Renato Grasso

Eye-head coordination for the steering of locomotion in humans: an anticipatory synergy

We investigated head and gaze orientation in six healthy volunteers walking along 90 degrees corner trajectories, both at light and with eyes closed. We found that head and eyes systematically deviated toward the future direction of the curved trajectory. Anticipation lead was about 1 s. Strikingly, the same behaviour was observed in darkness. In backward (BW) locomotion along the trajectory (from end- to start-point), gaze deviated toward the opposite direction, such that the forward locomotor pattern did not appear time-reversed. Orienting movements displayed higher amplitude, reproducibility and time lead in the forward (FW) direction at light. We suggest that anticipatory orienting synergies belong to the behavioural repertoire of human navigation and may reflect the need to prepare a stable reference frame for intended action.

The predictive brain: anticipatory control of head direction for the steering of locomotion.

THE control of head direction in humans walking along planned circular trajectories was investigated in this study. Five healthy volunteers were asked to walk at a constant speed along circular trajectories in the light and while blindfolded. Head and walking directions were analysed on a real-time basis. Head direction systematically anticipated changes in the direction of locomotion (by about 200ms). The anticipation interval depended on the curvature of the circle. In the light, head orientation was deviated with respect to the walking direction, toward the inner concavity of the performed trajectory. The results suggest that head direction is controlled on a step-by-step basis in a predictive fashion. A ‘go where you look’ strategy seems to underlie steering along circular trajectories.

The contribution of otoliths and semicircular canals to the perception of two‐dimensional passive whole‐body motion in humans

1 Perception of two‐dimensional (2–D) whole‐body passive motion in the horizontal plane was studied in twelve blindfolded healthy volunteers: pure rotation in place (180 deg), linear motion (4.5 m) and a semicircular trajectory (radius, 1.5 m; angular acceleration, 0.2 rad s−2) were applied in random sequence by means of a remote‐controlled robot equipped with a racing‐car seat. The seat orientation in the horizontal plane was controlled by the experimenter, independent of the robot trajectory. Thus different degrees of otolith–canal interaction were obtained. The maximal linear acceleration during the semicircular trajectory was 0.1 g; however, the linear acceleration vector was complex as it rotated relative to the subjects head. 2 In the first of two sessions, subjects were instructed to maintain an angular pointer oriented towards a remote (15 m) previously seen target during the passive movements. In the second session they had to make a drawing of the path of the perceived trajectory, after the movement was finished. 3 The results showed that, on average, the movement of the pointer matched the dynamics of the rotatory component of the 2‐D motion well. This suggests that, in the range of linear accelerations used in this study, no appreciable influence of otolith input on canal‐mediated perception of angular motion occurred. 4 The curvature of the drawn paths was mostly explained by the input to the semicircular canals. Subjects’ reconstruction of motion did not account for the directional dynamics of the input to the otoliths occurring during passive motion. 5 This finding proves that reconstructing trajectory in space does not imply a mathematically perfect transformation of the linear and angular motion‐related inputs into a Cartesian or polar 2‐D representation. Physiological constraints on the interaction between motion direction and change of heading play an important role in motion perception.

Development of Anticipatory Orienting Strategies During Locomotor Tasks in Children

Some basic problems related to the development of goal-directed locomotion in humans are reviewed here. A preliminary study is presented which was aimed at investigating the emergence of anticipatory head orienting strategies during goal-directed locomotion in children. Eight children ranging from 3.5 to 8 years had to walk along a 90 degrees right corner trajectory to reach a goal, both in light and in darkness. The instantaneous orientation in space of the head, trunk, hips and left foot antero/posterior axes was computed by means of an ELITE four-TV camera, 100 Hz system. The results showed that predictive head orienting movements can occur also in the youngest children. The head starts to rotate toward the goal before the corner point of the trajectory is reached. In children, the head peak rotation coincides with the trajectory corner while in adults the peak is attained before. In children, the walking speed is largely decreased in darkness. The results suggest that feedforward control of goal-directed locomotion appears very early in gait development and becomes increasingly important afterwards.

Two-thirds power law in human locomotion: role of ground contact forces.

Are there general rules for the generation of curvilinear motion of the end-effector? Form and kinematics of the arm trajectory are typically inter-related. A relationship between velocity and curvature of the endpoint path has been previously described and quantified as the two-thirds power law. Here we show that the two-thirds power law holds also for the foot trajectory (during the swing phase) in human locomotion for a wide range of walking speeds and gravitational loads, but air-stepping. In air-stepping, it was violated systematically. The results suggest that the power law represents a general constrain of biological motion, may be attributed to both mechanical and neural factors and can depend on natural interactions with external environment.

Plasticity of spinal centers in spinal cord injury patients: new concepts for gait evaluation and training.

