title: Top-Down Control of Visual Alpha Oscillations: Sources of Control Signals and Their Mechanisms of Action.
abstract: Alpha oscillations (8-12 Hz) are thought to inversely correlate with cortical excitability. Goal-oriented modulation of alpha has been studied extensively. In visual spatial attention, alpha over the region of visual cortex corresponding to the attended location decreases, signifying increased excitability to facilitate the processing of impending stimuli. In contrast, in retention of verbal working memory, alpha over visual cortex increases, signifying decreased excitability to gate out stimulus input to protect the information held online from sensory interference. According to the prevailing model, this goal-oriented biasing of sensory cortex is effected by top-down control signals from frontal and parietal cortices. The present study tests and substantiates this hypothesis by (a) identifying the signals that mediate the top-down biasing influence, (b) examining whether the cortical areas issuing these signals are task-specific or task-independent, and (c) establishing the possible mechanism of the biasing action. High-density human EEG data were recorded in two experimental paradigms: a trial-by-trial cued visual spatial attention task and a modified Sternberg working memory task. Applying Granger causality to both sensor-level and source-level data we report the following findings. In covert visual spatial attention, the regions exerting top-down control over visual activity are lateralized to the right hemisphere, with the dipoles located at the right frontal eye field (FEF) and the right inferior frontal gyrus (IFG) being the main sources of top-down influences. During retention of verbal working memory, the regions exerting top-down control over visual activity are lateralized to the left hemisphere, with the dipoles located at the left middle frontal gyrus (MFG) being the main source of top-down influences. In both experiments, top-down influences are mediated by alpha oscillations, and the biasing effect is likely achieved via an inhibition-disinhibition mechanism.
authors.affiliation: J. Crayton Pruitt Family Department of Biomedical Engineering, University of Florida Gainesville, FL, USA.
authors.forename: Chao
authors.initials: C
authors.lastname: Wang
authors.affiliation: J. Crayton Pruitt Family Department of Biomedical Engineering, University of Florida Gainesville, FL, USA.
authors.forename: Rajasimhan
authors.initials: R
authors.lastname: Rajagovindan
authors.affiliation: J. Crayton Pruitt Family Department of Biomedical Engineering, University of Florida Gainesville, FL, USA.
authors.forename: Sahng-Min
authors.initials: SM
authors.lastname: Han
authors.affiliation: J. Crayton Pruitt Family Department of Biomedical Engineering, University of Florida Gainesville, FL, USA.
authors.forename: Mingzhou
authors.initials: M
authors.lastname: Ding
keywords: EEG; alpha oscillaitons; granger causality; top-down control; visual spatial attention; working memory
references.citation: J Neurosci. 2003 Nov 26;23(34):10904-12
references.pmid: 14645486
references.citation: Neuroimage. 2010 Jun;51(2):867-76
references.pmid: 20188186
references.citation: PLoS Biol. 2014 Oct 21;12(10):e1001965
references.pmid: 25333286
references.citation: Trends Cogn Sci. 2012 Feb;16(2):129-35
references.pmid: 22209601
references.citation: Proc Natl Acad Sci U S A. 2006 Jan 10;103(2):449-54
references.pmid: 16407167
references.citation: J Neurosci. 2011 May 4;31(18):6750-8
references.pmid: 21543604
references.citation: Cereb Cortex. 2010 Apr;20(4):859-72
references.pmid: 19648173
references.citation: J Neurosci. 2015 Jan 28;35(4):1638-47
references.pmid: 25632139
references.citation: Int J Psychophysiol. 2003 Jan;47(1):65-74
references.pmid: 12543447
references.citation: J Cogn Neurosci. 2009 Jun;21(6):1146-61
references.pmid: 18752395
references.citation: Psychophysiology. 2000 Mar;37(2):163-78
references.pmid: 10731767
references.citation: Semin Neurol. 2000;20(4):463-70
references.pmid: 11149702
references.citation: Cereb Cortex. 2007 Feb;17(2):391-9
references.pmid: 16525126
references.citation: Eur J Neurosci. 2015 Apr;41(7):914-24
references.pmid: 25704649
references.citation: J Neurosci Methods. 2010 Feb 15;186(2):262-73
references.pmid: 19961876
references.citation: Hum Brain Mapp. 2008 May;29(5):603-12
