When up is down in 0g: how gravity sensing affects the timing of interceptive actions.

Humans are known to regulate the timing of interceptive actions by modeling, in a simplified way, Newtonian mechanics. Specifically, when intercepting an approaching ball, humans trigger their movements a bit earlier when the target arrives from above than from below. This bias occurs regardless of the ball's true kinetics, and thus appears to reflect an a priori expectation that a downward moving object will accelerate. We postulate that gravito-inertial information is used to tune visuomotor responses to match the target's most likely acceleration. Here we used the peculiar conditions of parabolic flight--where gravity's effects change every 20 s--to test this hypothesis. We found a striking reversal in the timing of interceptive responses performed in weightlessness compared with trials performed on ground, indicating a role of gravity sensing in the tuning of this response. Parallels between these observations and the properties of otolith receptors suggest that vestibular signals themselves might plausibly provide the critical input. Thus, in addition to its acknowledged importance for postural control, gaze stabilization, and spatial navigation, we propose that detecting the direction of gravity's pull plays a role in coordinating quick reactions intended to intercept a fast-moving visual target.

Distractor-induced saccade trajectory curvature reveals visual contralateral bias with respect to the dominant eye.

The functional consequences of the visual system lateralization referred to as "eye dominance" remain poorly understood. We previously reported shorter hand reaction times for targets appearing in the contralateral visual hemifield with respect to the dominant eye (DE). Here, we further explore this contralateral bias by studying the influence of laterally placed visual distractors on vertical saccade trajectories, a sensitive method to assess visual processing. In binocular conditions, saccade trajectory curvature was larger toward a distractor placed in the contralateral hemifield with respect to the DE (e.g., in the left visual hemifield for a participant with a right dominant eye) than toward one presented in the ipsilateral hemifield (in the right visual hemifield in our example). When two distractors were present at the same time, the vertical saccade showed curvature toward the contralateral side. In monocular conditions, when one distractor was presented, a similar larger influence of the contralateral distractor was observed only when the viewing eye was the DE. When the non dominant eye (NDE) was viewing, curvature was symmetric for both distractor sides. Interestingly, this curvature was as large as the one obtained for the contralateral distractor when the DE was viewing, suggesting that eye dominance consequences rely on inhibition mechanisms present when the DE is viewing. Overall, these results demonstrate that DE influences visual integration occurring around saccade production and support a DE-based contralateral visual bias.

Effect of weight-related labels on corticospinal excitability during observation of grasping: a TMS study.

Recent studies of corticospinal excitability during observation of grasping and lifting of objects of different weight have highlighted the role of agent's kinematics in modulating observer's motor excitability. Here, we investigate whether explicit weight-related information, provided by written labels on the objects, modulate the excitability of the observer's motor system and how this modulation is affected when there is a conflict between label and object's weight. We measured TMS-evoked motor potentials (MEPs) from right hand intrinsic muscles, while subjects were observing an actor lifting objects of different weights, in some trials labeled (heavy/light) in congruent or incongruent way. Results confirmed a weight-related modulation of MEPs based on kinematic cues. Interestingly, any conflict between the labels and the actual weight (i.e., explicit versus implicit information), although never consciously noticed by the observer, deeply affected the mirroring of others' actions. Our findings stress the automatic involvement of the mirror-neuron system.

Force requirements of observed object lifting are encoded by the observer's motor system: a TMS study.

Several transcranial magnetic stimulation (TMS) studies have reported facilitation of the primary motor cortex (M1) during the mere observation of actions. This facilitation was shown to be highly congruent, in terms of somatotopy, with the observed action, even at the level of single muscles. With the present study, we investigated whether this muscle-specific facilitation of the observer's motor system reflects the degree of muscular force that is exerted in an observed action. Two separate TMS experiments are reported in which corticospinal excitability was measured in the hand area of M1 while subjects observed the lifting of objects of different weights. The type of action 'grasping-and-lifting-the-object' was always identical, but the grip force varied according to the object's weight. In accordance to previous findings, excitability of M1 was shown to modulate in a muscle-specific way, such that only the cortical representation areas in M1 that control the specific muscles used in the observed lifting action became increasingly facilitated. Moreover, muscle-specific M1 facilitation was shown to modulate to the force requirements of the observed actions, such that M1 excitability was considerably higher when observing heavy object lifting compared with light object lifting. Overall, these results indicate that different levels of observed grip force are mirrored onto the observer's motor system in a highly muscle-specific manner. The measured force-dependent modulations of corticospinal excitability in M1 are hypothesized to be functionally relevant for scaling the observed grip force in the observer's own motor system. In turn, this mechanism may contribute, at least partly, to the observer's ability to infer the weight of the lifted object.

