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. 2019 Oct 4;9:14323. doi: 10.1038/s41598-019-50803-y

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© The Author(s) 2019

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Schematic diagram showing the processing of vestibular signals in the brainstem and cerebellum. Physical parameters sensed by the canals and otoliths are angular velocity (w) and gravito-inertial acceleration (GIA) respectively, and central estimates are angular velocity (w’), orientation of gravity (G’), and linear acceleration (A’). Canal inputs are processed through a direct pathway with gain g_D and an indirect (velocity storage, light grey box) pathway which functions as a leaky integrator with gain g_VS and time constant T_VS^33,52. The cerebellar NU (‘tilt estimator,’ dark grey box) inhibits velocity storage (grey arrow #1) by shortening the time constant of the velocity storage integrator. Brainstem regions associated with migraine (e.g., LC = locus coeruleus; DRN = dorsal raphe nucleus) also project to the vestibular nuclei (black arrow #2)–LC projections are noradrenergic⁵³ and DRN projections are serotonergic⁵⁴. Letters in bold are three-dimensional vectors.