Correction: Finger Muscle Attachments for an OpenSim Upper-Extremity Model.

[This corrects the article DOI: 10.1371/journal.pone.0121712.].

Fingertip forces and completion time for index finger and thumb touchscreen gestures.

Users actuate touchscreen computers by applying forces with their fingers to the touchscreen, although the amount and direction of the force is unknown. Our aim was to characterize the magnitude, direction and impulse of the force applied during single finger (tapping and sliding in four directions) and two finger gestures (stretch and pinch). Thirteen subjects performed repeated trials of each gesture. Mean(±SD) resultant force was 0.50(0.09)N for tap, 0.79(0.32)N to 1.18(0.47)N for sliding gestures, 1.47(0.63)N for pinch and 2.05(1.13)N for stretch. Mean resultant force was significantly less (p<0.04) for tap than for all gestures except slide right. The direction of force application was more vertical for the two-finger gestures as compared to the single- finger gestures. Tap was the fastest gesture to complete at 133(83)ms, followed by slide right at 421(181)ms. On average, participants took the longest to complete the stretch gesture at 920(398)ms. Overall, there are differences in forces, force direction, and completion times among touchscreen gestures that could be used to estimate musculoskeletal exposure and help forge guidelines to reduce risk of musculoskeletal injury.

A model for differential leg joint function during human running.

Locomotion requires coordination of leg joints to maintain stability and to maneuver. We studied leg joint function during constant-average-velocity running and the sagittal-plane maneuvers of step ascent and descent. We tested two hypotheses: (1) that leg joints perform distinct functions during locomotion; and (2) that humans select functional parameters to maximize intrinsic dynamic stability. We recorded whole-body kinematics and forces when participants stepped up or down a single vertical step, and found that leg joints show functional differences during both constant-average-velocity locomotion and maneuvers. The hip, knee and ankle function as a motor, damper, and spring, respectively. We therefore constructed a simplified computational model of a human leg with a motor, damper, and spring in series (MDS). The intrinsic dynamics of the model resulted in sustained locomotion on level ground within narrow parameter ranges. However, using parameters experimentally derived from humans, the model showed only short-term stability. Humans may not optimize intrinsic dynamic stability alone, but may instead choose mechanical and behavioral parameters appropriate for both constant-average-velocity locomotion and maneuvers. Understanding joint-level mechanical function during unsteady locomotion helps to understand how differential joint function contributes to whole-body performance, and could lead to improvements in rehabilitation, prosthetic and robotic design.

Index finger and thumb kinematics and performance measurements for common touchscreen gestures.

This study aimed to quantify differences in 7 touchscreen gestures. Eighteen participants performed index finger tapping, sliding in 4 orthogonal directions, and index finger and thumb pinch and stretch gestures on a touchscreen tablet computer. We hypothesized that two finger gestures would require longer task completion time and greater finger joint excursions than sliding gestures using only the index finger. We measured task completion times and finger joint kinematics. Tapping showed the fastest average (±SD) task completion time, 567(190) ms, of all gestures (p < 0.001). Pinch had faster task completion time, 765(277) ms, than all single-finger sliding gestures (p < 0.001). Stretch was faster to complete at 843(317) ms (p < 0.001) than all sliding gestures except slide right. Stretch demonstrated greater mean index finger metacarpophalangeal flexion/extension joint excursions, 63(16)°, compared to sliding gestures, 34(10)°, and tapping, 27(13)° (p < 0.01). Overall, two-finger gestures were faster to complete and showed greater joint excursions than single-finger sliding gestures.

Two-handed grip on a mobile phone affords greater thumb motor performance, decreased variability, and a more extended thumb posture than a one-handed grip.

