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. 2015 Feb 6;9:10. doi: 10.3389/fncom.2015.00010

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Copyright © 2015 Page, Walters and Stringer.

This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

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Proposed model architecture, with HD cell activity both sustained and shifted by connectivity incoporation axonal conduction delays Δt. This connectivity is reciprocal with a layer of combination (COMB) cells, which learn to respond to a particular combination of HD and AHV through competitive learning. During no head rotation (A), HD activity is stabilized through reciprocal connectivity with COMB cells representing the combination of current head direction and no angular head velocity (AHV = 0). When the head is rotating (B), activity is shifted through the HD layer via connectivity with different COMB cells representing the conjunction of current head direction with angular head velocity (AHV = x, where x is some non-zero value). Connectivity from these COMB cells back onto the HD layer projects with an offset from the original HD activity of 2Δtx; causing HD cell activity to accurately track true head direction given current angular head velocity.