references.pmid: 17557296
references.citation: Clin Neurophysiol. 2002 Aug;113(8):1279-85
references.pmid: 12140008
references.citation: Front Hum Neurosci. 2010 Nov 04;4:186
references.pmid: 21119777
references.citation: J Comput Neurosci. 2000 Nov-Dec;9(3):271-91
references.pmid: 11139043
references.citation: Nat Neurosci. 2011 May;14(5):656-61
references.pmid: 21441920
references.citation: J Cogn Neurosci. 1997 Jul;9(4):462-75
references.pmid: 23968211
references.citation: J Cogn Neurosci. 2003 Oct 1;15(7):948-60
references.pmid: 14614806
references.citation: Front Psychol. 2011 May 19;2:99
references.pmid: 21779257
references.citation: Nat Neurosci. 2000 Mar;3(3):284-91
references.pmid: 10700262
references.citation: PLoS One. 2010 Apr 22;5(4):e10298
references.pmid: 20421978
references.citation: Neuroimage. 2008 Jun;41(2):354-62
references.pmid: 18394927
references.citation: Neuroscience. 2011 Apr 28;180:165-80
references.pmid: 21310216
references.citation: Trends Cogn Sci. 2005 Oct;9(10):474-80
references.pmid: 16150631
references.citation: Science. 2012 Aug 10;337(6095):753-6
references.pmid: 22879517
references.citation: J Cogn Neurosci. 2000 Jan;12(1):174-87
references.pmid: 10769314
references.citation: Brain Res Cogn Brain Res. 1998 Jul;7(1):1-13
references.pmid: 9714705
references.citation: Trends Neurosci. 2014 Jul;37(7):357-69
references.pmid: 24836381
references.citation: Brain Res Rev. 2007 Jan;53(1):63-88
references.pmid: 16887192
references.citation: J Neurosci. 1993 Mar;13(3):1202-26
references.pmid: 8441008
references.citation: Neuron. 1999 Apr;22(4):751-61
references.pmid: 10230795
references.citation: Electroencephalogr Clin Neurophysiol. 1997 Dec;103(6):642-51
references.pmid: 9546492
references.citation: J Cogn Neurosci. 2014 Feb;26(2):422-32
references.pmid: 24116843
references.citation: Ann N Y Acad Sci. 2013 Aug;1296:88-107
references.pmid: 23909769
references.citation: Nat Rev Neurosci. 2002 Mar;3(3):201-15
references.pmid: 11994752
references.citation: Cereb Cortex. 2002 Aug;12(8):877-82
references.pmid: 12122036
references.citation: Cereb Cortex. 2016 Feb;26(2):517-29
references.pmid: 25205663
references.citation: Psychiatry Res. 2001 Oct 1;107(3):151-63
references.pmid: 11566431
references.citation: Neuroimage. 2008 Jan 1;39(1):492-9
references.pmid: 17936643
references.citation: J Neurosci. 2014 Nov 26;34(48):16117-25
references.pmid: 25429152
references.citation: Biol Cybern. 2000 Jul;83(1):35-45
references.pmid: 10933236
references.citation: Brain Res. 2007 Jun 4;1152:171-81
references.pmid: 17459348
references.citation: J Neurosci. 2012 Oct 10;32(41):14305-10
references.pmid: 23055501
references.citation: Nature. 2003 Jan 23;421(6921):370-3
references.pmid: 12540901
references.citation: Neuroimage. 2004 Mar;21(3):894-903
references.pmid: 15006656
references.citation: Clin Neurophysiol. 2011 Nov;122(11):2324-5
references.pmid: 21543253
references.citation: Neuron. 1998 Oct;21(4):761-73
references.pmid: 9808463
references.citation: Psychiatry Res. 1996 Oct 7;67(3):173-87
references.pmid: 8912957
references.citation: Brain Topogr. 1992 Winter;5(2):103-11
references.pmid: 1489638
references.citation: Curr Biol. 2006 Aug 8;16(15):1479-88
references.pmid: 16890523
references.citation: Science. 2005 Apr 29;308(5722):702-4
references.pmid: 15860630
references.citation: Proc Natl Acad Sci U S A. 2004 Jun 29;101(26):9849-54
references.pmid: 15210971
references.citation: Neuroreport. 2008 Jan 22;19(2):203-8
references.pmid: 18185109
references.citation: Neuron. 2012 Nov 21;76(4):838-46
references.pmid: 23177967
references.citation: J Cogn Neurosci. 2004 Jan-Feb;16(1):149-65
references.pmid: 15006044
references.citation: Nat Neurosci. 2001 Sep;4(9):953-7
references.pmid: 11528429
references.citation: Clin Neurophysiol. 2001 Sep;112(9):1672-5
references.pmid: 11514250
references.citation: Arch Neurol. 1987 Jun;44(6):621-5
references.pmid: 3579679
references.citation: Electroencephalogr Clin Neurophysiol. 1997 Nov;103(5):499-515
references.pmid: 9402881
references.citation: J Neurosci. 2008 Oct 1;28(40):9976-88
references.pmid: 18829955
references.citation: J Neurosci. 2009 May 6;29(18):5863-72
references.pmid: 19420253
references.citation: Neuroimage. 2014 Sep;98:306-13
references.pmid: 24807400
references.citation: Trends Cogn Sci. 2012 Dec;16(12):606-17
references.pmid: 23141428
