Search PubMed⌕ Search

Biomedical subjects

W P Medendorp

Publications and source records attributed to W P Medendorp.

16 recordsLinked to original sources
PubMed2005

Visuospatial memory computations during whole-body rotations in roll.

We used a memory-saccade task to test whether the location of a target, briefly presented before a whole-body rotation in roll, is stored in egocentric or in allocentric coordinates. To make this distinction, we exploited the fact that subjects, when tilted sideways in darkness, make systematic errors when indicating the direction of gravity (an allocentric task) even though they have a veridical percept of their self-orientation in space. We hypothesized that if spatial memory is coded allocentrically, these distortions affect the coding of remembered targets and their readout after a body rotation. Alternatively, if coding is egocentric, updating for body rotation becomes essential and errors in performance should be related to the amount of intervening rotation. Subjects (n = 6) were tested making saccades to remembered world-fixed targets after passive body tilts. Initial and final tilt angle ranged between -120 degrees CCW and 120 degrees CW. The results showed that subjects made large systematic directional errors in their saccades (up to 90 degrees ). These errors did not occur in the absence of intervening body rotation, ruling out a memory degradation effect. Regression analysis showed that the errors were closely related to the amount of subjective allocentric distortion at both the initial and final tilt angle, rather than to the amount of intervening rotation. We conclude that the brain uses an allocentric reference frame, possibly gravity-based, to code visuospatial memories during whole-body tilts. This supports the notion that the brain can define information in multiple frames of reference, depending on sensory inputs and task demands.

Adult↗
PubMed2004

Spatial transformations for eye-hand coordination.

Eye-hand coordination is complex because it involves the visual guidance of both the eyes and hands, while simultaneously using eye movements to optimize vision. Since only hand motion directly affects the external world, eye movements are the slave in this system. This eye-hand visuomotor system incorporates closed-loop visual feedback but here we focus on early feedforward mechanisms that allow primates to make spatially accurate reaches. First, we consider how the parietal cortex might store and update gaze-centered representations of reach targets during a sequence of gaze shifts and fixations. Recent evidence suggests that such representations might be compared with hand position signals within this early gaze-centered frame. However, the resulting motor error commands cannot be treated independently of their frame of origin or the frame of their destined motor command. Behavioral experiments show that the brain deals with the nonlinear aspects of such reference frame transformations, and incorporates internal models of the complex linkage geometry of the eye-head-shoulder system. These transformations are modeled as a series of vector displacement commands, rotated by eye and head orientation, and implemented between parietal and frontal cortex through efficient parallel neuronal architectures. Finally, we consider how this reach system might interact with the visually guided grasp system through both parallel and coordinated neural algorithms.

Animals↗
PubMed2003

Kinematic rules for upper and lower arm contributions to grasp orientation.

The purpose of the current study was to investigate the contribution of upper and lower arm torsion to grasp orientation during a reaching and grasping movement. In particular, we examined how the visuomotor system deals with the conflicting demands of coordinating upper and lower arm torsion and maintaining Donders' Law of the upper arm (a behavioral restriction of the axes of arm rotation to a two-dimensional "surface"). In experiment 1, subjects reached out and grasped a target block that was presented in one of 19 orientations (5 degrees clockwise increments from horizontal to vertical) at one position in a vertical presentation board. In experiment 2, target blocks were presented in one of three orientations (horizontal, three-quarter, and vertical) at nine different positions in the presentation board. If reach and grasp commands control the proximal and distal arms separately, then one would only expect the lower arm to contribute to grasp orientations and that Donders' Law would hold for the upper arm-independent of grasp orientations. Instead, as the required grasp orientation increased from horizontal to vertical, there was a significant clockwise torsional rotation in the upper arm, which accounted for 9% of the final vertical grasp orientation, and the lower arm, which accounted for 42%. A linear relationship existed between the torsional rotations of the upper and lower arm, which indicates that the components of the arm rotate in coordination with one another. The location-dependent aspects of upper and lower arm torsion remained invariant, however, yielding consistently shaped Donders' "surfaces" (with different torsional offsets) for different grasp orientations. These observations suggest that the entire arm-hand system contributes to grasp orientation, and therefore, the reach/grasp distinction is not directly reflected in proximal-distal kinematics but is better reflected in the distinction between these coordinated orienting rules and the location-dependent kinematic rules for the upper arm that result in Donders' Law for one given grasp orientation.

