Behavioral reference frames for planning human reaching movements

At some stage in the process of a sensorimotor transformation for a reaching movement, information about the current position of the hand and information about the location of the target must be encoded in the same frame of reference to compute the hand-to-target difference vector. Two main hypotheses have been proposed regarding this reference frame: an eye-centered and a body-centered frame. Here we evaluated these hypotheses using the pointing errors that subjects made when planning and executing arm movements to memorized targets starting from various initial hand positions while keeping gaze fixed in various directions. One group of subjects (n = 10) was tested without visual information about hand position during movement planning (unseen-hand condition); another group (n = 8) was tested with hand and target position simultaneously visible before movement onset (seen-hand condition). We found that both initial hand position and gaze fixation direction had a significant effect on the magnitude and direction of the pointing error. Errors were significantly smaller in the seen-hand condition. For both conditions, though, a reference frame analysis showed that the errors arose at an eye- or hand-centered stage or both, but not at a body-centered stage. As a common reference frame is required to specify a movement vector, these results suggest that an eye-centered mechanism is involved in integrating target and hand position in programming reaching movements. We discuss how simple gain elements modulating the eye-centered target and hand-position signals can account for these results.     

Separate adaptive mechanisms for controlling trajectory and final position in reaching

We examined control of the hand's trajectory (direction and shape) and final equilibrium position in horizontal planar arm movements by quantifying transfer of learned visuomotor rotations between two tasks that required aiming the hand to the same spatial targets. In a trajectory-reversal task ("slicing"), the hand reversed direction within the target and returned to the origin. In a positioning task ("reaching"), subjects moved the hand to the target and held it there; cursor feedback was provided only after movement ended to isolate learning of final position from trajectory direction. We asked whether learning acquired in one task would transfer to the other. Transfer would suggest that the hand's entire trajectory, including its endpoint, was controlled using a common spatial plan. Instead we found minimal transfer, suggesting that the brain used different representations of target position to specify the hand's initial trajectory and its final stabilized position. We also observed asymmetrical practice effects on hand trajectory, including systematic curvature of reaches made after rotation training and hypermetria of untrained slice reversals after reach training. These are difficult to explain with a unified control model, but were replicated in computer simulations that specified the hand's initial trajectory and its final equilibrium position. Our results suggest that the brain uses different mechanisms to plan the hand's initial trajectory and final position in point-to-point movements, that it implements these control actions sequentially, and that trajectory planning does not account for specific impedance values to be implemented about the final stabilized posture.     

Action, perception and postural planning when reaching for tools

The dorsal and ventral streams model of action and perception suggests that reaching to grasp a tool for use involves integrated operation of the two streams. Few attempts have been made to test the limits of this integration in normal subjects. Twenty normal subjects reached for tools or geometric objects which were rotated rapidly during reaching or immediately beforehand. In a first experiment it was shown that reaching for an inverted tool was slower than reaching for objects which required hand inversion due to proximity to a physical barrier. Also, for the right hand, tool rotation during reaching provoked a higher incidence of hand rotation in the wrong direction than did rotation of objects. In a second similar experiment, hand inversion when grasping objects was induced by the need to plan a future action rather than by proximity of a physical barrier. Despite this balancing of complexity of postural planning for tools and objects, hand rotation errors for both hands were more common for tools than objects. This was consistent with the two-stream model in suggesting that there was a process which produced rapid online tracking of stimulus rotation and this had to be overcome by a slower process which dictated grasping in accordance with knowledge of tool use.     