Holding a mobile computing device with two hands may affect thumb motor performance, joint postures, and device stability compared to holding the device and tapping the touchscreen with the thumb of the holding hand. We tested the hypotheses that holding a touchscreen mobile phone with two hands lead to increased thumb motor performance, different thumb postures, and decreased device movement relative to using one hand. Ten right-handed participants completed reciprocal thumb tapping tasks between emulated keys on a smartphone in either a one- (portrait) or two-handed (landscape) grip configuration. Effective index of performance measured from Fitts' Law was 9% greater (p < 0.001), movement time 7% faster (p < 0.001), and taps were 4% more precise (p < 0.016) for the two-handed grip. Tapping with a two-handed grip involved significantly different wrist and thumb postures than a one-handed grip. Variability of the computing device's movement was 36-63% lower for the two-handed grip compared to the one-handed grip condition (p < 0.001). The support for our hypotheses suggests that a two-handed grip results in increased performance and more extended wrist and thumb postures than a single-handed grip. Device designs that allow two-handed grips may afford increased performance relative to a one-handed grip.

Leg joint function during walking acceleration and deceleration.

Although constant-average-velocity walking has been extensively studied, less is known about walking maneuvers that change speed. We investigated the function of individual leg joints when humans walked at a constant speed, accelerated or decelerated. We hypothesized that leg joints make different functional contributions to maneuvers. Specifically, we hypothesized that the hip generates positive mechanical work (acting like a "motor"), the knee generates little mechanical work (acting like a "strut"), and the ankle absorbs energy during the first half of stance and generates energy during the second half (consistent with "spring"-like function). We recorded full body kinematics and kinetics, used inverse dynamics to estimate net joint moments, and decomposed joint function into strut-, motor-, damper-, and spring-like components using indices based on net joint work. Although overall leg mechanics were primarily strut-like, individual joints did not act as struts during stance. The hip functioned as a power generating "motor," and ankle function was consistent with spring-like behavior. Even though net knee work was small, the knee did not behave solely as a strut but also showed motor-, and damper-like function. Acceleration involved increased motor-like function of the hip and ankle. Deceleration involved decreased hip motor-like function and ankle spring-like function and increased damping at the knee and ankle. Changes to joint mechanical work were primarily due to changes in joint angular displacements and not net moments. Overall, joints maintain different functional roles during unsteady locomotion.

Finger muscle attachments for an OpenSim upper-extremity model.

We determined muscle attachment points for the index, middle, ring and little fingers in an OpenSim upper-extremity model. Attachment points were selected to match both experimentally measured locations and mechanical function (moment arms). Although experimental measurements of finger muscle attachments have been made, models differ from specimens in many respects such as bone segment ratio, joint kinematics and coordinate system. Likewise, moment arms are not available for all intrinsic finger muscles. Therefore, it was necessary to scale and translate muscle attachments from one experimental or model environment to another while preserving mechanical function. We used a two-step process. First, we estimated muscle function by calculating moment arms for all intrinsic and extrinsic muscles using the partial velocity method. Second, optimization using Simulated Annealing and Hooke-Jeeves algorithms found muscle-tendon paths that minimized root mean square (RMS) differences between experimental and modeled moment arms. The partial velocity method resulted in variance accounted for (VAF) between measured and calculated moment arms of 75.5% on average (range from 48.5% to 99.5%) for intrinsic and extrinsic index finger muscles where measured data were available. RMS error between experimental and optimized values was within one standard deviation (S.D) of measured moment arm (mean RMS error = 1.5 mm < measured S.D = 2.5 mm). Validation of both steps of the technique allowed for estimation of muscle attachment points for muscles whose moment arms have not been measured. Differences between modeled and experimentally measured muscle attachments, averaged over all finger joints, were less than 4.9 mm (within 7.1% of the average length of the muscle-tendon paths). The resulting non-proprietary musculoskeletal model of the human fingers could be useful for many applications, including better understanding of complex multi-touch and gestural movements.

Effects of spinal cord injury-induced changes in muscle activation on foot drag in a computational rat ankle model.

