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Visual control of reaching movements without vision of the limb. II. Evidence of fast unconscious processes correcting the trajectory of the hand to the final position of a double-step stimulus.

In this study, a visual target was localized by both limb and eye. The experimental procedure provided an opportunity to analyze the limb movement trajectories to the target whose location was displaced during saccades. Absence of visual information about position of the moving limb did not interfere with correction of the trajectory of pointing movements. These corrections reflect the new information about target position that becomes available at the end of the first saccade. Mean localization errors to stationary and to displaced targets were not significantly different. This result suggests that subjects were able to compare visual (retinal + eye position) information about the position of the target with information about the position of their moving limb derived from kinesthesis and/or efference copies of the motor commands. An analysis of velocity profiles indicates that the observed corrections of hand movement to target displacement could not be identified by an inflexion point in the trajectory. None of the subjects reported seeing the target change location. In other words, the motor command was adjustable despite the failure of changes in visual locus to reach consciousness.

Adult↗

The development of goal-directed reaching in infants: hand trajectory formation and joint torque control.

Nine young infants were followed longitudinally from 4 to 15 months of age. We recorded early spontaneous movements and reaching movements to a stationary target. Time-position data of the hand (endpoint), shoulder, and elbow were collected using an optoelectronic measurement system (ELITE). We analyzed the endpoint kinematics and the intersegmental dynamics of the shoulder and elbow joint to investigate how changes in proximal torque control determined the development of hand trajectory formation. Two developmental phases of hand trajectory formation were identified: a first phase of rapid improvements between 16 and 24 weeks of age, the time of reaching onset for all infants. During that time period the number of movement units per reach and movement time decreased dramatically. In a second phase (28-64 weeks), a period of "fine-tuning" of the sensorimotor system, we saw slower, more gradual changes in the endpoint kinematics. The analysis of the underlying intersegmental joint torques revealed the following results: first, the range of muscular and motion-dependent torques (relative to body weight) did not change significantly with age. That is, early reaching was not confined by limitations in producing task-adequate levels of muscular torque. Second, improvements in the endpoint kinematics were not accomplished by minimizing amplitude of muscle and reactive torques. Third, the relative timing of muscular and motion-dependent torque peaks showed a systematic development toward an adult timing profile with increasing age. In conclusion, the development toward invariant characteristics of the hand trajectory is mirrored by concurrent changes in the control of joint forces. The acquisition of stable patterns of intersegmental coordination is not achieved by simply regulating force amplitude, but more so by modulating the correct timing of joint force production and by the system's use of reactive forces. Our findings support the view that development of reaching is a process of unsupervised learning with no external or innate teacher prescribing the desired kinematics or kinetics of the movement.

Adult↗

Projected free fall trajectories. I. Theory and simulation.

How we manage to reconstruct the three-dimensional character of the world from the two-dimensional representations on our retinae has been a lively subject of research in the last ten or fifteen years. One principle that has emerged unifying many of these ideas is the need for constraints to allow the visual system to interpret the images it receives as three-dimensional. These constraints come from assumptions about the nature of the situation that produced the image. We have looked at how gravity can be used as a constraint in the case of a free fall trajectory projected onto an image plane by central projection. We have examined several possible methods for deriving the initial conditions of the trajectory from the two-dimensional projection, and examined their behavior under noisy and noiseless conditions, using both image simulations and videotapes of a real ball. We show that there are several ways to robustly compute the initial conditions of the parabolic trajectory from the image data in the presence of noise.

Humans↗

Projected free fall trajectories. II. Human experiments.

We are very adept at using the purely two-dimensional information we get from our retinae to manoever and react to the three-dimensional world: witness the tennis player returning a 100 mph serve. In another article (Saxberg 1987) we have shown how gravity can in theory be used as a constraint to determine the initial conditions of a three-dimensional free fall trajectory from the two-dimensional central projection of the trajectory. We have developed a simple video game to investigate what information is important to humans trying to solve a problem of this sort: predicting where a ball will fall. We show that humans do not seem to use the trajectory information suggested by the theoretical results in (Saxberg 1985), but rely on other sources of information, such as the size of the image on the projection surface.

Humans↗

Three dimensional arm trajectories.

Planar arm trajectories are characterized by a segmentation of the hand velocity profile and by a coupling between shape and speed. The question addressed in this paper is whether such coupling, observed in two dimensions, still holds in three dimensions. This matter was investigated experimentally by recording three dimensional "aimless" movements of the arm, particularly three dimensional scribbles, and the answer suggested by the experimental data is that only the "bending" of the trajectory is coupled with speed, whereas the "twisting" is independent of speed. The same behaviour was also found to characterize a computational model of trajectory formation which is based on the spatial composition of chains of planar strokes, overlapped in time.

