Attentional modulation of SSVEP power depends on the network tagged by the flicker frequency.

Modulation of the steady-state visual evoked potential (SSVEP) by attention was studied in detail using 15 'tag' frequencies in the range of 2.5-20 Hz. The stimuli were two series of random disc search arrays superimposed on two concentric color-marked annuli respectively. Two series of arrays were updated independently; one updated at one fixed frequency (flicker) and the other updated randomly according to a white noise distribution (random broadband flicker, rbbf). On each trial, the observer was instructed to attend one annulus and to detect a target (a triangle) that occasionally appeared in a random disc array in the attended annulus. The SSVEP results show that the choice of flicker frequency selects which cortical network synchronizes to the flicker two distinct cortical networks showed different effects of attention. SSVEP power and the effects of attention on SSVEP power strongly depend on both flicker frequency and radial position of rbbf annulus. At flicker frequencies in the delta band (2-4 Hz), and in the upper alpha band (10-11 Hz), an occipital-frontal network appears to phase-lock to the flicker when attending to the flicker, increasing the magnitude of the SSVEP. At flicker frequencies in the lower alpha band (8-10 Hz), a global response to a peripheral flickering stimulus, that includes parietal cortex and posterior frontal cortex, has higher amplitude when attention is directed away from the flickering peripheral stimulus and towards a competing rbbf stimulus in the fovea. Increases in SSVEP power when attention is directed to peripheral flicker are always associated with increases in phase locking. By contrast, at frequencies in the lower alpha band, increases in SSVEP power when attention is directed away from the flicker and towards foveal stimuli are not associated with changes in phase-locking. Thus, whether attention to a flicker stimulus increases or decreases SSVEP amplitude and phase locking depends on which of two cortical networks, which have distinct spatial and dynamic properties, is selected by the flicker frequency.

[1]  J. Bendat,et al.  Random Data: Analysis and Measurement Procedures , 1971 .

[2]  H. Saunders Literature Review : RANDOM DATA: ANALYSIS AND MEASUREMENT PROCEDURES J. S. Bendat and A.G. Piersol Wiley-Interscience, New York, N. Y. (1971) , 1974 .

[3]  D. Regan Steady-state evoked potentials. , 1977, Journal of the Optical Society of America.

[4]  M. Posner,et al.  Attention and the detection of signals. , 1980, Journal of experimental psychology.

[5]  J. Fermaglich Electric Fields of the Brain: The Neurophysics of EEG , 1982 .

[6]  K. Nakayama,et al.  Steady state visual evoked potentials in the alert primate , 1982, Vision Research.

[7]  C. Eriksen,et al.  Visual attention within and around the field of focal attention: A zoom lens model , 1986, Perception & psychophysics.

[8]  S Makeig,et al.  Auditory steady-state responses: threshold prediction using phase coherence. , 1987, Electroencephalography and clinical neurophysiology.

[9]  S A Hillyard,et al.  Spatial gradients of visual attention: behavioral and electrophysiological evidence. , 1988, Electroencephalography and clinical neurophysiology.

[10]  D. Regan,et al.  Human brain electrophysiology , 1989 .

[11]  S J Luck,et al.  Visual event-related potentials index focused attention within bilateral stimulus arrays. I. Evidence for early selection. , 1990, Electroencephalography and clinical neurophysiology.

[12]  M. Brandt,et al.  The effect of the phase of prestimulus alpha activity on the averaged visual evoked response. , 1991, Electroencephalography and clinical neurophysiology.

[13]  Robert Tibshirani,et al.  An Introduction to the Bootstrap , 1994 .

[14]  D. Tucker Spatial sampling of head electrical fields: the geodesic sensor net. , 1993, Electroencephalography and clinical neurophysiology.

[15]  S. Kobayashi,et al.  Electroencephalographic activity associated with shifts of visuospatial attention. , 1994, Brain : a journal of neurology.

[16]  P. Nunez,et al.  Neocortical Dynamics and Human EEG Rhythms , 1995 .

[17]  S. Hillyard,et al.  Spatial Selective Attention Affects Early Extrastriate But Not Striate Components of the Visual Evoked Potential , 1996, Journal of Cognitive Neuroscience.

