The fuzzy brain. Vagueness and mapping connectivity of the human cerebral cortex

While the past century of neuroscientific research has brought considerable progress in defining the boundaries of the human cerebral cortex, there are cases in which the demarcation of one area from another remains fuzzy. Despite the existence of clearly demarcated areas, examples of gradual transitions between areas are known since early cytoarchitectonic studies. Since multi-modal anatomical approaches and functional connectivity studies brought renewed attention to the topic, a better understanding of the theoretical and methodological implications of fuzzy boundaries in brain science can be conceptually useful. This article provides a preliminary conceptual framework to understand this problem by applying philosophical theories of vagueness to three levels of neuroanatomical research. For the first two levels (cytoarchitectonics and fMRI studies), vagueness will be distinguished from other forms of uncertainty, such as imprecise measurement or ambiguous causal sources of activation. The article proceeds to discuss the implications of these levels for the anatomical study of connectivity between cortical areas. There, vagueness gets imported into connectivity studies since the network structure is dependent on the parcellation scheme and thresholds have to be used to delineate functional boundaries. Functional connectivity may introduce an additional form of vagueness, as it is an organizational principle of the brain. The article concludes by discussing what steps are appropriate to define areal boundaries more precisely.

[1]  Karl J. Friston Functional and effective connectivity in neuroimaging: A synthesis , 1994 .

[2]  Jonathan A C Sterne,et al.  Sifting the evidence—what's wrong with significance tests? , 2001, BMJ : British Medical Journal.

[3]  E. J. Tehovnik,et al.  Direct and indirect activation of cortical neurons by electrical microstimulation. , 2006, Journal of neurophysiology.

[4]  R. Faull,et al.  Distribution of excitatory and inhibitory amino acid, sigma, monoamine, catecholamine, acetylcholine, opioid, neurotensin, substance P, adenosine and neuropeptide Y receptors in human motor and somatosensory cortex , 1991, Brain Research.

[5]  G. Gigerenzer Mindless statistics , 2004 .

[6]  P. Rakic Specification of cerebral cortical areas. , 1988, Science.

[7]  G. Smith,et al.  Die Cytoarchitektonik der Hirnrinde des erwachsenen Menschen. , 1927 .

[8]  Smith Ge,et al.  A New Topographical Survey of the Human Cerebral Cortex, being an Account of the Distribution of the Anatomically Distinct Cortical Areas and their Relationship to the Cerebral Sulci. , 1907 .

[9]  G. Frege Grundgesetze der Arithmetik , 1893 .

[10]  Yul-Wan Sung,et al.  Functional magnetic resonance imaging , 2004, Scholarpedia.

[11]  Davis Lewis,et al.  INDEX, CONTEXT, AND CONTENT , 1980 .

[12]  Ioannis A. Kakadiaris,et al.  Localization and Segmentation of Left Ventricle in Cardiac Cine-MR Images , 2009, IEEE Transactions on Biomedical Engineering.

[13]  M. Raichle,et al.  Anatomic Localization and Quantitative Analysis of Gradient Refocused Echo-Planar fMRI Susceptibility Artifacts , 1997, NeuroImage.

[14]  A. Fingelkurts,et al.  Functional connectivity in the brain—is it an elusive concept? , 2005, Neuroscience & Biobehavioral Reviews.

[15]  D. Pandya,et al.  Dorsolateral prefrontal cortex: comparative cytoarchitectonic analysis in the human and the macaque brain and corticocortical connection patterns , 1999, The European journal of neuroscience.

[16]  Rudolf Seising,et al.  From vagueness in medical thought to the foundations of fuzzy reasoning in medical diagnosis , 2006, Artif. Intell. Medicine.

[17]  Marisa O. Hollinshead,et al.  The organization of the human cerebral cortex estimated by intrinsic functional connectivity. , 2011, Journal of neurophysiology.

[18]  A. Schleicher,et al.  Mapping of human and macaque sensorimotor areas by integrating architectonic, transmitter receptor, MRI and PET data. , 1995, Journal of anatomy.

[19]  D. J. Felleman,et al.  Distributed hierarchical processing in the primate cerebral cortex. , 1991, Cerebral cortex.