Spinal cord injury (SCI) can lead to changes in muscle activation patterns and atrophy of affected muscles. Moderate levels of SCI are typically associated with foot drag during the swing phase of locomotion. Foot drag is often used to assess locomotor recovery, but the causes remain unclear. We hypothesized that foot drag results from inappropriate muscle coordination preventing flexion at the stance-to-swing transition. To test this hypothesis and to assess the relative contributions of neural and muscular changes on foot drag, we developed a two-dimensional, one degree of freedom ankle musculoskeletal model with gastrocnemius and tibialis anterior muscles. Anatomical data collected from sham-injured and incomplete SCI (iSCI) female Long-Evans rats as well as physiological data from the literature were used to implement an open-loop muscle dynamics model. Muscle insertion point motion was calculated with imposed ankle trajectories from kinematic analysis of treadmill walking in sham-injured and iSCI animals. Relative gastrocnemius deactivation and tibialis anterior activation onset times were varied within physiologically relevant ranges based on simplified locomotor electromyogram profiles. No-atrophy and moderate muscle atrophy as well as normal and injured muscle activation profiles were also simulated. Positive moments coinciding with the transition from stance to swing phase were defined as foot swing and negative moments as foot drag. Whereas decreases in activation delay caused by delayed gastrocnemius deactivation promote foot drag, all other changes associated with iSCI facilitate foot swing. Our results suggest that even small changes in the ability to precisely deactivate the gastrocnemius could result in foot drag after iSCI.

Extrinsic and intrinsic index finger muscle attachments in an OpenSim upper-extremity model.

Musculoskeletal models allow estimation of muscle function during complex tasks. We used objective methods to determine possible attachment locations for index finger muscles in an OpenSim upper-extremity model. Data-driven optimization algorithms, Simulated Annealing and Hook-Jeeves, estimated tendon locations crossing the metacarpophalangeal (MCP), proximal interphalangeal (PIP) and distal interphalangeal (DIP) joints by minimizing the difference between model-estimated and experimentally-measured moment arms. Sensitivity analysis revealed that multiple sets of muscle attachments with similar optimized moment arms are possible, requiring additional assumptions or data to select a single set of values. The most smooth muscle paths were assumed to be biologically reasonable. Estimated tendon attachments resulted in variance accounted for (VAF) between calculated moment arms and measured values of 78% for flex/extension and 81% for ab/adduction at the MCP joint. VAF averaged 67% at the PIP joint and 54% at the DIP joint. VAF values at PIP and DIP joints partially reflected the constant moment arms reported for muscles about these joints. However, all moment arm values found through optimization were non-linear and non-constant. Relationships between moment arms and joint angles were best described with quadratic equations for tendons at the PIP and DIP joints.

A data-driven design evaluation tool for handheld device soft keyboards.

Thumb interaction is a primary technique used to operate small handheld devices such as smartphones. Despite the different techniques involved in operating a handheld device compared to a personal computer, the keyboard layouts for both devices are similar. A handheld device keyboard that considers the physical capabilities of the thumb may improve user experience. We developed and applied a design evaluation tool for different geometries of the QWERTY keyboard using a performance evaluation model. The model utilizes previously collected data on thumb motor performance and posture for different tap locations and thumb movement directions. We calculated a performance index (PITOT, 0 is worst and 2 is best) for 663 designs consisting in different combinations of three variables: the keyboard's radius of curvature (R) (mm), orientation (O) (°), and vertical location on the screen (L). The current standard keyboard performed poorly (PITOT = 0.28) compared to other designs considered. Keyboard location (L) contributed to the greatest variability in performance out of the three design variables, suggesting that designers should modify this variable first. Performance was greatest for designs in the middle keyboard location. In addition, having a slightly upward curve (R = -20 mm) and orientated perpendicular to the thumb's long axis (O = -20°) improved performance to PITOT = 1.97. Poorest performances were associated with placement of the keyboard's spacebar in the bottom right corner of the screen (e.g., the worst was for R = 20 mm, O = 40°, L =  Bottom (PITOT = 0.09)). While this evaluation tool can be used in the design process as an ergonomic reference to promote user motor performance, other design variables such as visual access and usability still remain unexplored.

Compensations during Unsteady Locomotion.

