While it has been well established that optostatic and optokinetic cues contribute to the perception of vertical, it is unclear how the brain processes their combined presence with the nonvisual vestibular cues. Using a psychometric approach, we examined the percept of vertical in human participants ( n = 17) with their body and head upright, presented with a visual frame tilted at one of eight orientations (between ±45°, steps of 11.25°) or no frame, surrounded by an optokinetic roll-stimulus (velocity =  ±30°/s or stationary). Both cues demonstrate relatively independent biases on vertical perception, with a sinusoidal modulation by frame orientation of ~4° and a general shift of ~1–2° in the rotation direction of the optic flow. Variability was unaffected by frame orientation but was higher with than without optokinetic rotation. An optimal-observer model in which vestibular, optostatic, and optokinetic cues provide independent sources to vertical perception was unable to explain these data. In contrast, a model in which the optokinetic signal biases the internal representation of gravity, which is then optimally integrated with the optostatic cue, provided a good account, at the individual participant level. We conclude that optostatic and optokinetic cues interact differently with vestibular cues in the neural computations for vertical perception. NEW & NOTEWORTHY Static and dynamic visual cues are known to bias the percept of vertical, but how they interact with vestibular cues remains to be established. Guided by an optimal-observer model, the present results suggest that optokinetic information is combined with vestibular information into a single, vestibular-optokinetic estimate, which is integrated with an optostatically derived estimate of vertical.

The brain is thought to use rotation cues from both the vestibular and optokinetic system to disambiguate the gravito-inertial force, as measured by the otoliths, into components of linear acceleration and gravity direction relative to the head. Hence, when the head is stationary and upright, an erroneous percept of tilt arises during optokinetic roll stimulation (OKS) or when an artificial canal-like signal is delivered by means of galvanic vestibular stimulation (GVS). It is still unknown how this percept is affected by the combined presence of both cues or how it develops over time. Here, we measured the time course of the subjective visual vertical (SVV), as a proxy of perceived head tilt, in human participants ( n = 16) exposed to constant-current GVS (1 and 2 mA, cathodal and anodal) and constant-velocity OKS (30°/s clockwise and counterclockwise) or their combination. In each trial, participants continuously adjusted the orientation of a visual line, which drifted randomly, to Earth vertical. We found that both GVS and OKS evoke an exponential time course of the SVV. These time courses have different amplitudes and different time constants, 4 and 7 s respectively, and combine linearly when the two stimulations are presented together. We discuss these results in the framework of observer theory and Bayesian state estimation. NEW & NOTEWORTHY While it is known that both roll optokinetic stimuli and galvanic vestibular stimulation affect the percept of vertical, how their effects combine and develop over time is still unclear. Here we show that both effects combined linearly but are characterized by different time constants, which we discuss from a probabilistic perspective.

The brain can use multiple reference frames to code line orientation, including head-, object-, and gravity-centered references. If these frames change orientation, their representations must be updated to keep register with actual line orientation. We tested this internal updating during head rotation in roll, exploiting the rod-and-frame effect: The illusory tilt of a vertical line surrounded by a tilted visual frame. If line orientation is stored relative to gravity, these distortions should also affect the updating process. Alternatively, if coding is head- or frame-centered, updating errors should be related to the changes in their orientation. Ten subjects were instructed to memorize the orientation of a briefly flashed line, surrounded by a tilted visual frame, then rotate their head, and subsequently judge the orientation of a second line relative to the memorized first while the frame was upright. Results showed that updating errors were mostly related to the amount of subjective distortion of gravity at both the initial and final head orientation, rather than to the amount of intervening head rotation. In some subjects, a smaller part of the updating error was also related to the change of visual frame orientation. We conclude that the brain relies primarily on a gravity-based reference to remember line orientation during head roll.