[20]  F. Inglis How To Do Things With Words. , 1971 .

[21]  Damien A. Fair,et al.  Defining functional areas in individual human brains using resting functional connectivity MRI , 2008, NeuroImage.

[22]  Joseph T Rouse,et al.  Articulating the World: Experimental Systems and Conceptual Understanding , 2011 .

[23]  Peter A. Bandettini,et al.  Functional MRI: background, methodology, limits, and implementation , 2000 .

[24]  Hiroshi Fukuda,et al.  The Overlapping Community Structure of Structural Brain Network in Young Healthy Individuals , 2011, PloS one.

[25]  Jelliffe Vergleichende Lokalisationslehre der Grosshirnrinde , 1910 .

[26]  Karl J. Friston,et al.  Structural Covariance in the Human Cortex , 2005, The Journal of Neuroscience.

[27]  Katrin Amunts,et al.  Cortical Folding Patterns and Predicting Cytoarchitecture , 2007, Cerebral cortex.

[28]  Achille C. Varzi Vagueness, Logic, and Ontology , 2001 .

[29]  Achille C. Varzi Vagueness in geography , 2001 .

[30]  Peter Unger,et al.  The Problem of the Many , 1980 .

[31]  Wen-June Wang,et al.  Automated classification of multi-spectral MR images using Linear Discriminant Analysis , 2010, Comput. Medical Imaging Graph..

[32]  Barry Horwitz,et al.  The elusive concept of brain connectivity , 2003, NeuroImage.

[33]  Katrin Amunts,et al.  3 Architectonic Mapping of the Human Cerebral Cortex , 2002 .

[34]  Y Matsuda,et al.  Arrangement of orientation pinwheel centers around area 17/18 transition zone in cat visual cortex. , 2000, Cerebral cortex.

[35]  L. Laudan A Confutation of Convergent Realism , 1981, Philosophy of Science.

[36]  M. Young,et al.  Computational analysis of functional connectivity between areas of primate cerebral cortex. , 2000, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[37]  Bart Kosko,et al.  Neural networks and fuzzy systems: a dynamical systems approach to machine intelligence , 1991 .

[38]  Simon B. Eickhoff,et al.  Analysis of neural mechanisms underlying verbal fluency in cytoarchitectonically defined stereotaxic space—The roles of Brodmann areas 44 and 45 , 2004, NeuroImage.

[39]  P S Goldman-Rakic,et al.  Cytoarchitectonic definition of prefrontal areas in the normal human cortex: I. Remapping of areas 9 and 46 using quantitative criteria. , 1995, Cerebral cortex.

[40]  Qiao Zhang,et al.  Fuzziness - vagueness - generality - ambiguity , 1998 .

[41]  Krzysztof J. Gorgolewski,et al.  Pitfalls of Thresholding Statistical Maps in Presurgical fMRI Mapping , 2011 .

[42]  John F. Canny,et al.  A Computational Approach to Edge Detection , 1986, IEEE Transactions on Pattern Analysis and Machine Intelligence.

[43]  Geert Keil HALBGLATZEN STATT HALBWAHRHEITEN. ÜBER VAGHEIT, WAHRHEITS- UND AUFLÖSUNGSGRADE , 2010 .

[44]  Nancy Kanwisher,et al.  Divide and conquer: A defense of functional localizers , 2006, NeuroImage.

[45]  Markus Schneider,et al.  Vague Regions , 1997, SSD.

[46]  H. Barbas,et al.  Cortical structure predicts the pattern of corticocortical connections. , 1997, Cerebral cortex.

[47]  G. Edelman,et al.  A measure for brain complexity: relating functional segregation and integration in the nervous system. , 1994, Proceedings of the National Academy of Sciences of the United States of America.

[48]  Heidi Johansen-Berg,et al.  Tractography: Where Do We Go from Here? , 2011, Brain Connect..

[49]  Jonas Åkerman,et al.  Vagueness and Non-Indexical Contextualism , 2010 .

[50]  Yutaka Hata,et al.  Automated Gyrus Labeling Using Knowledge-based Fuzzy Inference Systems , 2007, 2007 IEEE International Conference on System of Systems Engineering.