Reference frames for reach planning in macaque dorsal premotor cortex

When a human or animal reaches out to grasp an object, the brain rapidly computes a pattern of muscular contractions that can acquire the target. This computation involves a reference frame transformation because the target's position is initially available only in a visual reference frame, yet the required control signal is a set of commands to the musculature. One of the core brain areas involved in visually guided reaching is the dorsal aspect of the premotor cortex (PMd). Using chronically implanted electrode arrays in two Rhesus monkeys, we studied the contributions of PMd to the reference frame transformation for reaching. PMd neurons are influenced by the locations of reach targets relative to both the arm and the eyes. Some neurons encode reach goals using limb-centered reference frames, whereas others employ eye-centered reference fames. Some cells encode reach goals in a reference frame best described by the combined position of the eyes and hand. In addition to neurons like these where a reference frame could be identified, PMd also contains cells that are influenced by both the eye- and limb-centered locations of reach goals but for which a distinct reference frame could not be determined. We propose two interpretations for these neurons. First, they may encode reach goals using a reference frame we did not investigate, such as intrinsic reference frames. Second, they may not be adequately characterized by any reference frame.     

Contribution of reference frames for movement planning in peripersonal space representation

The principal goal of our study is to gain an insight into the representation of peripersonal space. Two different experiments were conducted in this study. In the first experiment, subjects were asked to represent principal anatomical reference planes by drawing ellipses in the sagittal, frontal and horizontal planes. The three-dimensional hand-drawing movements, which were achieved with and without visual guidance, were considered as the expression of a cognitive process per se: the peripersonal space representation for action. We measured errors in the spatial orientation of ellipses with regard to the requested reference planes. For ellipses drawn without visual guidance, with eyes open and eyes closed, orientation errors were related to the reference planes. Errors were minimal for sagittal and maximal for horizontal plane. These disparities in errors were considerably reduced when subjects drew using a visual guide. These findings imply that different planes are centrally represented, and are characterized, by different errors when subjects use a body-centered frame for performing the movement and suggest that the representation of peripersonal space may be anisotropic. However, this representation can be modified when subjects use an environment-centered reference frame to produce the movement. In the second experiment, subjects were instructed to represent, with eyes open and eyes closed, sagittal, frontal and horizontal planes by pointing to virtual targets located in these planes. Disparities in orientation errors measured for pointing were similar to those found for drawing, implying that the sensorimotor representation of reference planes was not constrained by the type of motor tasks. Moreover, arm postures measured at pointing endpoints and at comparable spatial locations in drawing are strongly correlated. These results suggest that similar patterns of errors and arm posture correlation, for drawing and pointing, can be the consequence of using a common space representation and reference frame. These findings are consistent with the assumption of an anisotropic action-related representation of peripersonal space when the movement is performed in a body-centered frame.     

Auditory saccades from different eye positions in the monkey: implications for coordinate transformations

Auditory spatial information arises in a head-centered coordinate frame, whereas the saccade command signals generated by the superior colliculus (SC) are thought to specify target locations in an eye-centered frame. However, auditory activity in the SC appears to be neither head- nor eye-centered but in a reference frame that is intermediate between both of these reference frames. This neurophysiological finding suggests that auditory saccades might not fully compensate for changes in initial eye position. Here, we investigated whether the accuracy of saccades to sounds is affected by initial eye position in rhesus monkeys. We found that, on average, a 12 degrees horizontal shift in initial eye position produced only a 0.6 to 1.6 degrees horizontal shift in the endpoints of auditory saccades made to targets at a range of locations along the horizontal meridian. This shift was similar in size to the modest influence of eye position on visual saccades. This virtually complete compensation for initial eye position implies that auditory activity in the SC is read out in a manner that is appropriate for generating accurate saccades to sounds.     

Autoactivation of source dwell positions for HDR brachytherapy treatment planning

The most accurate classical dose optimization algorithms in HDR brachytherapy strongly depend on an appropriate selection of source dwell positions which fulfill user-defined geometrical boundary conditions which are relative to patient anatomy. Most anatomical situations, such as for prostate and head and neck tumors, are complex and can require geometries with 5-15 catheters with 48 possible dwell positions per catheter depending on the tumor volume. The manual selection of dwell positions using visual checks by trial and error is very time consuming. This can only be improved by the use of a technique which automatically recognizes and selects the optimum dwell positions for each catheter. We have developed an algorithm, termed an autoactivation algorithm, which improves implant planning by providing a facility for the necessary automatic recognition of HDR source dwell positions.     

