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Biomedical subjects

Martha Flanders

Publications and source records attributed to Martha Flanders.

At least 19 recordsLinked to original sources

Timing of muscle activation in a hand movement sequence.

Recent studies have described muscle synergies as overlapping, multimuscle groups defined by synchronous covariation in activation intensity. A different approach regards a synergy as a fixed temporal sequence of bursts of activity across groups of motoneurons. To pursue this latter definition, the present study used a principal component (PC) analysis tailored to reveal the across-muscle temporal synergies of human hand movement. Electromyographic (EMG) activity was recorded as subjects used a manual alphabet to spell a list of words. The analysis was applied to the EMG waveforms from 27 letter-to-letter transitions of equal duration. The first PC (of 27) represented the main temporal synergy; after practice, it began to account for more of the EMG variance (up to 40%). This main synergy began with a burst in the 4-finger extensor and a silent period in the flexors. There were then progressively later and shorter bursts in the thumb abductor, thumb flexor, little finger abductor, and finally the finger flexors. The results suggest that hand movements may be generated by activity waves unfolding in time. Because finger muscles are under relatively direct cortical control, this suggests a specific form of cortical pattern generation.

Adult↗

Oculomotor responses to gradual changes in target direction.

Smooth pursuit tracking of targets moving linearly (in one dimension) is well characterized by a model where retinal image motion drives eye acceleration. However, previous findings suggest that this model cannot be simply extended to two-dimensional (2D) tracking. To examine 2D pursuit, in the present study, human subjects tracked a target that moved linearly and then followed the arc of a circle. The subjects' gaze angular velocity accurately matched target angular velocity, but the direction of smooth pursuit always lagged behind the current target direction. Pursuit speed slowly declined after the onset of the curve (for about 500 ms), even though the target speed was constant. In a second experiment, brief perturbations were presented immediately prior to the beginning of the change in direction. The subjects' responses to these perturbations consisted of two components: (1) a response specific to the parameters of the perturbation and (2) a nonspecific response that always consisted of a transient decrease in gaze velocity. With the exception of this nonspecific response, pursuit behavior in response to the gradual changes in direction and to the perturbations could be explained by using retinal slip (image velocity) as the input signal. The retinal slip was parallel and perpendicular to the instantaneous direction of pursuit ultimately resulted in changes in gaze velocity (via gaze acceleration). Perhaps due to the subjects' expectations that the target will curve, the sensitivity to the image motion in the direction of pursuit was not as strong as the sensitivity to image motion perpendicular to gaze velocity.

Acceleration↗

Planning and drawing complex shapes.

Arm and hand movements are generally controlled using a combination of sensory-based and memory-based guidance mechanisms. This study examined similarities and differences in visually-guided and memory-guided arm movements, and sought to determine as to what extent certain control principles apply to each type of movement. In particular, the 2/3 power law is a principle that appears to govern the formation of complex, curved hand trajectories; it specifies that the tangential velocity should be proportional to the radius of curvature raised to an exponent of 1/3. A virtual reality system was used to project complex target paths in three-dimensional (3D) space. Human subjects first tracked (with the tip of a handheld pen) a single target moving along an unseen path. The entire target path then became visible and the subject traced the shape. Finally, the target shape disappeared and the subject was to draw it, in the same 3D space, from memory. Most aspects of the movements (speed, path size, shape and arm postures) were very similar across the three conditions. However, subjects adhered to the 2/3 power law most closely in the tracing condition, when the entire target path was visible. Also, only within the tracing condition, there were significant differences in the value of the exponent depending on the size and the spatial orientation of the trajectory. In the tracking and drawing conditions, the exponent was greater than 1/3, indicating that subjects spent more time in areas of tight curvature. This may represent a strategy for learning and remembering the complex shape.

Adult↗

Manual tracking in three dimensions.

