The gradient concept in neuroscience describes systematic and continuous progressions of features of cortical organization across the entire cortex. Recent multimodal studies revealed a macroscale gradient from primary sensory to transmodal association areas which is linked to increasing representational abstraction along the cortical hierarchy, and which is paralleled by microscale gradients of cytoarchitecture and gene expression profiles. Convergent or divergent evidence from these multimodal studies is then used to support inferences about the existence of one common or multiple scale-specific gradients of hierarchical information processing. This paper evaluates the validity of such inferences within the framework of multiscale modeling. In branches of physics and biology where multiscale modeling techniques are used, the simple averaging of microscale details can introduce errors in macroscale modeling if it ignores structures at the intermediate mesoscales of organization which affect system behavior. Conversely, information about mesoscale structures can be used to determine which microscale details are actually relevant to macroscale behavior. In this paper, I similarly argue that multiscale modeling of cortical gradients needs to take organization of mesoscale circuits into account if it affects structure-function relation these models describe. Information about these circuits provides crucial evidence for evaluating inferences from micro- and macroscale data to the role of cortical gradients in hierarchical information processing. My application of the multiscale modeling framework reveals that the gradient concept tracks multiple overlapping progressions of cortical properties, rather than one overall gradient of hierarchical information processing. I support this argument by proposing a mesoscale gradient of connectivity which describes architectural differences between granular and agranular circuits, and which helps us better understand the relation between neural connectivity and hierarchical information processing. Copyright © 2021 The Author(s). Published by Elsevier Inc. All rights reserved.
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