Egocentric and allocentric reference frames for catching a falling object.

When programming movement, one must account for gravitational acceleration. This is particularly important when catching a falling object because the task requires a precise estimate of time-to-contact. Knowledge of gravity's effects is intimately linked to our definition of 'up' and 'down'. Both directions can be described in an allocentric reference frame, based on visual and/or gravitational cues, or in an egocentric reference frame in which the body axis is taken as vertical. To test which frame humans use to predict gravity's effect, we asked participants to intercept virtual balls approaching from above or below with artificially controlled acceleration that could be congruent or not with gravity. To dissociate between these frames, subjects were seated upright (trunk parallel to gravity) or lying down (body axis orthogonal to the gravitational axis). We report data in line with the use of an allocentric reference frame and discuss its relevance depending on available gravity-related cues.

Visuo-motor coordination and internal models for object interception.

Intercepting and avoiding collisions with moving objects are fundamental skills in daily life. Anticipatory behavior is required because of significant delays in transforming sensory information about target and body motion into a timed motor response. The ability to predict the kinematics and kinetics of interception or avoidance hundreds of milliseconds before the event may depend on several different sources of information and on different strategies of sensory-motor coordination. What are exactly the sources of spatio-temporal information and what are the control strategies remain controversial issues. Indeed, these topics have been the battlefield of contrasting views on how the brain interprets visual information to guide movement. Here we attempt a synthetic overview of the vast literature on interception. We discuss in detail the behavioral and neurophysiological aspects of interception of targets falling under gravity, as this topic has received special attention in recent years. We show that visual cues alone are insufficient to predict the time and place of interception or avoidance, and they need to be supplemented by prior knowledge (or internal models) about several features of the dynamic interaction with the moving object.

Internal models and prediction of visual gravitational motion.

Baurès et al. [Baurès, R., Benguigui, N., Amorim, M.-A., & Siegler, I. A. (2007). Intercepting free falling objects: Better use Occam's razor than internalize Newton's law. Vision Research, 47, 2982-2991] rejected the hypothesis that free-falling objects are intercepted using a predictive model of gravity. They argued instead for "a continuous guide for action timing" based on visual information updated till target capture. Here we show that their arguments are flawed, because they fail to consider the impact of sensori-motor delays on interception behaviour and the need for neural compensation of such delays. When intercepting a free-falling object, the delays can be overcome by a predictive model of the effects of gravity on target motion.

Internal model of gravity influences configural body processing.

Human bodies are processed by a configural processing mechanism. Evidence supporting this claim is the body inversion effect, in which inversion impairs recognition of bodies more than other objects. Biomechanical configuration, as well as both visual and embodied expertise, has been demonstrated to play an important role in this effect. Nevertheless, the important factor of body inversion effect may also be linked to gravity orientation since gravity is one of the most fundamental constraints of our biology, behavior, and perception on Earth. The visual presentation of an inverted body in a typical body inversion paradigm turns the observed body upside down but also inverts the implicit direction of visual gravity in the scene. The orientation of visual gravity is then in conflict with the direction of actual gravity and may influence configural processing. To test this hypothesis, we dissociated the orientations of the body and of visual gravity by manipulating body posture. In a pretest we showed that it was possible to turn an avatar upside down (inversion relative to retinal coordinates) without inverting the orientation of visual gravity when the avatar stands on his/her hands. We compared the inversion effect in typical conditions (with gravity conflict when the avatar is upside down) to the inversion effect in conditions with no conflict between visual and physical gravity. The results of our experiment revealed that the inversion effect, as measured by both error rate and reaction time, was strongly reduced when there was no gravity conflict. Our results suggest that when an observed body is upside down (inversion relative to participants' retinal coordinates) but the orientation of visual gravity is not, configural processing of bodies might still be possible. In this paper, we discuss the implications of an internal model of gravity in the configural processing of observed bodies.

Asymmetry in visual information processing depends on the strength of eye dominance.

Unlike handedness, sighting eye dominance, defined as the eye unconsciously chosen when performing monocular tasks, is very rarely considered in studies investigating cerebral asymmetries. We previously showed that sighting eye dominance has an influence on visually triggered manual action with shorter reaction time (RT) when the stimulus appears in the contralateral visual hemifield with respect to the dominant eye (Chaumillon et al. 2014). We also suggested that eye dominance may be more or less pronounced depending on individuals and that this eye dominance strength could be evaluated through saccadic peak velocity analysis in binocular recordings (Vergilino-Perez et al. 2012). Based on these two previous studies, we further examine here whether the strength of the eye dominance can modulate the influence of this lateralization on manual reaction time. Results revealed that participants categorized as having a strong eye dominance, but not those categorized as having a weak eye dominance, exhibited the difference in RT between the two visual hemifields. This present study reinforces that the analysis of saccade peak velocity in binocular recordings provides an effective tool to better categorize the eye dominance. It also shows that the influence of eye dominance in visuo-motor tasks depends on its strength. Our study also highlights the importance of considering the strength of eye dominance in future studies dealing with brain lateralization.