Little is known about the manual tracking of targets that move in three dimensions. In the present study, human subjects followed, with the tip of a hand-held pen, a virtual target moving four times (period 5 s) around a novel, unseen path. Two basic types of target paths were used: a peanut-shaped Cassini ellipse and a quasi-spherical shape where four connected semicircles lay in orthogonal planes. The quasi-spherical shape was presented in three different sizes, and the Cassini shape was varied in spatial orientation and by folding it along one of the three bend axes. During the first cycle of Cassini shapes, the hand lagged behind the target by about 150 ms on average, which decreased to 100 ms during the last three cycles. Tracking performance gradually improved during the first 3 s of the first cycle and then stabilized. Tracking was especially good during the smooth, planar sections of the shapes, and time lag was significantly shorter when the tracking of a low-frequency component was compared to performance at a higher frequency (-88 ms at 0.2 Hz vs. -101 ms at 0.6 Hz). Even after the appropriate adjustment of the virtual target path to a virtual shape tracing condition, tracking in depth was poor compared to tracking in the frontal plane, resulting in a flattening of the hand path. In contrast to previous studies where target trajectories were linear or sinusoidal, these complex trajectories may have involved estimation of the overall shape, as well as prediction of target velocity.

Feedback↗

Haptic feature extraction.

This study examined the process by which the shape of a haptically explored object is synthesized from the geometric characteristics of simpler constituent elements, such as arcs and ellipses. Subjects traced the outlines of virtual objects by means of whole arm movements. Each object consisted of the union of a large central ellipse and two smaller circles, extending upward and outward from the top left and right sides of the base. The sizes of the two circles and the eccentricity of the elliptical base were varied. After exploring the object's contour in the absence of vision, subjects reproduced the sensed shape by means of freehand drawing. Speed and force were modulated during the exploratory phase in a manner that suggested that subjects reacted to rather than predicted changes in curvature. Also, subjects typically devoted more time to exploring the part of the contour encompassing the two smaller circles. During drawing, individual features of the explored shape were reproduced with varying degrees of fidelity. Aspects related to the size and location of the smaller circles were reproduced better than was the eccentricity of the ellipse forming the base. Since subjects spent proportionally less time exploring the base, these results suggest that subjects selectively focused attention to regions of high spatial contrast and that the exploratory strategy introduced distortions in the haptically sensed shapes.

Form Perception↗

Functional somatotopy in sensorimotor cortex.

In an effort to understand the highly distributed somatotopy of primary motor cortex, this review draws on principles derived from studies of auditory, visual and somatosensory cortical areas. In each case, a behaviorally important feature or function is overlaid in multiple locations on an underlying topographic map of the peripheral sensory surface. Recent studies of hand muscle synergies suggest the types of two-dimensional functional axes that might reasonably be mapped to the two-dimensional surface of the primary motor cortex. However, other research emphasizes that even a functional somatotopy must be extremely flexible.

Animals↗

Distortions in the visual perception of shape.

It is known that visual illusions lead to a distorted perception of the length and orientation of lines, but it is not clear how these illusions affect the appreciation of the shape of closed forms. In this study two experiments were performed to characterize distortions in the visual perception of the shape of quadrilaterals and the extent to which these distortions were similar to the distortions of haptically sensed shapes. In the first experiment human subjects were presented with two quadrilaterals side by side on a computer monitor. One was a reference shape; the other was rotated and distorted relative to the first. The subjects used the computer mouse to adjust the corners of the distorted quadrilateral to match the shape of the target quadrilateral. They made consistent errors on this task: the adjusted quadrilateral was about 2% wider and about 2% shorter than the veridical shape. Furthermore, subjects adjusted the inner angles of the quadrilateral to make them closer to 90 degrees . The first type of error was also present in a second experiment in which, in a two-alternative forced-choice paradigm, subjects viewed a reference shape and were asked to indicate which of two transiently presented quadrilaterals was closest to the target shape. The width/height errors and the inner angle errors were comparable to those described previously when subjects felt the outline of a quadrilateral and then drew its reproduction in the absence of vision, suggesting that the distortion occurs in the process of remembering the shape.

Adult↗

Smooth pursuit tracking of an abrupt change in target direction: vector superposition of discrete responses.