[51]  P. Hoyningen-Huene The Ultimate Argument Against Convergent Realism and Structural Realism: The Impasse Objection , 2013 .

[52]  C. Craver Explaining the Brain: Mechanisms and the Mosaic Unity of Neuroscience , 2007 .

[53]  A. Schleicher,et al.  Cyto- and Myeloarchitecture of Human Visual Cortex and the Periodical GABAA Receptor Distribution , 1993 .

[54]  John MacFarlane,et al.  Nonindexical contextualism , 2007, Synthese.

[55]  Olaf Sporns,et al.  Complex network measures of brain connectivity: Uses and interpretations , 2010, NeuroImage.

[56]  Manfred Pinkal Logic and lexicon - the semantics of the indefinite , 1995, Studies in linguistics and philosophy.

[57]  Klaas E. Stephan,et al.  The anatomical basis of functional localization in the cortex , 2002, Nature Reviews Neuroscience.

[58]  R. Savoy History and future directions of human brain mapping and functional neuroimaging. , 2001, Acta psychologica.

[59]  F. J. Pelletier Kinds, things, and stuff : mass terms and generics , 2010 .

[60]  E. Rosch,et al.  Family resemblances: Studies in the internal structure of categories , 1975, Cognitive Psychology.

[61]  Deirdre N. McCloskey,et al.  The Standard Error of Regressions , 1996 .

[62]  Katrin Amunts,et al.  Locating the functional and anatomical boundaries of human primary visual cortex , 2009, NeuroImage.

[63]  A. Campbell Histological Studies on the Localisation of Cerebral Function , 2009 .

[64]  B. Payne,et al.  Representation of the ipsilateral visual field in the transition zone between areas 17 and 18 of the cat's cerebral cortex , 1990, Visual Neuroscience.

[65]  Douglas H. Johnson The Insignificance of Statistical Significance Testing , 1999 .

[66]  J. Chason The Isocortex of Man , 1952 .

[67]  Andreas Bartels,et al.  fMRI and its interpretations: an illustration on directional selectivity in area V5/MT , 2008, Trends in Neurosciences.

[68]  Duncan J. Watts,et al.  Collective dynamics of ‘small-world’ networks , 1998, Nature.

[69]  Jonathan D. Power,et al.  A Parcellation Scheme for Human Left Lateral Parietal Cortex , 2010, Neuron.

[70]  L. Harlow,et al.  What if there were no significance tests , 1997 .

[71]  G. E. Smith A New Topographical Survey of the Human Cerebral Cortex, being an Account of the Distribution of the Anatomically Distinct Cortical Areas and their Relationship to the Cerebral Sulci. , 1907, Journal of anatomy and physiology.

[72]  G. Bonin,et al.  The isocortex of man , 1951 .

[73]  Timothy Endicott Vagueness in Law , 2001 .

[74]  C. Klein Images Are Not the Evidence in Neuroimaging , 2010, The British Journal for the Philosophy of Science.

[75]  A. Walker,et al.  A cytoarchitectural study of the prefrontal area of the macaque monkey , 1940 .

[76]  Russell A. Poldrack,et al.  In praise of tedious anatomy , 2007, NeuroImage.

[77]  E. Bullmore,et al.  Hierarchical Organization of Human Cortical Networks in Health and Schizophrenia , 2008, The Journal of Neuroscience.

[78]  A. Schleicher,et al.  Architectonics of the human cerebral cortex and transmitter receptor fingerprints: reconciling functional neuroanatomy and neurochemistry , 2002, European Neuropsychopharmacology.

[79]  T. Vicsek,et al.  Uncovering the overlapping community structure of complex networks in nature and society , 2005, Nature.

[80]  K. Fine Vagueness, truth and logic , 1975, Synthese.

[81]  P. Goldman-Rakic,et al.  Cytoarchitectonic definition of prefrontal areas in the normal human cortex: II. Variability in locations of areas 9 and 46 and relationship to the Talairach Coordinate System. , 1995, Cerebral cortex.

[82]  Peter F. Smith,et al.  Vagueness: A Reader , 1999 .