Adult↗
PubMed2003

Geometric computations underlying eye-hand coordination: orientations of the two eyes and the head.

Eye-hand coordination is geometrically complex. To compute the location of a visual target relative to the hand, the brain must consider every anatomical link in the chain from retinas to fingertips. Here we focus on the first three links, studying how the brain handles information about the angles of the two eyes and the head. It is known that people, even in darkness, reach more accurately when the eye looks toward the target, rather than right or left of it. We show that reaching is also impaired when the binocular fixation point is displaced from the target in depth: reaching becomes not just sloppy, but systematically inaccurate. Surprisingly, though, in normal Gaze-On-Target reaching we found no strong correlations between errors in aiming the eyes and hand onto the target site. We also asked people to reach when the head was not facing the target. When the eyes were on-target, people reached accurately, but when gaze was off-target, performance degraded. Taking all these findings together, we suggest that the brain's computational networks have learned the complex geometry of reaching for well-practiced tasks, but that the networks are poorly calibrated for less common tasks such as Gaze-Off-Target reaching.

Adult↗
PubMed2003

Ocular kinematics and eye-hand coordination.

Eye-hand coordination is complicated by the fact that the eyes are constantly in motion relative to the head. This poses problems in interpreting the spatial information gathered from the retinas and using this to guide hand motion. In particular, eye-centered visual information must somehow be spatially updated across eye movements to be useful for future actions, and these representations must then be transformed into commands appropriate for arm motion. In this review, we present evidence that early visuomotor representations for arm movement are remapped relative to the gaze direction during each saccade. We find that this mechanism holds for targets in both far and near visual space. We then show how the brain incorporates the three-dimensional, rotary geometry of the eyes when interpreting retinal images and transforming these into commands for arm movement. Next, we explore the possibility that hand-eye alignment is optimized for the eye with the best field of view. Finally, we describe how head orientation influences the linkage between oculocentric visual frames and bodycentric motor frames. These findings are framed in terms of our 'conversion-on-demand' model, in which only those representations selected for action are put through the complex visuomotor transformations required for interaction with objects in personal space, thus providing a virtual on-line map of visuomotor space.

Biomechanical Phenomena↗
PubMed2002

Visuomotor transformations for eye-hand coordination.

In recent years the scientific community has come to appreciate that the early cortical representations for visually guided arm movements are probably coded in a visual frame, i.e. relative to retinal landmarks. While this scheme accounts for many behavioral and neurophysiological observations, it also poses certain problems for manual control. For example, how are these oculocentric representations updated across eye movements, and how are they then transformed into useful commands for accurate movements of the arm relative to the body? Also, since we have two eyes, which is used as the reference point in eye-hand alignment tasks like pointing? We show that patterns of errors in human pointing suggest that early oculocentric representations for arm movement are remapped relative to the gaze direction during each saccade. To then transform these oculocentric representations into useful commands for accurate movements of the arm relative to the body, the brain correctly incorporates the three-dimensional, rotary geometry of the eyes when interpreting retinal images. We also explore the possibility that the eye-hand coordination system uses a strategy like ocular dominance, but switches alignment between the left and right eye in order to maximize eye-hand coordination in the best field of view. Finally, we describe the influence of eye position on eye-hand alignment, and then consider how head orientation influences the linkage between oculocentric visual frames and bodycentric motor frames. These findings are framed in terms of our 'conversion-on-demand' model, which suggests a virtual representation of egocentric space, i.e. one in which only those representations selected for action are put through the complex visuomotor transformations required for interaction with actual objects in personal space.

Arm↗
PubMed2002

Human gaze stabilization during active head translations.