The directional control of smooth pursuit eye movements was studied by presenting human subjects with targets that moved in a straight line at a constant speed and then changed direction abruptly and unpredictably. To minimize the probability of saccadic responses in the interval following the target's change in direction, target position was offset so as to eliminate position error after the reaction time. Smooth pursuit speed declined at a latency of 90 ms, whereas the direction of smooth pursuit began to change later (130 ms). The amplitude of the offset in target position did not affect the subsequent smooth pursuit response. In other experiments, the target's speed or acceleration was changed abruptly at the time of the change in direction. Step changes in speed elicited short-latency responses in smooth pursuit tracking but step changes in acceleration did not. In all instances, the earliest component of the response did not depend on the parameters of the stimulus. The data were fit with a model in which smooth pursuit resulted from the vector addition of two components, one representing a response to the arrest of the initial target motion and the other the response to the onset of target motion in the new direction. This model gave an excellent fit but further analysis revealed nonlinear interactions between the two vector components. These interactions represented directional anisotropies both in terms of the initial tracking direction (which was either vertical or 45 degrees ) and in terms of the cardinal directions (vertical and horizontal).

Brain↗

Muscular and postural synergies of the human hand.

Because humans have limited ability to independently control the many joints of the hand, a wide variety of hand shapes can be characterized as a weighted combination of just two or three main patterns of covariation in joint rotations, or "postural synergies." The present study sought to align muscle synergies with these main postural synergies and to describe the form of membership of motor units in these postural/muscle synergies. Seventeen joint angles and the electromyographic (EMG) activities of several hand muscles (both intrinsic and extrinsic muscles) were recorded while human subjects held the hand statically in 52 specific shapes (i.e., shaping the hand around 26 commonly grasped objects or forming the 26 letter shapes of a manual alphabet). Principal-components analysis revealed several patterns of muscle synergy, some of which represented either coactivation of all hand muscles, or reciprocal patterns of activity (above and below average levels) in the intrinsic index finger and thumb muscles or (to a lesser extent) in the extrinsic four-tendoned extensor and flexor muscles. Single- and multiunit activity was generally a multimodal function of whole hand shape. This implies that motor-unit activation does not align with a single synergy; instead, motor units participate in multiple muscle synergies. Thus it appears that the organization of the global pattern of hand muscle activation is highly distributed. This organization mirrors the highly fractured somatotopy of cortical hand representations and may provide an ideal substrate for motor learning and recovery from injury.

Electromyography↗

Interception of targets using brief directional cues.

There are time delays in visuomanual and oculomotor pathways, and some of these time delays may be due to the finite time required to process visual motion signals and to extract accurate information about the speed and direction of the motion. The present experiments were designed to ascertain the time required to obtain a reliable estimate of the direction of target motion. Subjects were asked to indicate the final direction of a moving target, which abruptly changed direction and shortly thereafter disappeared, by pointing to its expected emergence at the boundary of an occlusion. Subjects made small but consistent errors that overestimated the target's change in direction. These errors depended little on the amount of time the target was visible (ranging from 50 to 400 ms) after it changed direction. Pointing direction was strongly correlated with gaze, which was dominated by a saccade initiated shortly after the target changed direction. The pointing errors were explained by the fact that the saccade always intercepted the (occluded) target, but then continued in the same direction toward the boundary of the occlusion. The analysis reveals that target direction was estimated accurately even at the shortest viewing time.

Cues↗

Haptic synthesis of shapes and sequences.