Recent data on spinal cord plasticity after spinal cord injury (SCI) were reviewed to analyze the influence of training on the neurophysiological organization of locomotor spinal circuits in SCI patients. In particular, the authors studied the relationship between central pattern generators (CPGs) and motor neuron pool activation during gait. An analysis of the relations between locomotor recovery and compensatory mechanisms focuses on the hierarchical organization of gait parameters and allows characterizing kinematic parameters that are highly stable during different gait conditions and in recovered gait after SCI. The importance of training characteristics and the use of robotic/automated devices in gait recovery is analyzed and discussed. The role of CPG in defining kinematic gait parameters is summarized, and spatio-temporal maps of EMG activity during gait are used to clarify the role of CPG plasticity in sustaining gait recovery.

Circular trajectory formation during blind locomotion: a test for path integration and motor memory

Abstract Eight healthy subjects were asked to walk blindfolded along circular paths of different radii after several practice trials with vision. Their task was to stop after completing two full revolutions. They always walked counter-clockwise (CCW) in (a) a control condition (CONTROL), including the instructions mentioned above, (b) with the further instruction to count backwards in twos (MENTAL), (c) with the instruction to count loudly (LOUD). The movement of two markers lying along the head naso-occipital axis was recorded by means of an ELITE system. Total walked distance (DISTANCE), total head turning angle (ANGLE) and average radius (RADIUS) of the trajectories performed were measured. All subjects were able to perform approximately circular trajectories. They consistently overshot the ideal radius independently of the condition and circle size, undershot the total angle and overshot total distance. The LOUD condition induced greater errors in the performance but only on total distance (P<0.05). A strong correlation was found between the errors in radius and total distance but not between distance and total angle. Principal components analysis suggested that radius and distance share a common source of errors while total angle produced independent errors. The results indicate that (a) circular trajectories can be generated starting from spatial and/or motor memory, without the aid of visual information; (b) the task needs some attentional control and does not involve simple automatic processing of afferent information; (c) different sensory information or different processing modes are probably involved in the estimation of the curvature and length of the walked path on the one hand, and of the total rotation angle on the other.

Spatial, not temporal cues drive predictive orienting movements during navigation: a virtual reality study.

A fundamental property of the human brain is the ability to make predictions of future sensory and motor events. We have recently found that steering manoeuvres when walking along curvilinear trajectories are controlled by an anticipatory guidance of the direction of head (and eyes). However it is unclear whether a time-related or space-related signal triggers such anticipatory head orienting movements. By simulating navigation along a multi-legged virtual corridor we show that anticipatory orienting movements are triggered (in standing subjects) by reaching specific locations rather than by the time to the approaching corridors bend. Similar to what happens in car driving, specific spatial features of the route rather than time to collision seem to drive steering.

Quantitative analysis of human walking trajectory on a circular path in darkness

Thirteen normal (eight young and five older) subjects and a patient who was removed left sided acoustic neurinoma were tested to walk blindfolded along circular paths. They were asked to walk completing two revolutions and to stop when they judged they had returned to the initial position with their head faced to the initial directions. Movements of two markers on the subjects head were recorded by three dimensional motion analyzing system (ELITE system) at 50 Hz which allowed us to measure (a) total walked distance, (b) average radius of the trajectory, and (c) cumulative angle of rotation. Eight young subjects were tested on three circles with radii 0.5, 0.9, and 1.15 m, in two conditions (control, and with mental arythmetic), only to clockwise direction. Five older subjects and a patient were tested on a circle of 0.9 m radius in two conditions, but to both directions, counterclockwise and clockwise. Walked trajectories of young subjects were smooth, whereas those of older subjects tended to be polygonal. Young subjects overshot the ideal distance (6.6%) and ideal radius (9.5%), whereas they undershot the ideal angle (5.1%). There was no effect of circle size or condition on these variables. On the other hand, there was a significant effect of condition on average radius in the older group. The performance of older subjects seemed to be affected by the concurrent mental task. Comparing the counterclockwise walk, the older subjects undershot the turning angle much more than the young subjects which suggest deficits in the vestibular function with aging. The patient showed larger radius and smaller angle while she turned to the healthy side (clockwise) than to the affected side (counterclockwise). Lack of unilateral vestibular information seemed to have affected the circular walking trajectory.

Integration of somatosensory and vestibular inputs in perceiving the direction of passive whole-body motion

We investigated the contribution of somatosensory and vestibular inputs in the detection of the direction of linear whole-body displacement (1.5 m) at low levels of linear acceleration (peak acceleration 0.2 m/s2), in normal subjects. Subjects sat on a mobile robot wearing opaque goggles and headphones. They indicated the direction of motion by using a laser pointer held by the right hand at the level of the chin. Adaptation to a long-lasting static head turn (45 degrees to the right) was used to modify the perceived head orientation relative to the trunk. After about 15 min the head and trunk were perceived to be aligned with each other. After adaptation subjects pointed in the same direction as in the control condition in spite of the change in the perception of the head orientation. Because space orientated reactions to vestibular stimuli were previously shown to be coded in the perceived head reference frame, these results indicate that somatosensory cues are also integrated in the perception of linear motion. Unexpectedly, after adaptation, trunk and head perceived orientations were attracted toward the direction of the imposed motion. This suggests that the internal representation of body configuration depends also upon available cues from the extrapersonal space.