The Global Signal and Observed Anticorrelated Resting State Brain Networks

1 Departments of Radiology, 2 Neurology, 3 Anatomy and Neurobiology, and 4 Biomedical Engineering, Washington University in St. Louis, St. Louis, Missouri Submitted 17 July 2008; accepted in final form 23 March 2009 Resting state studies of spontaneous fluctuations in the functional MRI (fMRI) blood...

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Published in	Journal of neurophysiology Vol. 101; no. 6; pp. 3270 - 3283
Main Authors	Fox, Michael D, Zhang, Dongyang, Snyder, Abraham Z, Raichle, Marcus E
Format	Journal Article
Language	English
Published	United States Am Phys Soc 01.06.2009 American Physiological Society
Subjects	Brain - blood supply Brain - physiology Brain Mapping Humans Image Processing, Computer-Assisted - methods Magnetic Resonance Imaging - methods Models, Neurological Movement - physiology Nerve Net - blood supply Nerve Net - physiology Neural Pathways - blood supply Neural Pathways - physiology Oxygen - blood Perceptual Masking - physiology Regression Analysis Rest - physiology Statistics as Topic
Online Access	Get full text
ISSN	0022-3077 1522-1598
DOI	10.1152/jn.90777.2008

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Abstract	1 Departments of Radiology, 2 Neurology, 3 Anatomy and Neurobiology, and 4 Biomedical Engineering, Washington University in St. Louis, St. Louis, Missouri Submitted 17 July 2008; accepted in final form 23 March 2009 Resting state studies of spontaneous fluctuations in the functional MRI (fMRI) blood oxygen level dependent (BOLD) signal have shown great promise in mapping the brain's intrinsic, large-scale functional architecture. An important data preprocessing step used to enhance the quality of these observations has been removal of spontaneous BOLD fluctuations common to the whole brain (the so-called global signal). One reproducible consequence of global signal removal has been the finding that spontaneous BOLD fluctuations in the default mode network and an extended dorsal attention system are consistently anticorrelated, a relationship that these two systems exhibit during task performance. The dependence of these resting-state anticorrelations on global signal removal has raised important questions regarding the nature of the global signal, the validity of global signal removal, and the appropriate interpretation of observed anticorrelated brain networks. In this study, we investigate several properties of the global signal and find that it is, indeed, global, not residing preferentially in systems exhibiting anticorrelations. We detail the influence of global signal removal on resting state correlation maps both mathematically and empirically, showing an enhancement in detection of system-specific correlations and improvement in the correspondence between resting-state correlations and anatomy. Finally, we show that several characteristics of anticorrelated networks including their spatial distribution, cross-subject consistency, presence with modified whole brain masks, and existence before global regression are not attributable to global signal removal and therefore suggest a biological basis. Address for reprint requests and other correspondence: (E-mail: foxmdphd{at}gmail.com )
AbstractList	Resting state studies of spontaneous fluctuations in the functional MRI (fMRI) blood oxygen level dependent (BOLD) signal have shown great promise in mapping the brain's intrinsic, large-scale functional architecture. An important data preprocessing step used to enhance the quality of these observations has been removal of spontaneous BOLD fluctuations common to the whole brain (the so-called global signal). One reproducible consequence of global signal removal has been the finding that spontaneous BOLD fluctuations in the default mode network and an extended dorsal attention system are consistently anticorrelated, a relationship that these two systems exhibit during task performance. The dependence of these resting-state anticorrelations on global signal removal has raised important questions regarding the nature of the global signal, the validity of global signal removal, and the appropriate interpretation of observed anticorrelated brain networks. In this study, we investigate several properties of the global signal and find that it is, indeed, global, not residing preferentially in systems exhibiting anticorrelations. We detail the influence of global signal removal on resting state correlation maps both mathematically and empirically, showing an enhancement in detection of system-specific correlations and improvement in the correspondence between resting-state correlations and anatomy. Finally, we show that several characteristics of anticorrelated networks including their spatial distribution, cross-subject consistency, presence with modified whole brain masks, and existence before global regression are not attributable to global signal removal and therefore suggest a biological basis. Resting state studies of spontaneous fluctuations in the functional MRI (fMRI) blood oxygen level dependent (BOLD) signal have shown great promise in mapping the brain's intrinsic, large-scale functional architecture. An important data preprocessing step used to enhance the quality of these observations has been removal of spontaneous BOLD fluctuations common to the whole brain (the so-called global signal). One reproducible consequence of global signal removal has been the finding that spontaneous BOLD fluctuations in the default mode network and an extended dorsal attention system are consistently anticorrelated, a relationship that these two systems exhibit during task performance. The dependence of these resting-state anticorrelations on global signal removal has raised important questions regarding the nature of the global signal, the validity of global signal removal, and the appropriate interpretation of observed anticorrelated brain networks. In this study, we investigate several properties of the global signal and find that it is, indeed, global, not residing preferentially in systems exhibiting anticorrelations. We detail the influence of