This study investigated how binocular gaze is controlled to compensate for self-generated translational movements of the head where geometric requirements dictate that the ideal gaze signal needs to be modulated by the inverse of target distance. Binocular gaze (eye plus head) was measured for visual and remembered targets at various distances in six human subjects during active head translations at frequencies of 0.25, 0.5, 1.0, and 1.5 Hz. We found that, during head translations, gaze changes were achieved by a combination of eye and head rotations. Accordingly, stabilization performance was characterized by the gaze response parameters sensitivity and phase, where sensitivity is defined as the ratio of gaze velocity and translational eye velocity and where phase refers to the phase delay of gaze velocity relative to translational eye velocity. In the analysis, we used a binocular coordinate system yielding a version and a vergence component. We examined how frequency and target distance, estimated from the vergence angle, affected sensitivity and phase of the version component of the gaze signal and compared the results to the requirements for ideal performance. The relation between gaze sensitivity and the inverse of distance was characterized by a linear regression analysis. The ratio of the slope of the linear regression and the slope required for ideal stabilization provided a measure for the degree of "distance compensation." The results show that distance compensation was better for a visual target than for remembered targets in darkness, and behaved according to low-pass characteristics in both target conditions. It declined from 1.00 to 0.84 for visual targets and from 0.87 to 0.57 for remembered targets in the frequency range 0.25-1.5 Hz. The intercept obtained from the regression yielded the gaze response at zero vergence and specified a "default sensitivity" of gaze compensation. Default sensitivity increased with frequency from 0.02 at 0.25 Hz to 0.10 degrees/cm at 1.5 Hz for visual targets and from 0.04 to 0.16 degrees/cm in darkness. The phase delays of the gaze response increased on average from 2 to 7 degrees in the frequency range 0.25-1.5 Hz. In comparison with earlier passive studies, active translation compensation in the dark is superior at all frequencies where comparison was possible. We conclude that a nonvestibular signal with low-pass characteristics contributes to gaze during active head translations.

Adult↗
PubMed2001

Three-dimensional head and upper arm orientations during kinematically redundant movements and at rest.

The three rotational degrees of freedom of the head and the upper arm exceed the number needed in a two-dimensional (2-D) facing or pointing task, respectively. Previous studies reported a reduction of the number of degrees of freedom from three to two, with one degree of freedom being a unique function of the other two (Donders' law). This study investigated whether three-dimensional (3-D) orientations of the head and arm are the same at rest and during movement for corresponding pointing or facing directions. Two separate experiments were performed: one focused on head orientations, the other focused on upper arm orientations. We instructed subjects to direct the nose or to point the extended arm in the direction of targets, which appeared in a quasi-random order at 2-s intervals. The head and upper arm orientations at rest were described by a 2-D surface with a scatter less than 3 or 4 degrees, respectively. Both for the arm and the head, orientations started and ended near the 2-D surface, but for a number of the target pairs, the orientations deviated from those predicted by the 2-D surface during movement in a way that was consistent and reproducible for movements between each target pair. For upper arm movements, we often found that deviations of arm orientations from the 2-D surface increased with increasing movement velocity. Such a positive correlation between deviation and movement velocity was not found for head movements. These results clearly indicate violations of Donders' law during movement and argue against several models for movement control found in the literature.

Adult↗
PubMed2001

The subjective vertical and the sense of self orientation during active body tilt.

Previous testing of the ability to set a luminous line to the direction of gravity in passively-tilted subjects, in darkness, has revealed a remarkable pattern of systematic errors at tilts beyond 60 degrees, as if body tilt is undercompensated or underestimated (Aubert or A-effect). We investigated whether these consistent deviations from orientation constancy can be avoided during active body tilt, where more potential cues about body tilt (e.g. proprioception and efference copy) are available. The effects of active body tilt on the subjective vertical and on the perception of self tilt were studied in six subjects. After adopting a laterally-tilted posture, while standing in a dark room, they indicated the subjective vertical by adjusting a visual line and gave their verbal estimate of head orientation, expressed on a clock scale. Head roll tilts covered the range from -150 degrees to +150 degrees. The subjective vertical results showed no sign of improvement. Actively-tilted subjects still exhibited the same pattern of systematic errors that characterised their performance during passive tilt. Random errors in this task showed a steep monotonic increase with tilt angle, as in earlier passive tilt experiments. By contrast, verbal head-tilt estimates in the active experiments showed a clear improvement and were now almost devoid of systematic errors, but the noise level remained high. Various models are discussed in an attempt to clarify how these task-related differences and the selective improvement of the self-tilt estimates in the active experiments may have come about.

Adult↗
PubMed2000

Kinematic strategies for upper arm-forearm coordination in three dimensions.