Adult↗

Selective actions of anesthetic agents on membrane potential trajectory in bulbar respiratory neurons of cats.

The effects of two anesthetic agents, halothane and thiopental, on the membrane potential trajectory of respiratory-related neurons in the ventral respiratory group were investigated in decerebrate cats, of which the carotid sinus and vagal afferents were denervated. Infusion of halothane (2% for 90 s) depolarized the membrane in nearly half of the inspiratory (12/21), post-inspiratory (10/26) and expiratory (4/6) neurons and caused hyperpolarization in the rest of the population. Thiopental (2.5 mg/kg i.v.) produced depolarization in 11 inspiratory and 10 post-inspiratory neurons and hyperpolarization in 1 expiratory, 4 inspiratory and 7 post-inspiratory neurons. In both hyperpolarized and depolarized neurons, reduction of the respiratory membrane potential fluctuations and an increase of input resistance were commonly observed. Both drugs depressed spontaneous firing in most of the neurons studied. An increase of firing was observed in 9 out of 47 depolarized cells. These two contrasting effects on the membrane potential trajectory occurred similarly in the known groups of respiratory neurons, but the response of a given cell was consistent for the two anesthetic agents. The present results demonstrate that the anesthetic drugs exert various influences on the ventral respiratory group neuron population in maintaining the membrane potential trajectory and discharge activity. This may reflect a functional heterogeneity in the bulbar respiratory network of neurons.

Action Potentials↗

Simulation of head movement trajectories: model and fit to main sequence.

A sixth order nonlinear model for horizontal head rotations in humans is presented and investigated using experimental results on head movement trajectories and neck muscle EMG. The controller signals, structured in accordance with time optimal control theory, are parameterized, and controller signal parameter variations show a dominating influence on different aspects of the head movement trajectory. The model fits the common head acceleration types over a wide range of amplitudes, and also less common (dynamic overshoot) trajectories.

Electromyography↗

A neural network model for limb trajectory formation.

This paper deals with the problem of representing and generating unconstrained aiming movements of a limb by means of a neural network architecture. The network produced time trajectories of a limb from a starting posture toward targets specified by sensory stimuli. Thus the network performed a sensory-motor transformation. The experimenters trained the network using a bell-shaped velocity profile on the trajectories. This type of profile is characteristic of most movements performed by biological systems. We investigated the generalization capabilities of the network as well as its internal organization. Experiments performed during learning and on the trained network showed that: (i) the task could be learned by a three-layer sequential network; (ii) the network successfully generalized in trajectory space and adjusted the velocity profiles properly; (iii) the same task could not be learned by a linear network; (iv) after learning, the internal connections became organized into inhibitory and excitatory zones and encoded the main features of the training set; (v) the model was robust to noise on the input signals; (vi) the network exhibited attractor-dynamics properties; (vii) the network was able to solve the motor-equivalence problem. A key feature of this work is the fact that the neural network was coupled to a mechanical model of a limb in which muscles are represented as springs. With this representation the model solved the problem of motor redundancy.

Extremities↗

Discrete and continuous planning of hand movements and isometric force trajectories.

We have previously demonstrated that, in preparing themselves to aim voluntary impulses of isometric elbow force to unpredictable targets, subjects selected default values for amplitude and direction according the range of targets that they expected. Once a specific target appeared, subjects specified amplitude and direction through parallel processes. Amplitude was specified continuously from an average or central default; direction was specified stochastically from one of the target directions. Using the same timed response paradigm, we now report three experiments to examine how the time available for processing target information influences trajectory characteristics in two-degree-of-freedom forces and multijoint movements. We first sought to determine whether the specification of force direction could also take the form of a discrete stochastic process in pulses of wrist muscle force, where direction can vary continuously. With four equiprobable targets (two force amplitudes in each of two directions separated by 22 degrees or 90 degrees), amplitude was specified from a central default value for both narrow and wide target separations as a continuous variable. Direction, however, remained specified as a discrete variable for wide target separations. For narrow target separations, the directional distribution of default responses suggested the presence of both discrete and central values. We next examined point-to-point movements in a multijoint planar hand movement task with targets at two distances and two directions but at five directional separations (from 30 degrees to 150 degrees separation). We found that extent was again specified continuously from a central default. Direction was specified discretely from alternative default directions when target separation was wide and continuously from a central default when separation was narrow. The specification of both extent and direction evolved over a 200-ms time period beginning about 100 ms after target presentation. As in elbow force pulses, extent was specified progressively in both correct and wrong direction responses through a progressive improvement in the scaling of acceleration and velocity peaks to the target. On the other hand, movement time and hand path straightness did not change significantly in the course of specification. Thus, the specification of movement time and linearity, global features of the trajectories, are given priority over the specific values of extent and direction. In a third experiment, we varied the distances between unidirectional target pairs and found that movement extent is specified discretely, like direction, when the disparity in distances is large. The implications of these findings for contextual effects on trajectory planning are discussed. The independence of extent and direction specification and the prior setting of response duration and straightness provide critical support for the hypothesis that point-to-point movements are planned vectorially.

