6 218 (1997).","DOI":"10.1006\/nimg.1997.0291"},{"key":"e_1_3_1_31_2","unstructured":"The Functional Imaging Laboratory and C.D.F. were supported by the Wellcome Trust. E.M. and J.D. were supported by a Programme Grant from the Medical Research Council (UK). E.M. was also supported by a Research Training Grant from The Swiss National Research Foundation and a personal grant from the Janggen-Poehn-Stiftung (Switzerland)."}],"container-title":["Science"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.science.org\/doi\/pdf\/10.1126\/science.289.5482.1206","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,1,13]],"date-time":"2024-01-13T04:37:08Z","timestamp":1705120628000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.science.org\/doi\/10.1126\/science.289.5482.1206"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2000,8,18]]},"references-count":30,"journal-issue":{"issue":"5482","published-print":{"date-parts":[[2000,8,18]]}},"alternative-id":["10.1126\/science.289.5482.1206"],"URL":"https:\/\/doi.org\/10.1126\/science.289.5482.1206","relation":{},"ISSN":["0036-8075","1095-9203"],"issn-type":[{"value":"0036-8075","type":"print"},{"value":"1095-9203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2000,8,18]]}}}