Locomotion in a complex environment is often not steady, but the mechanisms used by animals to power and control unsteady locomotion (stability and maneuverability) are not well understood. We use behavioral, morphological, and impulsive perturbations to determine the compensations used during unsteady locomotion. At the level both of the whole-body and of joints, quasi-stiffness models are useful for describing adjustments to the functioning of legs and joints during maneuvers. However, alterations to the mechanics of legs and joints often are distinct for different phases of the step cycle or for specific joints. For example, negotiating steps involves independent changes of leg stiffness during compression and thrust phases of stance. Unsteady locomotion also involves parameters that are not part of the simplest reduced-parameter models of locomotion (e.g., the spring-loaded inverted pendulum) such as moments of the hip joint. Extensive coupling among translational and rotational parameters must be taken into account to stabilize locomotion or maneuver. For example, maneuvers with morphological perturbations (increased rotational inertial turns) involve changes to several aspects of movement, including the initial conditions of rotation and ground-reaction forces. Coupled changes to several parameters may be employed to control maneuvers on a trial-by-trial basis. Compensating for increased rotational inertia of the body during turns is facilitated by the opposing effects of several mechanical and behavioral parameters. However, the specific rules used by animals to control translation and rotation of the body to maintain stability or maneuver have not been fully characterized. We initiated direct-perturbation experiments to investigate the strategies used by humans to maintain stability following center-of-mass (COM) perturbations. When walking, humans showed more resistance to medio-lateral perturbations (lower COM displacement). However, when running, humans could recover from the point of maximum COM displacement faster than when walking. Consequently, the total time necessary for recovery was not significantly different between walking and running. Future experiments will determine the mechanisms used for compensations during unsteady locomotion at the behavioral, joint, and muscle levels. Using reduced-parameter models will allow common experimental and analytical frameworks for the study of both stability and maneuverability and the determination of general control strategies for unsteady locomotion.

Compensations for increased rotational inertia during human cutting turns.

Locomotion in a complex environment is often not steady state, but unsteady locomotion (stability and maneuverability) is not well understood. We investigated the strategies used by humans to perform sidestep cutting turns when running. Previous studies have argued that because humans have small yaw rotational moments of inertia relative to body mass, deceleratory forces in the initial velocity direction that occur during the turning step, or 'braking' forces, could function to prevent body over-rotation during turns. We tested this hypothesis by increasing body rotational inertia and testing whether braking forces during stance decreased. We recorded ground reaction force and body kinematics from seven participants performing 45 deg sidestep cutting turns and straight running at five levels of body rotational inertia, with increases up to fourfold. Contrary to our prediction, braking forces remained consistent at different rotational inertias, facilitated by anticipatory changes to body rotational speed. Increasing inertia revealed that the opposing effects of several turning parameters, including rotation due to symmetrical anterior-posterior forces, result in a system that can compensate for fourfold changes in rotational inertia with less than 50% changes to rotational velocity. These results suggest that in submaximal effort turning, legged systems may be robust to changes in morphological parameters, and that compensations can involve relatively minor adjustments between steps to change initial stance conditions.

Tablet Keyboard Configuration Affects Performance, Discomfort and Task Difficulty for Thumb Typing in a Two-Handed Grip.

When holding a tablet computer with two hands, the touch keyboard configuration imposes postural constraints on the user because of the need to simultaneously hold the device and type with the thumbs. Designers have provided users with several possible keyboard configurations (device orientation, keyboard layout and location). However, potential differences in performance, usability and postures among these configurations have not been explored. We hypothesize that (1) the narrower standard keyboard layout in the portrait orientation leads to lower self-reported discomfort and less reach than the landscape orientation; (2) a split keyboard layout results in better overall outcomes compared to the standard layout; and (3) the conventional bottom keyboard location leads to the best outcomes overall compared to other locations. A repeated measures laboratory experiment of 12 tablet owners measured typing speed, discomfort, task difficulty, and thumb/wrist joint postures using an active marker system during typing tasks for different combinations of device orientation (portrait and landscape), keyboard layout (standard and split), and keyboard location (bottom, middle, top). The narrower standard keyboard with the device in the portrait orientation was associated with less discomfort (least squares mean (and S.E.) 2.9±0.6) than the landscape orientation (4.5±0.7). Additionally, the split keyboard decreased the amount of reaching required by the thumb in the landscape orientation as defined by a reduced range of motion and less MCP extension, which may have led to reduced discomfort (2.7±0.6) compared to the standard layout (4.5±0.7). However, typing speed was greater for the standard layout (127±5 char./min.) compared to the split layout (113±4 char./min.) regardless of device orientation and keyboard location. Usage guidelines and designers can incorporate these findings to optimize keyboard design parameters and form factors that promote user performance and usability for thumb interaction.