Adult↗

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

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.

Adult↗

Spatiotemporal consistency of human mandibular and head-neck movement trajectories during jaw opening-closing tasks.

Previous studies of spatial and temporal coordination between human mandibular and head-neck movements during single as well as rhythmic jaw opening-closing tasks suggest that these movements are regulated by central nervous commands common for jaw and neck muscles. The present study evaluated the spatiotemporal consistency of concomitant mandibular and head-neck movements during repeated single jaw opening-closing tasks, in short- as well as long-term perspectives and at different speeds. The subjects were seated in an upright position without head support. They were instructed to perform single maximal jaw opening-closing movements, repeated ten times for fast and slow speed, respectively. Recordings were made at two sessions. A wireless optoelectronic technique was used to record three-dimensional mandibular and head-neck movements. The spatiotemporal consistency of repeated movement trajectories of the mandible (both in relation to the head and in space) and of the head-neck was quantitatively assessed by a spatiotemporal index (STI). In addition, mean movement trajectory patterns were compared for speed and recording sessions (expressed as correlation coefficient, r). The results showed relatively small STI (high degree of spatiotemporal consistency) and high r (reproducible trajectory patterns) values, both in short- and long-term perspectives, for concomitant mandibular and head-neck movements. The data were not generally speed related. In conclusion, the present results suggest a high degree of spatiotemporal consistency of the kinematic patterns of mandibular and head-neck movements during jaw opening-closing, in short- as well as long-term perspectives. They also indicate that underlying neural processes are invariant in nature. The results give further support to the notion of a tight functional coupling between the human jaw and the neck motor systems during natural jaw function.

Adult↗

Adaptation of arm trajectory during continuous drawing movements in different dynamic environments.

Human subjects can readily adapt their movement trajectories to different dynamic or visuomotor environments. The focus of the current study was to determine whether subjects could simultaneously adapt to multiple dynamic environments. Subjects ( n=5) drew ellipses continuously for 70 s using a torquable manipulandum under six distinct dynamic conditions, representing the combination of load type (spring, viscous, and inertia) and load direction (assisting and opposing). Each subject performed two control, ten load, and five washout trials. A significant effect of force condition on the trajectory of the movement was found in 26 of 30 cases (6 conditions x 5 subjects); the magnitude of the distortion differed across the conditions. The extent of adaptation also differed across the loads. Opposing inertia and viscosity led to fast adaptation. However, assisting inertia and viscosity were associated with relatively slow adaptation. The results of adaptation to the stiffness conditions were not consistent. Following sudden removal of the load we saw an additional disturbance of the trajectory (after-effect), which was often the mirror image of the original distortion. The shape and size of the after-effect were different across load conditions. These results show that human subjects can adapt to a variety of different dynamic transformations and that the time-course of adaptation is dependent on both the state space and the direction of the load.

Adaptation, Physiological↗

Visuo-vestibular interaction in the reconstruction of travelled trajectories.

We recently published a study of the reconstruction of passively travelled trajectories from optic flow. Perception was prone to illusions in a number of conditions, and was not always veridical in the others. Part of the illusionary reconstructed trajectories could be explained by assuming that subjects base their reconstruction on the ego-motion percept built during the stimulus' initial moments. In the current paper, we test this hypothesis using a novel paradigm: if the final reconstruction is governed by the initial percept, providing additional, extra-retinal information that modifies the initial percept should predictably alter the final reconstruction. The extra-retinal stimulus was tuned to supplement the information that was under-represented or ambiguous in the optic flow; the subjects were physically displaced or rotated at the onset of the visual stimulus. A highly asymmetric velocity profile (high acceleration, very low deceleration) was used. Subjects were required to guide an input device (in the form of a model vehicle; we measured position and orientation) along the perceived trajectory. We show for the first time that a vestibular stimulus of short duration can influence the perception of a much longer-lasting visual stimulus. Perception of the ego-motion translation component in the visual stimulus was improved by a linear physical displacement, perception of the ego-motion rotation component by a physical rotation. This led to a more veridical reconstruction in some conditions, but to a less veridical reconstruction in other conditions.

Acceleration↗

Initiation of rapid reach-and-grasp balance reactions: is a pre-formed visuospatial map used in controlling the initial arm trajectory?

