Moving and interacting with the environment require a reference for orientation and a scale for calibration in space and time. There is a wide variety of environmental clues and calibrated frames at different locales, but the reference of gravity is ubiquitous on Earth. The pull of gravity on static objects provides a plummet which, together with the horizontal plane, defines a three-dimensional Cartesian frame for visual images. On the other hand, the gravitational acceleration of falling objects can provide a time-stamp on events, because the motion duration of an object accelerated by gravity over a given path is fixed. Indeed, since ancient times, man has been using plumb bobs for spatial surveying, and water clocks or pendulum clocks for time keeping. Here we review behavioral evidence in favor of the hypothesis that the brain is endowed with mechanisms that exploit the presence of gravity to estimate the spatial orientation and the passage of time. Several visual and non-visual (vestibular, haptic, visceral) cues are merged to estimate the orientation of the visual vertical. However, the relative weight of each cue is not fixed, but depends on the specific task. Next, we show that an internal model of the effects of gravity is combined with multisensory signals to time the interception of falling objects, to time the passage through spatial landmarks during virtual navigation, to assess the duration of a gravitational motion, and to judge the naturalness of periodic motion under gravity

Gravity is crucial for spatial perception, postural equilibrium, and movement generation. The vestibular apparatus is the main sensory system involved in monitoring gravity. Hair cells in the vestibular maculae respond to gravitoinertial forces, but they cannot distinguish between linear accelerations and changes of head orientation relative to gravity. The brain deals with this sensory ambiguity (which can cause some lethal airplane accidents) by combining several cues with the otolith signals: angular velocity signals provided by the semicircular canals, proprioceptive signals from muscles and tendons, visceral signals related to gravity, and visual signals. In particular, vision provides both static and dynamic signals about body orientation relative to the vertical, but it poorly discriminates arbitrary accelerations of moving objects. However, we are able to visually detect the specific acceleration of gravity since early infancy. This ability depends on the fact that gravity effects are stored in brain regions which integrate visual, vestibular, and neck proprioceptive signals and combine this information with an internal model of gravity effects.

During gradual speed changes, humans exhibit a sudden discontinuous switch from walking to running at a specific speed, and it has been suggested that different gaits may be associated with different functioning of neuronal networks. In this study we recorded the EMG activity of leg muscles at slow increments and decrements in treadmill belt speed and at different levels of body weight unloading. In contrast to normal walking at 1 g, at lower levels of simulated gravity (<0.4 g) the transition between walking and running was generally gradual, without systematic abrupt changes in either intensity or timing of EMG patterns. This phenomenon depended to a limited extent on the gravity simulation technique, although the exact level of the appearance of smooth transitions (0.4-0.6 g) tended to be lower for the vertical than for the tilted body weight support system. Furthermore, simulations performed with a half-center oscillator neuromechanical model showed that the abruptness of motor patterns at gait transitions at 1 g could be predicted from the distinct parameters anchored already in the normal range of walking and running speeds, whereas at low gravity levels the parameters of the model were similar for the two human gaits. A lack of discontinuous changes in the pattern of speed-dependent locomotor characteristics in a hypogravity environment is consistent with the idea of a continuous shift in the state of a given set of central pattern generators, rather than the activation of a separate set of central pattern generators for each distinct gait.