Proprioceptive guidance and motor planning of reaching movements to unseen targets

The ability to make accurate reaching movements toward proprioceptively defined target locations was studied in seven normal subjects who were trained to reach to five different targets in a horizontal plane, with no vision of hand or target. The task consisted of moving a handle from a fixed origin to each target location, fast and accurately. Target locations were learned in training sessions that utilized acoustic cuing. Most movements were rapid, with a bell-shaped velocity profile. The error in target reproduction, which constituted the difference between the position consciously identified as the correct target location and the real target location, was calculated in each trial. This was compared with the error in preprogrammed reaching, which constituted the difference between the point in space where the initial fast movement toward the target ended and the target location. The absence of significant differences between these two error types indicated that the transformation from an internal representation of target location into a motor program for reaching to it did not introduce an additional reaching error. Learning of target locations was done only with the right hand, yet, reaching of both hands was tested. This allowed a comparison between the subjects' ability to utilize a transformed spatial code (reaching with the untrained hand) and their ability to use a direct sensory-motor code (reaching with the trained hand). While transformation of the spatial code was found to reduce it's accuracy, utilization of this code in motor programming again did not appear to introduce an additional error.     

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° or 90°), 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° to 150° 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. Received: 6 August 1996 / Accepted: 18 December 1996     

Reconstruction of applicator positions from multiple-view images for accurate superficial hyperthermia treatment planning

In the current clinical practice, prior to superficial hyperthermia treatments (HT), temperature probes are placed in tissue to document a thermal dose. To investigate whether the painful procedure of catheter placement can be replaced by superficial HT planning, we study if the specific absorption rate (SAR) coverage is predictive for treatment outcome. An absolute requirement for such a study is the accurate reconstruction of the applicator setup. The purpose of this study was to investigate the feasibility of the applicator setup reconstruction from multiple-view images. The accuracy of the multiple-view reconstruction method has been assessed for two experimental setups using six lucite cone applicators (LCAs) representing the largest array applied at our clinic and also the most difficult scenario for the reconstruction. For the two experimental setups and 112 distances, the mean difference between photogrametry reconstructed and manually measured distances was 0.25 ± 0.79?mm (mean±1 standard deviation). By a parameter study of translation T (mm) and rotation R (°) of LCAs, we showed that these inaccuracies are clinically acceptable, i.e. they are either from ±1.02 mm error in translation or ±0.48° in rotation, or combinations expressed by 4.35R(2) + 0.97T(2) = 1. We anticipate that such small errors will not have a relevant influence on the SAR distribution in the treated region. The clinical applicability of the procedure is shown on a patient with a breast cancer recurrence treated with reirradiation plus superficial hyperthermia using the six-LCA array. The total reconstruction procedure of six LCAs from a set of ten photos currently takes around 1.5 h. We conclude that the reconstruction of superficial HT setup from multiple-view images is feasible and only minor errors are found that will have a negligible influence on treatment planning quality.     

Manual asymmetries in the directional coding of reaching: further evidence for hemispatial effects and right hemisphere dominance for movement planning

Directional coding of hand movements is of primary importance in the proactive control of goal-directed aiming. At the same time, manual reaction times are known to be asymmetric when reaching at lateralized targets. Generally, ipsilateral movements and left hand advantages are interpreted using the classical model of interhemispheric transmission for simple visuomotor integration, but the use of this model was recently challenged when applied to reaching movements, arguing that attentional and biomechanical effects could also account for such asymmetries. In this work, we aimed at controlling both visual attention orienting and movement mechanical constraints in order to clarify the origin of manual reaction time asymmetries and hemispatial effects in the directional coding of reaching. Choice reaction time pointing tasks were assessed in two experiments in which identical movements were compared in different conditions of target lateralization and different conditions of head, eye and hand position. Results suggested that biomechanical constraints could account for hemispatial effects for movement execution but not for movement direction coding. These results are discussed in the light of models of interhemispheric cooperation and the right hemisphere dominance for spatial processing. Electronic Publication     