references.citation: Eur J Neurosci. 2005 Dec;22(11):2917-26
references.pmid: 16324126
references.citation: Neuroimage. 2010 Apr 15;50(3):1292-302
references.pmid: 20035885
references.citation: Hum Brain Mapp. 2009 Feb;30(2):532-40
references.pmid: 18095279
references.citation: Annu Rev Neurosci. 2000;23:315-41
references.pmid: 10845067
references.citation: J Neurosci. 2008 Oct 1;28(40):10056-61
references.pmid: 18829963
references.citation: Ann Neurol. 1981 Oct;10(4):309-25
references.pmid: 7032417
references.citation: Proc Natl Acad Sci U S A. 1998 Jul 7;95(14):8410-3
references.pmid: 9653200
references.citation: Int J Psychophysiol. 2005 Aug;57(2):97-103
references.pmid: 15967528
references.citation: J Neurosci. 2006 Sep 13;26(37):9494-502
references.pmid: 16971533
references.citation: Neuron. 2006 Jun 1;50(5):791-8
references.pmid: 16731516
references.citation: Neuroimage. 2014 Oct 15;100:125-34
references.pmid: 24945669
references.citation: Neuroimage. 2010 Apr 1;50(2):577-88
references.pmid: 20026279
references.citation: J Neurosci. 2000 Mar 15;20(6):RC63
references.pmid: 10704517
references.citation: Cortex. 2003 Sep-Dec;39(4-5):897-911
references.pmid: 14584558
references.citation: Nat Rev Neurosci. 2001 Oct;2(10):704-16
references.pmid: 11584308
references.citation: Cereb Cortex. 2005 May;15(5):670-8
references.pmid: 15342429
references.citation: Brain Res Cogn Brain Res. 1999 Mar;7(4):493-501
references.pmid: 10076094
references.citation: Brain Res Brain Res Rev. 1999 Apr;29(2-3):169-95
references.pmid: 10209231
references.citation: Neuroimage. 1999 Dec;10(6):695-704
references.pmid: 10600415
references.citation: Neuroimage. 2003 Jul;19(3):797-809
references.pmid: 12880808
references.citation: J Neurosci. 2012 May 16;32(20):7034-41
references.pmid: 22593071
references.citation: Electroencephalogr Clin Neurophysiol. 1994 Mar;90(3):229-41
references.pmid: 7511504
references.citation: Neuroimage. 2003 Feb;18(2):247-56
references.pmid: 12595179
references.citation: J Clin Neurophysiol. 2002 Apr;19(2):91-112
references.pmid: 11997721
references.citation: Electroencephalogr Clin Neurophysiol. 1989 Feb;72(2):184-7
references.pmid: 2464490
references.citation: Brain Topogr. 2004 Summer;16(4):233-8
references.pmid: 15379219
references.citation: J Cogn Neurosci. 2011 Jun;23(6):1379-94
references.pmid: 20459310
references.citation: Neuroimage. 2014 May 1;91:300-10
references.pmid: 24434679
references.citation: J Neurosci. 2008 Mar 5;28(10):2667-79
references.pmid: 18322110
references.citation: Front Psychol. 2011 Oct 05;2:241
references.pmid: 22007179
references.citation: Eur Psychiatry. 2008 Jun;23(4):281-8
references.pmid: 18455371
references.citation: Proc Natl Acad Sci U S A. 2006 Dec 19;103(51):19530-4
references.pmid: 17151200
references.citation: J Neurosci. 2003 May 15;23 (10 ):3990-8
references.pmid: 12764083
references.citation: Neuroimage. 2014 Feb 15;87:242-51
references.pmid: 24185020
references.citation: Front Psychol. 2011 Sep 05;2:204
references.pmid: 21922012
references.citation: Biol Cybern. 2001 Aug;85(2):145-57
references.pmid: 11508777
references.citation: Science. 1966 Aug 5;153(3736):652-4
references.pmid: 5939936
references.citation: Neurology. 1980 Mar;30(3):327-30
references.pmid: 7189037
references.citation: Hum Brain Mapp. 2007 Aug;28(8):785-92
references.pmid: 17266103
references.citation: BMC Neurosci. 2011 Jan 06;12:2
references.pmid: 21211016
references.citation: Cereb Cortex. 1996 Jan-Feb;6(1):11-20
references.pmid: 8670634
references.citation: Neuroimage. 2011 Sep 15;58(2):339-61
references.pmid: 21477655
references.citation: Nat Neurosci. 2000 Apr;3(4):399-403
references.pmid: 10725931
references.citation: Neuroimage. 2004 Jan;21(1):340-51
references.pmid: 14741672
references.citation: Neuroimage. 2008 Aug 15;42(2):928-35
references.pmid: 18595738
references.citation: J Neurosci Methods. 2007 Oct 15;166(1):41-52
references.pmid: 17698205
references.citation: Brain Res. 2009 Dec 15;1303:97-110
references.pmid: 19782056
references.citation: Nat Neurosci. 2008 Jan;11(1):103-7
references.pmid: 18066057
references.citation: Hum Brain Mapp. 2009 Oct;30(10):3417-25
references.pmid: 19384888
references.citation: Science. 2009 May 29;324(5931):1207-10
references.pmid: 19478185
references.citation: Acta Psychol (Amst). 1999 Apr;101(2-3):159-78
references.pmid: 10344184