In order to recover balance by grasping an object for support, the CNS must rapidly move the hand toward a specific target (handhold) in the environment. The early latency (80-140 ms) of these grasping reactions would seem to preclude a role for online visual feedback in the control of the initial limb movement; however, some studies have shown that vision can influence initiation of lower-limb postural reactions at similar latency. This study explored the role of vision in controlling the initial trajectory of grasping reactions triggered by sudden unpredictable medio-lateral platform translation. Healthy young adults were instructed to recover balance by grasping a marked section of a handrail, located to their right. To reinforce a dependence on arm reactions, movement of the feet was prevented by barriers. Liquid-crystal goggles were used to occlude vision during response initiation (200 ms interval starting at perturbation onset, PO). Results showed that the initial grasping trajectory (first 100 ms) and associated muscle activation were heavily modulated to take into account the direction and speed of the perturbation-induced body motion in relation to the handrail. This modulation was unaffected by occlusion of vision at PO, indicating that information about the rail location obtained prior to PO was incorporated into the control. These findings are consistent with the view that the CNS tunes the initial arm trajectory by combining an egocentric spatial map, formed prior to PO, with online feedback about the body motion from non-visual inputs (somatosensory and/or vestibular). This prevents potential delays associated with visual processing and ensures very rapid onset of arm movement that is directed appropriately even though the position of the body is perturbed unpredictably with respect to the target.

Adult↗

Single joint perturbation during gait: neuronal control of movement trajectory.

The aim of this study was to investigate the effect of single joint displacement on the pattern of leg muscle electromyographic (EMG) activity during locomotion. For the first time, unilateral rotational hip or knee joint displacements were applied by a driven orthotic device at three phases of swing during locomotion on a treadmill. The response pattern of bilateral leg muscle activation with respect to the timing and selection of muscles was almost identical for displacements of upper (hip joint) or lower (knee joint) leg. The leg muscle EMG responses were much stronger when the displacement was directed against the physiological movement trajectory, compared with when the displacement was reinforcing, especially during mid swing. It is suggested that these response patterns are designed to restore physiological movement trajectory rather than to correct a single joint position. Displacements released at initial or terminal swing, assisting or resisting the physiological movement trajectory, were followed by similar and rather unspecific response patterns. This was interpreted as being directed to stabilise body equilibrium.

Adult↗

Magnetoencephalographic signals predict movement trajectory in space.

Brain-machine interface (BMI) efforts have been focused on using either invasive implanted electrodes or training-extensive conscious manipulation of brain rhythms to control prosthetic devices. Here we demonstrate an excellent prediction of movement trajectory by real-time magnetoencephalography (MEG). Ten human subjects copied a pentagon for 45 s using an X-Y joystick while MEG signals were being recorded from 248 sensors. A linear summation of weighted contributions of the MEG signals yielded a predicted movement trajectory of high congruence to the actual trajectory (median correlation coefficient: r=0.91 and 0.97 for unsmoothed and smoothed predictions, respectively). This congruence was robust since it remained high in cross-validation analyses (based on the first half of data to predict the second half; median correlation coefficient: r=0.76 and 0.85 for unsmoothed and smoothed predictions, respectively).

Brain↗

Trajectory of contact region on the fingerpad gives the illusion of haptic shape.

When one explores a solid object with a fingertip, a contact region is usually defined. When the trajectory of this region on the fingerpad is artificially controlled so as to resemble the trajectory that is normally present while exploring a real object, the experience of shape is created. In order to generate appropriate local deformation trajectories, we built a servo-controlled mechanism that rolled a flat plate on the fingerpad during the manual exploration of virtual surfaces so that the plate was kept tangent to a virtual shape at the point of virtual contact. An experiment was then designed to test which mode of exploration maximized the shape information gain: active versus semi-active exploration, where semi-active exploration is when one hand touches passively and the other moves the target object, and the use of single versus multiple points of contact. We found that subjects were able to perform curvature discrimination at levels comparable to those achieved when using direct manual contact with real objects, and that the highly simplified stimulus provided by the device was a sufficient cue to give the illusion of touching three-dimensional surfaces.

Analysis of Variance↗

Head orientation is aligned with take-off trajectory as chicks jump.

Chicks of up to one week of age were video recorded while jumping across gaps of different widths and depths. The orientation of the head in the sagittal plane was measured at the point where jumps began. As the gap became wider, or the far side of it was raised, chicks' heads rotated upwards, but this change in head orientation did not keep the far edge of the gap fixated at a particular elevation in the visual field. Except with narrow gaps, which chicks often stepped over, there was a consistent relationship between head orientation and the initial trajectory of a jump. Each 1 degree change in trajectory was accompanied by an approximately 1 degree change in head orientation. It is proposed that the visual control of head orientation and position in relation to the dimensions of a gap contributes to adjusting a chick's posture and, therefore, its trajectory at take-off.

Animals↗