Involvement of ipsilateral parieto-occipital cortex in the planning of reaching movements: Evidence by TMS

Involvement of the ipsilateral hemisphere during planning of reaching movements is still matter of debate. While it has been demonstrated that the contralateral hemisphere is dominant in visuo-motor integration, involvement of the ipsilateral hemisphere has also been proposed. Furthermore, a dominant role for left posterior parietal cortex has been shown in this process, independently of the hand and visual field involved. In this study, the possible involvement of ipsilateral parieto-occipital cortex in planning of reaching movements was investigated by transcranial magnetic stimulation (TMS). TMS was applied on four points of the parietal and occipital cortex at 50% (Time 1), 75% (Time 2) and 90% (Time 3) of reaction time from a go-signal to hand movement. The only effect observed was an increase in reaction time when a region around the parieto-occipital junction was stimulated at Time 2. These results provide further support to the hypothesis that, in the posterior parietal cortex, planning of reaching movements also relies on the ipsilateral hemisphere, in addition to the contralateral or dominant one.     

Perception of maximum reaching height reflects impending changes in reaching ability and improvements transfer to unpracticed reaching tasks

Perception of whether a given behavior is possible typically reflects a person’s action capabilities even before the behavior is performed and even when the person has undergone recent changes to their action capabilities. Importantly, perception of affordances for a given behavior also reflects impending changes to action capabilities. Two experiments investigated perception of affordances for reaching when the means of reaching would bring about changes in reaching ability. Experiment 1 found that perception of maximum reaching height is relative to impending changes brought on by a change in posture and use of a hand-held tool. Experiment 2 found that practice performing a reaching task in which reaching ability is unchanged is sufficient to bring about improvements in perception of maximum reaching height when the means of reaching would change reaching ability. The results are discussed in the context of the prospectivity of perception of affordances, and the experiences that may be necessary to bring about (transfer of) improvements in perception of affordances.     

Control of neural prostheses for grasping and reaching

In recent years several neural prostheses have been developed and tested as orthoses or as therapeutic systems for hemiplegic and tetraplegic subjects aiming to improve the upper extremities function. The use of neural prostheses demonstrated that the targeted group of subjects could significantly benefit from functional electrical stimulation that is integrated in goal directed movements. In this paper the control for neural prostheses is explained using available systems that apply either surface or implantable interfaces to sensory-motor systems. Further more, a new strategy that has been tested for control of reaching and grasping within a neural prosthesis especially designed for neurorehabilitation is described. This, so-called, coordination strategy was based on mimicking the output space model of natural control determined in reach/grasp/release movements of healthy humans.     

Target selection for visually guided reaching in macaque

We examined target selection for visually guided reaching in monkeys using a visual search task in which an odd-colored target was presented with distractors. The colors of the target and distractors were randomly switched in each trial between red and green, and the number of distractors was varied. Previous studies of saccades and attention have shown that target selection in this task is easier when a greater number of homogenous distractors is present. We found that monkeys made fewer reaches to distractors and that reaches to the target were completed more quickly when a greater number of homogenous distractors was present. When the target was presented in a sparse array of distractors, reaches had longer movement durations and greater trajectory curvature. Reaching errors were directed more often to a distractor adjacent to the target, suggesting a spatially coarse-to-fine progression during target selection. Reaches were also influenced by the properties of trials in the recent past. When the colors of the target and distractors remained the same from trial to trial rather than switching, reaches were completed more quickly and accurately, indicating that color priming across trials facilitates target selection. Moreover, when difficult search trials were randomly intermixed with easier trials without distractors, reach latencies were influenced by the difficulty of previous trials, indicating that motor initiation strategies are gradually adjusted based on accumulated experience. Overall, these results are consistent with reaching results in humans, indicating that the monkey provides a sound model for understanding the neural underpinnings of reach target selection.     

