Parietal rTMS distorts the mental number line: Simulating ‘spatial’ neglect in healthy subjects

Patients with left-sided visuospatial neglect, typically after damage to the right parietal lobe, show a systematic bias towards larger numbers when asked to bisect a numerical interval. This has been taken as further evidence for a spatial representation of numbers, perhaps akin to a mental number line with smaller numbers represented to the left and larger numbers to the right. Previously, contralateral neglect-like symptoms in physical line bisection have been induced in healthy subjects with repetitive transcranial magnetic stimulation (rTMS) over right posterior parietal lobe. Here we used rTMS over parietal and occipital sites in healthy subjects to investigate spatial representations in a number bisection task. Subjects were asked to name the midpoint of numerical intervals without calculating. On control trials subjects' behaviour was similar to performance reported in physical line bisection experiments. Subjects underestimated the midpoint of the numerical interval. Repetitive transcranial magnetic stimulation produced representational neglect-like symptoms in number bisection when applied over right posterior parietal cortex (right PPC). Repetitive TMS over right PPC shifted the perceived midpoint of the numerical interval significantly to the right while occipital TMS had no effect on bisection performance. Our study therefore provides further evidence that subjects use spatial representations, perhaps akin to a mental number line, in basic numerical processing tasks. Furthermore, we showed that the right posterior parietal cortex is crucially involved in spatial representation of numbers.

[1]  A. Cowey,et al.  The role of the parietal cortex in visual attention—hemispheric asymmetries and the effects of learning: a magnetic stimulation study , 1998, Neuropsychologia.

[2]  J. Tanji,et al.  Numerical representation for action in the parietal cortex of the monkey , 2002, Nature.

[3]  S. Dehaene,et al.  Topographical Layout of Hand, Eye, Calculation, and Language-Related Areas in the Human Parietal Lobe , 2002, Neuron.

[4]  H. Lüders,et al.  American Electroencephalographic Society Guidelines for Standard Electrode Position Nomenclature , 1991, Journal of clinical neurophysiology : official publication of the American Electroencephalographic Society.

[5]  S. Dehaene,et al.  THREE PARIETAL CIRCUITS FOR NUMBER PROCESSING , 2003, Cognitive neuropsychology.

[6]  J. Marshall,et al.  Spatial cognition: evidence from visual neglect , 2003, Trends in Cognitive Sciences.

[7]  J. Mattingley,et al.  Parietal neglect and visual awareness , 1998, Nature Neuroscience.

[8]  Stanislas Dehaene,et al.  Cerebral Pathways for Calculation: Double Dissociation between Rote Verbal and Quantitative Knowledge of Arithmetic , 1997, Cortex.

[9]  Alvaro Pascual-Leone,et al.  Transcranial magnetic stimulation: a neurochromometrics of mind. , 2003 .

[10]  Z. Nahas,et al.  Motor threshold in transcranial magnetic stimulation: a comparison of a neurophysiological method and a visualization of movement method. , 1998, The journal of ECT.

[11]  Y. Rossetti,et al.  Prism adaptation improves representational neglect , 2001, Neuropsychologia.

[12]  Eliot Hazeltine,et al.  Dissociable Contributions of Prefrontal and Parietal Cortices to Response Selection , 2002, NeuroImage.

[13]  Y. Rossetti,et al.  Ameliorating neglect with prism adaptation: visuo-manual and visuo-verbal measures , 2002, Neuropsychologia.

[14]  J. Knott,et al.  Regarding the American Electroencephalographic Society guidelines for standard electrode position nomenclature: a commentary on the proposal to change the 10-20 electrode designators. , 1993, Journal of clinical neurophysiology : official publication of the American Electroencephalographic Society.

[15]  Klaus Willmes,et al.  On The Impact of Different Number Representations in the Number Bisection Task , 2002, Cortex.

[16]  G. Orban,et al.  Parietal Representation of Symbolic and Nonsymbolic Magnitude , 2003, Journal of Cognitive Neuroscience.

[17]  Jordan Grafman,et al.  Handbook of Neuropsychology , 1991 .

[18]  M. Rushworth,et al.  The left parietal and premotor cortices: motor attention and selection , 2003, NeuroImage.