This study addressed the question of how the three-dimensional (3-D) control strategy for the upper arm depends on what the forearm is doing. Subjects were instructed to point a laser-attached in line with the upper arm-toward various visual targets, such that two-dimensional (2-D) pointing directions of the upper arm were held constant across different tasks. For each such task, subjects maintained one of several static upper arm-forearm configurations, i. e., each with a set elbow angle and forearm orientation. Upper arm, forearm, and eye orientations were measured with the use of 3-D search coils. The results confirmed that Donders' law (a behavioral restriction of 3-D orientation vectors to a 2-D "surface") does not hold across all pointing tasks, i.e., for a given pointing target, upper arm torsion varied widely. However, for any one static elbow configuration, torsional variance was considerably reduced and was independent of previous arm position, resulting in a thin, Donders-like surface of orientation vectors. More importantly, the shape of this surface (which describes upper arm torsion as a function of its 2-D pointing direction) depended on both elbow angle and forearm orientation. For pointing with the arm fully extended or with the elbow flexed in the horizontal plane, a Listing's-law-like strategy was observed, minimizing shoulder rotations to and from center at the cost of position-dependent tilts in the forearm. In contrast, when the arm was bent in the vertical plane, the surface of best fit showed a Fick-like twist that increased continuously as a function of static elbow flexion, thereby reducing position-dependent tilts of the forearm with respect to gravity. In each case, the torsional variance from these surfaces remained constant, suggesting that Donders' law was obeyed equally well for each task condition. Further experiments established that these kinematic rules were independent of gaze direction and eye orientation, suggesting that Donders' law of the arm does not coordinate with Listing's law for the eye. These results revive the idea that Donders' law is an important governing principle for the control of arm movements but also suggest that its various forms may only be limited manifestations of a more general set of context-dependent kinematic rules. We propose that these rules are implemented by neural velocity commands arising as a function of initial arm orientation and desired pointing direction, calculated such that the torsional orientation of the upper arm is implicitly coordinated with desired forearm posture.

Biomechanical Phenomena↗
PubMed2000

Context compensation in the vestibuloocular reflex during active head rotations.

The vestibuloocular reflex (VOR) needs to modulate its gain depending on target distance to prevent retinal slip during head movements. We investigated gain modulation (context compensation) for binocular gaze stabilization in human subjects during voluntary yaw and pitch head rotations. Movements of each eye were recorded, both when attempting to maintain gaze on a small visual target at straight-ahead in a darkened room and after its disappearance (remembered target). In the analysis, we relied on a binocular coordinate system yielding a version and a vergence component. We examined how frequency and target distance, approached here by using vergence angle, affected the gain and phase of the version component of the VOR and compared the results to the requirements for ideal performance. Linear regression analysis on the version gain-vergence relationship yielded a slope representing the influence of target proximity and an intercept corresponding to the response at zero vergence ("default gain"). The slope of the fitted relationship, divided by the geometrically required slope, provided a measure for the quality of version context compensation ("context gain"). In both yaw and pitch experiments, we found default version gains close to one even for the remembered target condition, indicating that the active VOR for far targets is already close to ideal without visual support. In near target experiments, the presence of visual feedback yielded near unity context gains, indicating close to optimal performance (retinal slip <0.4 degrees /s). For remembered targets, the context gain deteriorated but was still superior to performance in corresponding passive studies reported in the literature. In general, context compensation in the remembered target paradigm was better for vertical than for horizontal head rotations. The phase delay of version eye velocity relative to head velocity was small (approximately 2 degrees) for both horizontal and vertical head movements. Analysis of the vergence data from the near target experiments showed that context compensation took into account that the two eyes require slightly different VORs. In the DISCUSSION, comparison of the present default VOR gains and context gains with data from earlier passive studies has led us to propose a limited role for efference copies during self-generated movements. We also discuss how our analysis can provide a framework for evaluating two different hypotheses for the generation of binocular VOR eye movements.

Adult↗
PubMed1999

Pointing to remembered visual targets after active one-step self-displacements within reaching space.

We studied pointing movements to remembered visual targets in a completely darkened room with and without self-made step movements in order to investigate in which coordinate system and to what extent target representations relative to the body are updated for self-induced egomotion. A small red-light-emitting diode on the fingertip provided visual feedback about fingertip position at all times. We asked subjects to make pointing movements that started 2 s after disappearance of a visual target. In this interval of 2 s the subject did or did not make a step. The pointing errors without a step showed that subjects undershot faraway targets in a systematic way, whereas they sometimes overshot nearby targets. We found that the step causes larger pointing errors both in amplitude and direction with a bias in the direction of the step. We explored three different versions of a descriptive model in which polar coordinates were used to describe the pointing movement, and in which either Cartesian or polar coordinates were used to update target position relative to the shoulder for the step. The results suggest that incorporation of the step displacement in the new target position relative to the subject is done in a Cartesian frame of reference. Moreover, the amplitude of the step displacement tends to be underestimated by subjects.

