title: Secondary instabilities modulate cortical complexity in the mammalian brain.
abstract: Disclosing the origin of convolutions in the mammalian brain remains a scientific challenge. Primary folds form before we are born: they are static, well defined, and highly preserved across individuals. Secondary folds occur and disappear throughout our entire life time: they are dynamic, irregular, and highly variable among individuals. While extensive research has improved our understanding of primary folding in the mammalian brain, secondary folding remains understudied and poorly understood. Here, we show that secondary instabilities can explain the increasing complexity of our brain surface as we age. Using the nonlinear field theories of mechanics supplemented by the theory of finite growth, we explore the critical conditions for secondary instabilities. We show that with continuing growth, our brain surface continues to bifurcate into increasingly complex morphologies. Our results suggest that even small geometric variations can have a significant impact on surface morphogenesis. Secondary bifurcations, and with them morphological changes during childhood and adolescence, are closely associated with the formation and loss of neuronal connections. Understanding the correlation between neuronal connectivity, cortical thickness, surface morphology, and ultimately behavior, could have important implications on the diagnostics, classification, and treatment of neurological disorders.
authors.affiliation: Chair of Applied Mechanics, Department of Mechanical Engineering, University of Erlangen-Nuremberg, 91058 Erlangen, Germany.
authors.forename: Silvia
authors.initials: S
authors.lastname: Budday
authors.affiliation: Chair of Applied Mechanics, Department of Mechanical Engineering, University of Erlangen-Nuremberg, 91058 Erlangen, Germany.
authors.forename: Paul
authors.initials: P
authors.lastname: Steinmann
authors.affiliation: Departments of Mechanical Engineering and Bioengineering, Stanford University, Stanford, CA 94305, USA.
authors.forename: Ellen
authors.initials: E
authors.lastname: Kuhl
keywords: bifurcation; brain; cortical folding; instability; morphogenesis; period-doubling