[19]  Rita Sloan Berndt,et al.  Language and aphasia , 2001 .

[20]  Peter Brugger,et al.  Stimulus-response compatibility in representational space , 1998, Neuropsychologia.

[21]  D. Boisson,et al.  Does Action Make the Link Between Number and Space Representation? , 2004, Psychological science.

[22]  K. Zilles,et al.  The Neural Basis of Vertical and Horizontal Line Bisection Judgments: An fMRI Study of Normal Volunteers , 2001, NeuroImage.

[23]  E. Bisiach,et al.  Timing of right parietal and frontal cortex activity in visuo-spatial perception: a TMS study in normal individuals , 2001, Neuroreport.

[24]  Alan Cowey,et al.  Temporal aspects of visual search studied by transcranial magnetic stimulation , 1997, Neuropsychologia.

[25]  Y. Rossetti,et al.  Sensorimotor effects on central space representation: prism adaptation influences haptic and visual representations in normal subjects , 2004, Neuropsychologia.

[26]  E. Wassermann Risk and safety of repetitive transcranial magnetic stimulation: report and suggested guidelines from the International Workshop on the Safety of Repetitive Transcranial Magnetic Stimulation, June 5-7, 1996. , 1998, Electroencephalography and clinical neurophysiology.

[27]  S. Dehaene,et al.  Understanding dissociations in dyscalculia: a brain imaging study of the impact of number size on the cerebral networks for exact and approximate calculation. , 2000, Brain : a journal of neurology.

[28]  M. Perenin,et al.  Prism adaptation to a rightward optical deviation rehabilitates left hemispatial neglect , 1998, Nature.

[29]  J. Grafman,et al.  The calculating brain: an fMRI study , 2000, Neuropsychologia.

[30]  Marie-Pascale Noël,et al.  Images of numbers, or “when 98 is upper left and 6 sky blue” , 1992, Cognition.

[31]  J. Marshall,et al.  Completion in Visuo-Spatial Neglect: A Case Study , 1994, Cortex.

[32]  S. Dehaene,et al.  Differential Contributions of the Left and Right Inferior Parietal Lobules to Number Processing , 1999, Journal of Cognitive Neuroscience.

[33]  M. Gazzaniga,et al.  The new cognitive neurosciences , 2000 .

[34]  Y. Rossetti,et al.  Reducing spatial neglect by visual and other sensory manipulations: non-cognitive (physiological) routes to the rehabilitation of a cognitive disorder , 2002 .

[35]  J. Marshall,et al.  Completion in Visuo-Spatial Neglect: A Case Study , 1994, Cortex.

[36]  Mark E. McCourt,et al.  Visuospatial attention in line bisection: stimulusmodulation of pseudoneglect , 1999, Neuropsychologia.

[37]  S. Dehaene,et al.  The mental representation of parity and number magnitude. , 1993 .

[38]  Ernst Pöppel,et al.  Compression and lateral shift of mental coordinate systems in a line bisection task , 1988, Neuropsychologia.

[39]  Y. Rossetti,et al.  Simulating unilateral neglect in normals using prism adaptation: implications for theory , 2003, Neuropsychologia.

[40]  K. Priftis,et al.  Brain damage: Neglect disrupts the mental number line , 2002, Nature.

[41]  F. Restle Speed of Adding and Comparing Numbers. , 1970 .

[42]  Alan Cowey,et al.  Spatial neglect in near and far space investigated by repetitive transcranial magnetic stimulation. , 2002, Brain : a journal of neurology.

[43]  M. H. Fischer,et al.  Number processing induces spatial performance biases , 2001, Neurology.

[44]  M HERSHENSON,et al.  Reaction time as a measure of intersensory facilitation. , 1962, Journal of experimental psychology.

[45]  Michael D. Dodd,et al.  Perceiving numbers causes spatial shifts of attention , 2003, Nature Neuroscience.

[46]  M. McCourt,et al.  Pseudoneglect: a review and meta-analysis of performance factors in line bisection tasks , 2000, Neuropsychologia.

[47]  Lisa Cipolotti,et al.  Disturbances of number processing and calculation , 2001 .