Thumb motor performance varies with thumb and wrist posture during single-handed mobile phone use.

Design features of mobile computing technology such as device size and key location may affect thumb motor performance during single-handed use. Since single-handed use requires the thumb posture to vary with key location, we hypothesize that motor performance is associated with thumb and wrist joint postures. A repeated measures laboratory experiment of 10 right-handed participants measured thumb and wrist joint postures during reciprocal tapping tasks between two keys for different key pairs among 12 emulated keys. Fitts' effective index of performance and joint postures at contact with each key were averaged across trials for each key. Thumb motor performance varied for different keys, with poorest performances being associated with excessive thumb flexion such as when tapping on keys closest to the base of the thumb in the bottom right corner of the phone. Motor performance was greatest when the thumb was in a typical resting posture, neither significantly flexed nor fully extended with slight CMC joint abduction and supination, such as when tapping on keys located in the top right and middle left areas on the phone. Grip was also significantly affected by key location, with the most extreme differences being between the top left and bottom right corners of the phone. These results suggest that keypad designs aimed at promoting performance for single-handed use should avoid placing frequently used functions and keys close to the base of the thumb and instead should consider key locations that require a thumb posture away from its limits in flexion/extension, as these postures promote motor performance.

Quantitative metrics of spinal cord injury recovery in the rat using motion capture, electromyography and ground reaction force measurement.

Toward improving the quantitative tools available for evaluation of locomotion after a spinal cord injury, we characterized selected biomechanical and physiological parameters that could be used to assess the level of recovery of locomotion after a mid-thoracic spinal cord lateral hemisection. Specifically we defined quantitative measures of muscle activation and coordination, body weight support, propulsive force, and pre-toe contact activation. Generation of this ensemble of recovery measures was based on kinematics, ground reaction forces, and EMG in rats from the hindlimb ipsilateral to the hemisection during quadrupedal running on a trackway. We derived muscle activation levels using inverse dynamics and static optimization applied to a model of the hindlimb musculoskeletal system. Rats exhibited a phased recovery pattern: progressive recovery of general muscle activity beginning within 2-3 days post-injury, followed by recovery of propulsive force and intralimb coordination of antagonistic muscles 12-13 days post-injury. Even at 12-13 days post-injury however, body weight support and the normal pre-paw contact EMG burst were significantly impaired. These data are consistent with a differential rate of recovery of general motor pool recruitment, and coordination among motor pools. The results demonstrate the discriminative potential of these physiologically based measures in quantifying the progressive recovery of gait performance after a lateral spinal cord hemisection.

Task-level strategies for human sagittal-plane running maneuvers are consistent with robotic control policies.

The strategies that humans use to control unsteady locomotion are not well understood. A "spring-mass" template comprised of a point mass bouncing on a sprung leg can approximate both center of mass movements and ground reaction forces during running in humans and other animals. Legged robots that operate as bouncing, "spring-mass" systems can maintain stable motion using relatively simple, distributed feedback rules. We tested whether the changes to sagittal-plane movements during five running tasks involving active changes to running height, speed, and orientation were consistent with the rules used by bouncing robots to maintain stability. Changes to running height were associated with changes to leg force but not stance duration. To change speed, humans primarily used a "pogo stick" strategy, where speed changes were associated with adjustments to fore-aft foot placement, and not a "unicycle" strategy involving systematic changes to stance leg hip moment. However, hip moments were related to changes to body orientation and angular speed. Hip moments could be described with first order proportional-derivative relationship to trunk pitch. Overall, the task-level strategies used for body control in humans were consistent with the strategies employed by bouncing robots. Identification of these behavioral strategies could lead to a better understanding of the sensorimotor mechanisms that allow for effective unsteady locomotion.

Axon regeneration can facilitate or suppress hindlimb function after olfactory ensheathing glia transplantation.