Estimating time to contact and impact velocity when catching an accelerating object with the hand.

To catch a moving object with the hand requires precise coordination between visual information about the target's motion and the muscle activity necessary to prepare for the impact. A key question remains open as to if and how a human observer uses velocity and acceleration information when controlling muscles in anticipation of impact. Participants were asked to catch the moving end of a swinging counterweighted pendulum, and resulting muscle activities in the arm were measured. The authors also simulated muscle activities that would be produced according to different tuning strategies. By comparing data with simulations, the authors provide evidence that human observers use online information about velocity but not acceleration when preparing for impact.

Are there any left-right asymmetries in saccade parameters? Examination of latency, gain, and peak velocity.

PURPOSE: Hemispheric specialization in saccadic control is still under debate. Here we examine the latency, gain, and peak velocity of reactive and voluntary leftward and rightward saccades to assess the respective roles of eye and hand dominance. METHODS: Participants with contrasting hand and eye dominance were asked to make saccades toward a target displayed at 5°, 10°, or 15° left or right of the central fixation point. In separate sessions, reactive and voluntary saccades were elicited by Gap-200, Gap-0, Overlap-600, and Antisaccade procedures. RESULTS: Left-right asymmetries were not found in saccade latencies but appeared in saccade gain and peak velocity. Regardless of the dominant hand, saccades directed to the ipsilateral side relative to the dominant eye had larger amplitudes and faster peak velocities. CONCLUSIONS: Left-right asymmetries can be explained by naso-temporal differences for some subjects and by eye dominance for others. Further investigations are needed to examine saccadic parameters more systematically in relation to eye dominance. Indeed, any method that allows one to determine ocular dominance from objective measures based on saccade parameters should greatly benefit clinical applications, such as monovision surgery.

Cortical dynamics of anticipatory mechanisms in interception: a neuromagnetic study.

Humans demonstrate an amazing ability for intercepting and catching moving targets, most noticeably in fast-speed ball games. However, the few studies exploring the neural bases of interception in humans and the classical studies on visual motion processing and visuomotor interactions have reported rather long latencies of cortical activations that cannot explain the performances observed in most natural interceptive actions. The aim of our experiment was twofold: (1) describe the spatio-temporal unfolding of cortical activations involved in catching a moving target and (2) provide evidence that fast cortical responses can be elicited by a visuomotor task with high temporal constraints and decide if these responses are task or stimulus dependent. Neuromagnetic brain activity was recorded with whole-head coverage while subjects were asked to catch a free-falling ball or simply pay attention to the ball trajectory. A fast, likely stimulus-dependent, propagation of neural activity was observed along the dorsal visual pathway in both tasks. Evaluation of latencies of activations in the main cortical regions involved in the tasks revealed that this entire network of regions was activated within 40 msec. Moreover, comparison of experimental conditions revealed similar patterns of activation except in contralateral sensorimotor regions where common and catch-specific activations were differentiated.

Anticipating the effects of gravity when intercepting moving objects: differentiating up and down based on nonvisual cues.

Intercepting an object requires a precise estimate of its time of arrival at the interception point (time to contact or "TTC"). It has been proposed that knowledge about gravitational acceleration can be combined with first-order, visual-field information to provide a better estimate of TTC when catching falling objects. In this experiment, we investigated the relative role of visual and nonvisual information on motor-response timing in an interceptive task. Subjects were immersed in a stereoscopic virtual environment and asked to intercept with a virtual racket a ball falling from above or rising from below. The ball moved with different initial velocities and could accelerate, decelerate, or move at a constant speed. Depending on the direction of motion, the acceleration or deceleration of the ball could therefore be congruent or not with the acceleration that would be expected due to the force of gravity acting on the ball. Although the best success rate was observed for balls moving at a constant velocity, we systematically found a cross-effect of ball direction and acceleration on success rate and response timing. Racket motion was triggered on average 25 ms earlier when the ball fell from above than when it rose from below, whatever the ball's true acceleration. As visual-flow information was the same in both cases, this shift indicates an influence of the ball's direction relative to gravity on response timing, consistent with the anticipation of the effects of gravity on the flight of the ball.