Knowledge of head direction cell function has progressed remarkably in recent years. The predominant theory that they form an attractor has been confirmed by several experiments. Candidate pathways that may convey visual input have been identified. The pre-subicular circuitry that conveys head direction signals to the medial entorhinal cortex, potentially sustaining path integration by grid cells, has been resolved. Although the neuronal substrate of the attractor remains unknown in mammals, a simple head direction network, whose structure is astoundingly similar to neuronal models theorized decades earlier, has been identified in insects. Finally, recent experiments have revealed that these cells do not encode head direction in the horizontal plane only, but also in vertical planes, thus providing a 3D orientation signal.
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