Adult↗
PubMed1999

Donders' law in torticollis.

We investigated head movements of patients with spasmodic torticollis toward targets in various directions. These patients, whose severe dystonia was reflected in an abnormal resting head position, appeared to retain a Donders'-type strategy for the control of the rotational degrees of freedom of the head. As in normals, rotation vectors, representing head orientation, were confined to a curved surface, which specifies how head torsion depends on gaze direction. The orientation of the surface in body coordinates, which was very stereotyped in normals, was different for patients. The same Donders surface was found for head movements and for stationary head postures, indicating that the same neural mechanism governs its implementation in both tasks. To interpret our results, we propose a conceptual scheme incorporating the basal ganglia, which are thought to be involved in the etiology of torticollis, and an implementation stage for Donders' law.

Aged↗
PubMed1998

Off-centric rotation axes in natural head movements: implications for vestibular reafference and kinematic redundancy.

Until now, most studies concerning active head movements in three dimensions have used the classical rotation vector description. Although this description yields both the orientation of the head rotation axis and the amount of rotation, it is incomplete because it cannot specify the location of this rotation axis in space. The latter is of importance for a proper picture of the vestibular consequences of active head movements and has relevance for the problem of how the brain deals with the inherent kinematic redundancy of the multijoint head-neck system. With this in mind, we have extended the rotation vector description by applying the helical axes approach, which yields both the classical rotation vector as well as the location of the rotation axis in space. Subjects (n = 7), whose head movements were recorded optically, were instructed to shift gaze naturally to targets in 12 different directions at an eccentricity of 40 degrees. The results demonstrate that the axes for these head movements occupy consistently different spatial locations. For purely horizontal movements, the rotation axis is located near a point midway between the two ear canals. For gaze shifts in other directions, the rotation axes are located below the ear canals along two circles, one for movements with an upward component (up circle), the other (typically larger in size) for movements with a downward component (down circle). Purely vertical movement (up and down) axes were located on the lower pole of the up and down circles, respectively. It was found that both circles, the upper poles of which coincided, became larger in size as movement amplitude increased, which means that the axis location shifts to lower and more eccentric locations with respect to the skull for larger flexion and extension movements. Although this pattern could be recognized in most subjects, there were consistent intersubject differences in the absolute size of the circles, their increase with movement amplitude, and in the relative sizes of the up and down circles. Because multiple vertebrae are involved in head movements, there are theoretically many possibilities to execute a certain head movement. The differences in circle patterns among subjects indicate different strategies in resolving this kinematic redundancy problem, a fact that was not apparent from the classical rotation vector part of our description, which yielded a rather uniform picture. A simple model suggests that the downward shift of the location of the rotation axis requires a modulation in vestibulo-ocular reflex gain of </=10% to maintain fixation of a near target during vertical head movement. The involvement of the otolith system in this process remains to be determined.

Adult↗
PubMed1997

Motor unit firing behavior in human arm flexor muscles during sinusoidal isometric contractions and movements.

Simultaneous recordings of action potentials (APs) of multiple single motor units (MUs) were obtained in brachialis and biceps (caput breve) muscles during sinusoidally modulated isometric contractions of elbow flexor muscles and during sinusoidal flexion/extension movements in the elbow against a preload in the extension direction. The results show that MUs typically fire in one short burst for each sinusoidal cycle. The mean phase lead of the bursts of APs relative to a sinusoidally modulated isometric torque in the elbow joint or relative to sinusoidal movements in the elbow increases gradually with frequency. The increase of the mean phase lead during isometric contractions was very similar for all MUs and could be explained well by modeling the force production of MUs with a second-order linear low-pass system. For sinusoidal flexion/extension movements each MU reveals a specific, reproducible phase lead as a function of frequency. However, there is a large variability in phase behavior between MUs. Also, the modulation of the firing rate for sinusoidal isometric contractions versus sinusoidal movements appeared to be different for various MUs. In simultaneous recordings some MUs clearly revealed a larger firing rate in each burst for movements relative to isometric contractions, whereas other MUs revealed a smaller firing rate. This suggests that some MUs are preferentially activated during movements whereas others are preferably activated during isometric contractions. The results demonstrate task-dependent changes in the relative activation of MUs within a single muscle for sinusoidal isometric contractions and movements.

Action Potentials↗