Haptic perception of shape is based on kinesthetic and tactile information synthesized across space and time. We studied this process by having subjects move along the edges of multisided shapes and then remember and reproduce the shapes. With eyes closed, subjects moved a robot manipulandum whose force field was programmed to simulate a quadrilateral boundary in a horizontal plane. When subjects then reproduced the quadrilateral using the same manipulandum, with eyes still closed but now with the force field set to zero, they made consistent errors, overestimating the lengths of short segments and underestimating long ones, as well as overestimating acute angles and underestimating obtuse ones. Consequently their reproductions were more regular than the shapes they had experienced. When subjects felt the same quadrilaterals with the same manipulandum but drew them on a vertical screen with visual feedback, they made similar errors, indicating that their distortions reflected mainly perceptual rather than motor processes. In a third experiment, subjects explored the 3 sides of an open shape in a fixed order. The results revealed a temporal pattern of interactions, where the lengths and angles of previously explored segments influenced the drawing of later segments. In all tasks, our subjects were as accurate as subjects in earlier studies who haptically explored only single lines or angles, suggesting that the mental processes that synthesize haptic data from multiple segments into complete shapes do not introduce any net error.

Adolescent↗

Time constants in the perception of a change in the direction of motion in humans.

Motion signals are subject to spatio-temporal filtering at early stages of processing. In general, motion can be characterized by two parameters: speed and direction. This study sought to determine the time constants for the filtering of the directional component of the motion signal. In a forced-choice discrimination task, subjects were asked to choose the more abrupt change in direction of a target that moved through two 90 degrees corners. At each corner, direction of motion was low-pass filtered. Subjects were able to reliably perform this task if the filter time constants differed by >20 ms.

Choice Behavior↗

Coarticulation in fluent fingerspelling.

In speech, the phenomenon of coarticulation (differentiation of phoneme production depending on the preceding or following phonemes) suggests an organization of movement sequences that is not strictly serial. In the skeletal motor system, however, evidence for comparable fluency has been lacking. Thus the present study was designed to quantify coarticulation in the hand movement sequences of sign language interpreters engaged in fingerspelling. Records of 17 measured joint angles were subjected to discriminant and correlation analyses to determine to what extent and in what manner the hand shape for a particular letter was influenced by the hand shapes for the preceding or the following letters. Substantial evidence of coarticulation was found, revealing both forward and reverse influences across letters. These influences could be further categorized as assimilation (tending to reduce the differences between sequential hand shapes) or dissimilation (tending to emphasize the differences between sequential hand shapes). The proximal interphalangeal (PIP) joints of the index and middle fingers tended to show dissimilation, whereas at the same time (i.e., during the spelling of the same letters) the joints of the wrist and thumb tended to show assimilation. The index and middle finger PIP joints have been shown previously to be among the most important joints for computer recognition of the 26 letter shapes, and therefore the dissimilation may have served to enhance visual discrimination. The simultaneous occurrence of dissimilation in some joints and assimilation in others demonstrates an unprecedented level of parallel control of individual joint rotations in an essentially serial task.

Acoustic Stimulation↗

Drawing sequences of segments in 3D: kinetic influences on arm configuration.

Complex movements are generally thought to consist of a series of simpler elements. If this is so, how does the sensorimotor system assemble the pieces? This study recorded and evaluated sequences of arm movements to various targets placed in three-dimensional (3D) space. Subjects performed sequences consisting of single, double, or triple segments with the same first target but with different second targets. The data analysis focused on the first movement segment and evaluated hand path curvature, the hand's final approach to the first target, and the whole arm postures at the beginning and end. Although some idiosyncratic differences in approach were observed, only the final arm posture depended, in a consistent way, on which particular movement was to follow as the second segment. This provided evidence for "coarticulation" of the two segments, only at the level of arm posture, and simulations revealed that this anticipatory modification improved the energetic efficiency of the second segment. Data from movements through five consecutive triple segments (i.e., 5 triangles) were assessed to determine whether kinematic constraints, such as Donders' law, apply to repetitive drawing movements. Although such constraints could prevent the accumulation of changes in arm posture, this was not observed. Instead, in most cases, the elbow was a little bit higher at the end of each triangle than at the beginning. Taken together, the results suggest that coarticulation may facilitate the joining of two segments and the efficiency of the second movement, but does not extend over the drawing of several segments.

Adult↗

Biological constraints simplify the recognition of hand shapes.