global signal removal on resting state correlation maps both mathematically and empirically, showing an enhancement in detection of system-specific correlations and improvement in the correspondence between resting-state correlations and anatomy. Finally, we show that several characteristics of anticorrelated networks including their spatial distribution, cross-subject consistency, presence with modified whole brain masks, and existence before global regression are not attributable to global signal removal and therefore suggest a biological basis.Resting state studies of spontaneous fluctuations in the functional MRI (fMRI) blood oxygen level dependent (BOLD) signal have shown great promise in mapping the brain's intrinsic, large-scale functional architecture. An important data preprocessing step used to enhance the quality of these observations has been removal of spontaneous BOLD fluctuations common to the whole brain (the so-called global signal). One reproducible consequence of global signal removal has been the finding that spontaneous BOLD fluctuations in the default mode network and an extended dorsal attention system are consistently anticorrelated, a relationship that these two systems exhibit during task performance. The dependence of these resting-state anticorrelations on global signal removal has raised important questions regarding the nature of the global signal, the validity of global signal removal, and the appropriate interpretation of observed anticorrelated brain networks. In this study, we investigate several properties of the global signal and find that it is, indeed, global, not residing preferentially in systems exhibiting anticorrelations. We detail the influence of global signal removal on resting state correlation maps both mathematically and empirically, showing an enhancement in detection of system-specific correlations and improvement in the correspondence between resting-state correlations and anatomy. Finally, we show that several characteristics of anticorrelated networks including their spatial distribution, cross-subject consistency, presence with modified whole brain masks, and existence before global regression are not attributable to global signal removal and therefore suggest a biological basis. 1 Departments of Radiology, 2 Neurology, 3 Anatomy and Neurobiology, and 4 Biomedical Engineering, Washington University in St. Louis, St. Louis, Missouri Submitted 17 July 2008; accepted in final form 23 March 2009 Resting state studies of spontaneous fluctuations in the functional MRI (fMRI) blood oxygen level dependent (BOLD) signal have shown great promise in mapping the brain's intrinsic, large-scale functional architecture. An important data preprocessing step used to enhance the quality of these observations has been removal of spontaneous BOLD fluctuations common to the whole brain (the so-called global signal). One reproducible consequence of global signal removal has been the finding that spontaneous BOLD fluctuations in the default mode network and an extended dorsal attention system are consistently anticorrelated, a relationship that these two systems exhibit during task performance. The dependence of these resting-state anticorrelations on global signal removal has raised important questions regarding the nature of the global signal, the validity of global signal removal, and the appropriate interpretation of observed anticorrelated brain networks. In this study, we investigate several properties of the global signal and find that it is, indeed, global, not residing preferentially in systems exhibiting anticorrelations. We detail the influence of global signal removal on resting state correlation maps both mathematically and empirically, showing an enhancement in detection of system-specific correlations and improvement in the correspondence between resting-state correlations and anatomy. Finally, we show that several characteristics of anticorrelated networks including their spatial distribution, cross-subject consistency, presence with modified whole brain masks, and existence before global regression are not attributable to global signal removal and therefore suggest a biological basis. Address for reprint requests and other correspondence: (E-mail: foxmdphd{at}gmail.com )
Author	Zhang, Dongyang Raichle, Marcus E Snyder, Abraham Z Fox, Michael D
AuthorAffiliation	1 Departments of Radiology, 2 Neurology, 3 Anatomy and Neurobiology, and 4 Biomedical Engineering, Washington University in St. Louis, St. Louis, Missouri
AuthorAffiliation_xml	– name: 1 Departments of Radiology, 2 Neurology, 3 Anatomy and Neurobiology, and 4 Biomedical Engineering, Washington University in St. Louis, St. Louis, Missouri
Author_xml	– sequence: 1 fullname: Fox, Michael D – sequence: 2 fullname: Zhang, Dongyang – sequence: 3 fullname: Snyder, Abraham Z – sequence: 4 fullname: Raichle, Marcus E
BackLink	https://www.ncbi.nlm.nih.gov/pubmed/19339462$$D View this record in MEDLINE/PubMed
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Notes	ObjectType-Article-1 SourceType-Scholarly Journals-1 ObjectType-Feature-2 content type line 23 M. D. Fox and D. Zhang contributed equally to this study.
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Snippet	1 Departments of Radiology, 2 Neurology, 3 Anatomy and Neurobiology, and 4 Biomedical Engineering, Washington University in St. Louis, St. Louis, Missouri... Resting state studies of spontaneous fluctuations in the functional MRI (fMRI) blood oxygen level dependent (BOLD) signal have shown great promise in mapping...
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SubjectTerms	Brain - blood supply Brain - physiology Brain Mapping Humans Image Processing, Computer-Assisted - methods Magnetic Resonance Imaging - methods Models, Neurological Movement - physiology Nerve Net - blood supply Nerve Net - physiology Neural Pathways - blood supply Neural Pathways - physiology Oxygen - blood Perceptual Masking - physiology Regression Analysis Rest - physiology Statistics as Topic
Title	The Global Signal and Observed Anticorrelated Resting State Brain Networks
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