Reports based primarily on anatomical evidence suggest that olfactory ensheathing glia (OEG) transplantation promotes axon regeneration across a complete spinal cord transection in adult rats. Based on functional, electrophysiological, and anatomical assessments, we found that OEG promoted axon regeneration across a complete spinal cord transection and that this regeneration altered motor responses over time. At 7 months after transection, 70% of OEG-treated rats showed motor-evoked potentials in hindlimb muscles after transcranial electric stimulation. Furthermore, a complete spinal cord retransection performed 8 months after injury demonstrated that this axon regeneration suppressed locomotor performance and decreased the hypersensitive hindlimb withdrawal response to mechanical stimulation. OEG transplantation alone promoted reorganization of lumbosacral locomotor networks and, when combined with long-term training, enhanced some stepping measures. These novel findings demonstrate that OEG promote regeneration of mature axons across a complete transection and reorganization of spinal circuitry, both of which contribute to sensorimotor function.

Application of a rat hindlimb model: a prediction of force spaces reachable through stimulation of nerve fascicles.

A device to generate standing or locomotion through chronically placed electrodes has not been fully developed due in part to limitations of clinical experimentation and the high number of muscle activation inputs of the leg. We investigated the feasibility of functional electrical stimulation paradigms that minimize the input dimensions for controlling the limbs by stimulating at nerve fascicles, utilizing a model of the rat hindlimb, which combined previously collected morphological data with muscle physiological parameters presented herein. As validation of the model, we investigated the suitability of a lumped-parameter model for the prediction of muscle activation during dynamic tasks. Using the validated model, we found that the space of forces producible through activation of muscle groups sharing common nerve fascicles was nonlinearly dependent on the number of discrete muscle groups that could be individually activated (equivalently, the neuroanatomical level of activation). Seven commonly innervated muscle groups were sufficient to produce 78% of the force space producible through individual activation of the 42 modeled hindlimb muscles. This novel, neuroanatomically derived reduction in input dimension emphasizes the potential to simplify controllers for functional electrical stimulation to improve functional recovery after a neuromuscular injury.

Unconstrained three-dimensional reaching in rhesus monkeys.

To better understand normative behavior for quantitative evaluation of motor recovery after injury, we studied arm movements by non-injured rhesus monkeys during a food-retrieval task. While seated, monkeys reached, grasped, and retrieved food items. We recorded three-dimensional kinematics and muscle activity, and used inverse dynamics to calculate joint moments due to gravity, segmental interactions, and to the muscles and tissues of the arm. Endpoint paths showed curvature in three dimensions, suggesting that maintaining straight paths was not an important constraint. Joint moments were dominated by gravity. Generalized muscle and interaction moments were less than half of the gravitational moments. The relationships between shoulder and elbow resultant moments were linear during both reach and retrieval. Although both reach and retrieval required elbow flexor moments, an elbow extensor (triceps brachii) was active during both phases. Antagonistic muscles of both the elbow and hand were co-activated during reach and retrieval. Joint behavior could be described by lumped-parameter models analogous to torsional springs at the joints. Minor alterations to joint quasi-stiffness properties, aided by interaction moments, result in reciprocal movements that evolve under the influence of gravity. The strategies identified in monkeys to reach, grasp, and retrieve items will allow the quantification of prehension during recovery after a spinal cord injury and the effectiveness of therapeutic interventions.

Extensive spontaneous plasticity of corticospinal projections after primate spinal cord injury.

Although axonal regeneration after CNS injury is limited, partial injury is frequently accompanied by extensive functional recovery. To investigate mechanisms underlying spontaneous recovery after incomplete spinal cord injury, we administered C7 spinal cord hemisections to adult rhesus monkeys and analyzed behavioral, electrophysiological and anatomical adaptations. We found marked spontaneous plasticity of corticospinal projections, with reconstitution of fully 60% of pre-lesion axon density arising from sprouting of spinal cord midline-crossing axons. This extensive anatomical recovery was associated with improvement in coordinated muscle recruitment, hand function and locomotion. These findings identify what may be the most extensive natural recovery of mammalian axonal projections after nervous system injury observed to date, highlighting an important role for primate models in translational disease research.