Kinematics of reaching and implications for handedness in rhesus monkey infants

Kinematic studies of reaching in human infants using two-dimensional (2-D) and three-dimensional (3-D) recordings have complemented behavioral studies of infant handedness by providing additional evidence of early right asymmetries. Right hand reaches have been reported to be straighter and smoother than left hand reaches during the first year. Although reaching has been a popular measure of handedness in primates, there has been no systematic comparison of left and right hand reach kinematics. We investigated reaching in infant rhesus monkeys using the 2-D motion analysis software MaxTRAQ Lite+ (Innovision Systems). Linear mixed-effects models revealed that left hand reaches were smoother, but not straighter, than right hand reaches. An early left bias matches previous findings of a left hand preference for reaching in adult rhesus monkeys. Additional work using this kind of kinematic approach will extend our understanding of primate handedness beyond traditional studies measuring only frequency or bouts of hand use.     

Infants and adults reaching in the dark

It has been shown that infants over the age of 6 months will reach for an object in complete darkness. This experiment measured the reaching movements of 9- to 16-month-old infants and adults under several different conditions of illumination to investigate the role of vision and stored visual representations in reach control. In one condition, participants reached for an object with the lights on. In a second condition, participants reached for an object glowing in the dark (glowing condition). This allowed us to measure the effects of vision of the arm and vision of the reach space. We also looked at the effect of removing vision of the object on reach control: in the final two conditions, participants reached for an object in complete darkness (0-s dark) and in complete darkness after a 4-s delay (4-s dark). We compared the kinematics of a reach (e.g. average speed, reach straightness) between the four illumination conditions. The results showed that infants reached faster and decelerated for a shorter period of time in the dark (0- and 4-s dark) than in the light. By comparison, adults reached slower and decelerated for a longer period of time in the dark (0- and 4-s dark) than in the light. We did not find any effect of the glowing condition compared to full vision on infant reaching movements. These results suggest that infant reaching movements only become compromised when the target is not visible, whereas vision of the hand and the reach space are less significant. Without online visual feedback, an infant reach in the dark appears to be driven by feedforward mechanisms and control may be affected by an immature ability to form and/or retain visual spatial memory.     

Basic features of phasic activation for reaching in vertical planes

The purpose of this study was to fully characterize the timing and intensity of the phasic portion of the electromyographic (EMG) waveform for reaching movements in vertical planes. Electromyographic activity was simultaneously recorded from nine superficial elbow and/or shoulder muscles while human subjects made rapid arm movements. Hand paths comprised 20 directions in a sagittal plane and 20 directions in a frontal plane. In order to focus on the more phasic aspects of muscle activation, estimates of postural EMG activity were subtracted from the EMG traces recorded during rapid reaches. These postural estimates were obtained from activity recorded during very slow reaches to the same targets. After subtraction of this postural activity, agonist or antagonist burst patterns were often observed in the phasic EMG traces. For nearly all muscles and all subjects, the relation between phasic EMG intensity and movement direction was a function with multiple peaks. For all muscles, the timing of phasic EMG bursts varied as a function of movement direction: the data from each muscle exhibited a gradual temporal shift of activity over a certain range of directions. This gradual temporal shift has no obvious correspondence to the mechanical requirements of the task and might represent a neuromuscular control strategy in which burst timing contributes to the specification of movement direction.     

Visually guided reaching for food in binocularly deprived cats.

Four visually deprived cats and four controls were tested in reaching for food from a cage. In each trial the animals in the groups responded immediately after unscreening the cage. However, in reaction time of forelimb extension the deprived group showed proportionally fewer responses with shorter reaction time than the control group.