(a) Equivalence principle: the otolith organs are sensitive to the gravitoinertial acceleration (GIA), equal to the difference between the gravity vector (GA) and the translational acceleration (TA). (b) Naming conventions of the head's translation and rotation axes. FB, forward-backward; LR, leftward-rightward. (c) Representation of the motion protocols used by Laurens et al. [9]. GIA along the LR axis, represented by a swinging pendulum (bottom), is identical in the 3 protocols (translation, tilt, and off-vertical axis rotation [OVAR]). ((d)–(o)) Responses from a translation-selective cell (red) and a tilt-selective cell (green) during left-right (LR) translation ((d) and (h)), roll tilt ((e) and (i)), and constant velocity OVAR ((f), (g), (j), and (k)). (l), (m), (n), and (o) show the corresponding yaw velocity (detected by horizontal canals, blue), roll velocity (detected by vertical canals, cyan), and GIA along the LR axis (detected by otolith organs [OTO], black). Gray curves: fit to the LR translation response (shown in (d), translation cell) or the roll tilt response (shown in (i), tilt cell) (reproduced with permission from [5]).