Large-Scale Gradients in Human Cortical Organization
Advances in neuroimaging technologies and analytics have enabled the discovery of gradients in microstructure, connectivity, gene expression, and function in the human cerebral cortex. The notion that functional processing hierarchies are confined to sensorimotor systems is challenged by recent descriptions of global hierarchies, extending throughout transmodal association areas. An innovative line of research has uncovered a cortical hierarchy in the temporal domain that accounts for spatially distributed functional specialization. Recent advances in mapping cortical areas in the human brain provide a basis for investigating the significance of their spatial arrangement. Here we describe a dominant gradient in cortical features that spans between sensorimotor and transmodal areas. We propose that this gradient constitutes a core organizing axis of the human cerebral cortex, and describe an intrinsic coordinate system on its basis. Studying the cortex with respect to these intrinsic dimensions can inform our understanding of how the spectrum of cortical function emerges from structural constraints. Recent advances in mapping cortical areas in the human brain provide a basis for investigating the significance of their spatial arrangement. Here we describe a dominant gradient in cortical features that spans between sensorimotor and transmodal areas. We propose that this gradient constitutes a core organizing axis of the human cerebral cortex, and describe an intrinsic coordinate system on its basis. Studying the cortex with respect to these intrinsic dimensions can inform our understanding of how the spectrum of cortical function emerges from structural constraints. For more than a century, neuroscientists have studied the cerebral cortex by delineating individual cortical areas (see Glossary) and mapping their function [1Vogt C. Vogt O. Allgemeinere ergebnisse unserer hirnforschung.J. Psychol. Neurol. 1919; 25: 279-468Google Scholar]. This agenda has substantially advanced in recent years, as automated parcellation methods improve and data sets of unprecedented size and quality become available [2Amunts K. Zilles K. Architectonic mapping of the human brain beyond Brodmann.Neuron. 2015; 88: 1086-1107Abstract Full Text Full Text PDF PubMed Scopus (230) Google Scholar, 3Glasser M.F. et al.A multi-modal parcellation of human cerebral cortex.Nature. 2016; 536: 171-178Crossref PubMed Scopus (1992) Google Scholar, 4Eickhoff S.B. et al.Topographic organization of the cerebral cortex and brain cartography.Neuroimage. 2017; (Published online February 20, 2017)https://doi.org/10.1016/j.neuroimage.2017.02.018Crossref Scopus (82) Google Scholar]. Nevertheless, our understanding of how the complex structure of the cerebral cortex emerges and gives rise to its elaborate functions remains fragmentary. To complement the description of individual cortical areas, we propose an inquiry into the significance of their spatial arrangement, asking the basic question: Why are cortical areas located where they are? Early formulations of this question date to theories from classical neuroanatomy [1Vogt C. Vogt O. Allgemeinere ergebnisse unserer hirnforschung.J. Psychol. Neurol. 1919; 25: 279-468Google Scholar, 5Brockhaus H. Die cyto-und myeloarchitektonik des cortex claustralis und des claustrum beim menschen.J. Psychol. Neurol. 1940; 49: 249-348Google Scholar, 6Sanides F. Die Architektonik des Menschlichen Stirnhirns. Springer, 1962Crossref Google Scholar, 7Pandya D. et al.Cerebral Cortex: Architecture, Connections, and the Dual Origin Concept. Oxford University Press, 2015Crossref Google Scholar]. They state that the spatial layout of cortical areas is not arbitrary, but a consequence of developmental mechanisms, shaped through evolutionary selection. The location of an area among its neighbors thus provides insight into its microstructural characteristics [6Sanides F. Die Architektonik des Menschlichen Stirnhirns. Springer, 1962Crossref Google Scholar], its connections to other parts of the brain [7Pandya D. et al.Cerebral Cortex: Architecture, Connections, and the Dual Origin Concept. Oxford University Press, 2015Crossref Google Scholar], and eventually its position in global processing hierarchies [8Mesulam M.M. From sensation to cognition.Brain. 1998; 121: 1013-1052Crossref PubMed Scopus (2101) Google Scholar]. Consider, for example, the well-researched visual system of the macaque monkey [9Felleman D.J. Van Essen D.C. Distributed hierarchical processing in the primate cerebral cortex.Cereb. Cortex. 1991; 1: 1-47Crossref PubMed Scopus (5448) Google Scholar, 10Markov N.T. et al.A weighted and directed interareal connectivity matrix for macaque cerebral cortex.Cereb. Cortex. 2014; 24: PubMed Scopus Google Scholar]. the visual visual features are and with from other areas are on their of microstructural and the of their connections as H. in the of Neurol. PubMed Scopus Google Scholar]. The we that an position in the visual processing hierarchy and thus of its microstructural and features is to its from the visual area K. et gradients in structural 2015; PubMed Scopus Google Scholar, a gradient in the of the 2015; PubMed Scopus Google Scholar]. we propose that the spatial of areas a global gradient between sensorimotor and transmodal is a of human cortical gradient is an axis of in cortical areas in a spatially that other with respect to the of the we is a between the of areas that their position the gradient and their position the cortical This of cortical location a in we to the cerebral cortex with respect to its intrinsic this we on cortical microstructure, connectivity, and gene expression, to a dominant gradient of cortical we how this gradient can how the spectrum of functions from cortical we propose a intrinsic coordinate system of the human cerebral cortex. in the is for a global gradient in human cortical spans between sensorimotor and transmodal and is in cortical microstructure, connectivity, and gene these cortical features have to areas, the of an gradient not the of areas, but their spatial into The of cortical gradients is in classical neuroanatomy [1Vogt C. Vogt O. Allgemeinere ergebnisse unserer hirnforschung.J. Psychol. Neurol. 1919; 25: 279-468Google Scholar, 5Brockhaus H. Die cyto-und myeloarchitektonik des cortex claustralis und des claustrum beim menschen.J. Psychol. Neurol. 1940; 49: 249-348Google Scholar, 6Sanides F. Die Architektonik des Menschlichen Stirnhirns. Springer, 1962Crossref Google Scholar, 7Pandya D. et al.Cerebral Cortex: Architecture, Connections, and the Dual Origin Concept. 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