This study sought to identify constraints that might lead to a concise system of recognizing fingerspelling hand shapes. Previous studies of grasping suggested that hand shape is controlled using combinations of a small number of neuromuscular synergies, but fingerspelling shapes appear to be more highly individuated and, therefore, might require a larger number of degrees of freedom. Static hand postures of the American Sign Language manual alphabet were recorded by measuring 17 joint angles. Principal components (PCs) analysis was compared to the use of subsets of individual variables (i.e., joint angles) for reduction in degrees of freedom. The first four PCs were similar across subjects. Classification using weightings from these four components was 86.6% accurate, while classification using four individual variables was 88.5% accurate (thumb abduction, as well as flexion at the index and middle finger proximal interphalangeal joints and the ring finger metacarpalphalangeal joint). When chosen for each subject, particular four-variable subsets yielded correct rates above 95%. This superior performance of variable subsets over PC weighting vectors suggests that the reduction in degrees of freedom is due to biomechanical and neuromuscular constraints rather than synergistic control. Thus, in future application to dynamic fingerspelling, reasonable recognition accuracy might be achieved with a significant reduction in both computational and measured degrees of freedom.

Cluster Analysis↗

Using arm configuration to learn the effects of gyroscopes and other devices.

Previous studies have perturbed the association between motor commands and arm movements by applying forces to the arm during two-dimensional movements. These studies have revealed that, when the normal hand path is perturbed, subjects gradually adapt their motor commands to return to this path. The present study used the spin of a gyroscope to create a complex perturbation, as subjects reached to targets presented in three dimensions. Hand path did not change, but the whole-arm geometry ("arm configuration" in four dimensions) was altered. Over a series of several hundred reaches to various targets, subjects gradually returned the arm movement to its normal configuration. Furthermore, during the course of this learning, subjects used a strategy that involved manipulating arm posture. A similar strategy was observed when subjects made reaching movements with a rod attached to the upper arm to change its inertial characteristics. In both cases, the gradual return to the normal arm movement was accomplished without an increase in kinetic energy, suggesting that arm postures and movements (kinematics) and muscular forces (kinetics) may be mutually optimized. In contrast to previous studies, the present results highlight the role of arm configuration (rather than hand path) in learning and control.

Adaptation, Physiological↗

Choosing a wavelet for single-trial EMG.

A wavelet analysis was developed to measure the timing of multiunit bursts in surface electromyograms (EMGs) from single trials. EMG data were taken from eleven elbow and/or shoulder muscles during reaching movements in six different directions, at a range of speeds. A relatively simple wavelet (db2) was chosen, and the analysis focused on wavelet coefficients at an intermediate scale (D3), where the wavelet length approximately matched the wavelengths present in EMG bursts. Burst times were identified from the peaks of the coefficient traces and were plotted as a function of movement time. Linear regression revealed significant relations in most cases, and thus served to validate the wavelet burst identification. With a few exceptions, burst timing scaled in a manner approximately similar to the scaling of movement time. As shown previously with different analytical methods, both within and across joints, EMG bursts were not confined to distinct 'agonist' and 'antagonist' time frames, but instead showed a variety of phases relative to speed or joint torque.

Arm↗

Patterns of hand motion during grasping and the influence of sensory guidance.

This study was aimed at describing temporal synergies of hand movement and determining the influence of sensory cues on the control of these synergies. Subjects were asked to reach to and grasp various objects under three experimental conditions: (1) memory-guided movements, in which the object was not in view during the movement; (2) virtual object, in which a virtual image of the object was in view but the object was not physically present; and (3) real object, in which the object was in view and physically present. Motion of the arm and of 15 degrees of freedom of the hand was recorded. A principal components analysis was developed to provide a concise description of the spatiotemporal patterns underlying the motion. Vision of the object during the reaching movement had no influence on the kinematics, and the effect of the physical presence of the object became manifest primarily after the fingers had contacted the object. Two principal components accounted for >75% of the variance. For both components, there was a strong positive correlation in the rotations of metacarpophalangeal and proximal interphalangeal joints of the fingers. The first principal component exhibited a pattern of finger extension reversing to flexion, whereas the second principal component became important only in the second half of the reaching movement.

Arm↗