[18]  S. Hillyard,et al.  Selective attention to stimulus location modulates the steady-state visual evoked potential. , 1996, Proceedings of the National Academy of Sciences of the United States of America.

[19]  T. Collura Neocortical Dynamics and Human EEG Rhythms , 1996 .

[20]  D H Brainard,et al.  The Psychophysics Toolbox. , 1997, Spatial vision.

[21]  R. Desimone,et al.  Neural mechanisms of spatial selective attention in areas V1, V2, and V4 of macaque visual cortex. , 1997, Journal of neurophysiology.

[22]  D G Pelli,et al.  The VideoToolbox software for visual psychophysics: transforming numbers into movies. , 1997, Spatial vision.

[23]  W. Singer,et al.  The response of cat visual cortex to flicker stimuli of variable frequency , 1998, The European journal of neuroscience.

[24]  Matthias M. Müller,et al.  The time course of cortical facilitation during cued shifts of spatial attention , 1998, Nature Neuroscience.

[25]  D. P. Russell,et al.  Investigating neural correlates of conscious perception by frequency-tagged neuromagnetic responses. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[26]  P. Nunez,et al.  Spatial filtering and neocortical dynamics: estimates of EEG coherence , 1998, IEEE Transactions on Biomedical Engineering.

[27]  Matthias M. Müller,et al.  Effects of spatial selective attention on the steady-state visual evoked potential in the 20-28 Hz range. , 1998, Brain research. Cognitive brain research.

[28]  S. Hillyard,et al.  Event-related brain potentials in the study of visual selective attention. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[29]  D. P. Russell,et al.  Increased Synchronization of Neuromagnetic Responses during Conscious Perception , 1999, The Journal of Neuroscience.

[30]  G. Woodman,et al.  Event-related potential studies of attention , 2000, Trends in Cognitive Sciences.

[31]  Richard B. Silberstein,et al.  Steady state visually evoked potential, brain resonances and cognitive processes , 2000 .

[32]  G. V. Simpson,et al.  Anticipatory Biasing of Visuospatial Attention Indexed by Retinotopically Specific α-Bank Electroencephalography Increases over Occipital Cortex , 2000, The Journal of Neuroscience.

[33]  J. Bendat,et al.  Random Data: Analysis and Measurement Procedures , 1987 .

[34]  P. Nunez,et al.  Steady state visually evoked potential (SSVEP) topography in a graded working memory task. , 2001, International journal of psychophysiology : official journal of the International Organization of Psychophysiology.

[35]  R. Desimone,et al.  Modulation of Oscillatory Neuronal Synchronization by Selective Visual Attention , 2001, Science.

[36]  Francesca Pei,et al.  Neural correlates of object-based attention. , 2002, Journal of vision.

[37]  W. Klimesch,et al.  Frequency characteristics of evoked and oscillatory electroencephalic activity in a human memory scanning task , 2002, Neuroscience Letters.

[38]  S. A. Hillyard,et al.  Sustained division of the attentional spotlight , 2003, Nature.

[39]  Anil K. Seth,et al.  The power of human brain magnetoencephalographic signals can be modulated up or down by changes in an attentive visual task , 2003, Proceedings of the National Academy of Sciences of the United States of America.

[40]  D. Somers,et al.  Multiple Spotlights of Attentional Selection in Human Visual Cortex , 2004, Neuron.

[41]  P. Apkarian,et al.  Multiple spatial-frequency tuning of electrical responses from human visual cortex , 1978, Experimental Brain Research.

[42]  J. Reynolds,et al.  Attentional modulation of visual processing. , 2004, Annual review of neuroscience.

[43]  Ramesh Srinivasan,et al.  Internal and External Neural Synchronization during Conscious Perception , 2004, Int. J. Bifurc. Chaos.

[44]  W. Klimesch,et al.  Alpha phase synchronization predicts P1 and N1 latency and amplitude size. , 2005, Cerebral cortex.

[45]  Sabine Weiss,et al.  Quantification of phase synchronization phenomena and their importance for verbal memory processes , 2005, Biological Cybernetics.