[48]  C. Price The functional anatomy of word comprehension and production , 1998, Trends in Cognitive Sciences.

[49]  H. Karnath New insights into the functions of the superior temporal cortex , 2001, Nature Reviews Neuroscience.

[50]  Martin H. Fischer,et al.  Spatial representations in number processing--evidence from a pointing task , 2003 .

[51]  J. Marshall,et al.  The bisection of horizontal and radial lines: a case study of normal controls and ten patients with left visuospatial neglect. , 1993, The International journal of neuroscience.

[52]  Jason B Mattingley,et al.  Effects of prismatic adaptation on judgements of spatial extent in peripersonal and extrapersonal space , 2003, Neuropsychologia.

[53]  Y. Rossetti,et al.  Cognitive bias induced by visuo‐motor adaptation to prisms: a simulation of unilateral neglect in normal individuals? , 2000, Neuroreport.

[54]  Domenico Passafiume,et al.  Calculation Disturbances in Adults with Focal Hemispheric Damage , 1982, Cortex.

[55]  M. Goodale,et al.  The visual brain in action , 1995 .

[56]  A. Kleinschmidt,et al.  Dissociating neural correlates of cognitive components in mental calculation. , 2001, Cerebral cortex.

[57]  Stanislas Dehaene,et al.  Approximate quantities and exact number words: dissociable systems , 2003, Neuropsychologia.

[58]  Filippo Brighina,et al.  Contralateral neglect induced by right posterior parietal rTMS in healthy subjects , 2000, Neuroreport.

[59]  Patrik Vuilleumier,et al.  The Number Space and Neglect , 2004, Cortex.

[60]  E. Bisiach,et al.  Unilateral Neglect of Representational Space , 1978, Cortex.

[61]  John C. Marshall,et al.  How Long is a Piece of String? A Study of Line Bisection in a Case of Visual Neglect , 1988, Cortex.

[62]  Richard S. J. Frackowiak,et al.  The neural correlates of the verbal component of working memory , 1993, Nature.

[63]  Vincent Walsh A theory of magnitude: common cortical metrics of time, space and quantity , 2003, Trends in Cognitive Sciences.

[64]  Y. Rossetti,et al.  After-effects of visuo-manual adaptation to prisms on body posture in normal subjects , 2002, Experimental Brain Research.

[65]  FRANCIS GALTON,et al.  Visualised Numerals , 1880, Nature.

[66]  S. Dehaene,et al.  Interactions between number and space in parietal cortex , 2005, Nature Reviews Neuroscience.

[67]  Carlo Miniussi,et al.  Parietal Lobe Contribution to Mental Rotation Demonstrated with rTMS , 2003, Journal of Cognitive Neuroscience.

[68]  A. Kleinschmidt,et al.  A Supramodal Number Representation in Human Intraparietal Cortex , 2003, Neuron.

[69]  D. Perani,et al.  The anatomy of unilateral neglect after right-hemisphere stroke lesions. A clinical/CT-scan correlation study in man , 1986, Neuropsychologia.

[70]  E. Spelke,et al.  Sources of mathematical thinking: behavioral and brain-imaging evidence. , 1999, Science.

[71]  Philippe Pinel,et al.  Distributed and Overlapping Cerebral Representations of Number, Size, and Luminance during Comparative Judgments , 2004, Neuron.

[72]  Wim Fias,et al.  Oculomotor bias induced by number perception. , 2004, Experimental psychology.

[73]  Stanislas Dehaene,et al.  Two mental calculation systems: A case study of severe acalculia with preserved approximation , 1991, Neuropsychologia.

[74]  K. Zilles,et al.  Line bisection judgments implicate right parietal cortex and cerebellum as assessed by fMRI , 2000, Neurology.

[75]  R T Knight,et al.  Cortical substrates supporting visual search in humans. , 1991, Cerebral cortex.

[76]  Matthew F. S. Rushworth,et al.  The Mental Number Line and the Human Angular Gyrus , 2001, NeuroImage.

[77]  R. Rafal,et al.  "Both" means more than "two": localizing and counting in patients with visuospatial neglect , 1999, Nature Neuroscience.