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1.  Origins of submovements in movements of elderly adults 
Background
Slowness is a well-recognized feature of movements in aging. One of the possible reasons for slowness suggested by previous research is production of corrective submovements that compensate for shortened primary submovement to the target. Here, we re-examine this traditional interpretation and argue that the majority of submovements in older adults may be a consequence rather than the cause of slowness.
Methods
Pointing movements in young and older adults were recorded. Conditions for submovement emergence were manipulated by using small and large targets and three movement modes: discrete (required stopping on the target), reciprocal (required reversal on the target), and passing (required crossing the target and stopping after that). Movements were parsed into a primary and secondary submovement based on zero-crossings of velocity (type 1 submovements), acceleration (type 2 submovements), and jerk (type 3 submovements). In the passing mode, secondary submovements were analyzed only after crossing the target to exclude that they were accuracy adjustments.
Results
Consistent with previous research, the primary submovement was shortened and total secondary submovement incidence was increased in older adults. However, comparisons across conditions suggested that many submovements were non-corrective in both groups. Type 1 submovements were non-corrective because they were more frequent for large than small targets. They predominantly emerged due to arm stabilization and energy dissipation during motion termination in the discrete and passing mode. Although type 2 and 3 submovements were more frequent for small than large targets, this trend was also observed in the passing mode, suggesting that many of these submovements were non-corrective. Rather, they could have been velocity fluctuations associated predominantly with low speed of movements to small targets.
Conclusion
The results question the traditional interpretation of frequent submovements in older adults as corrective adjustments. Rather, the increased incidence of submovements in older adults is directly related to low movement speed observed in aging, whereas the relationship between submovement incidence and target size is a result of speed-accuracy trade-off. Aging-related declines in muscular control that may contribute to the disproportional increases in submovement incidence during slow movements of older adults are discussed.
doi:10.1186/1743-0003-5-28
PMCID: PMC2628348  PMID: 19014548
2.  Human basal ganglia and the dynamic control of force during on-line corrections 
Natural movements are corrected in part by the generation of submovements, occuring early in a movement such that they amend an ongoing action. Submovements are associated with activity of the basal ganglia, implying a role for the structures in error correction. In parallel, the basal ganglia are linked to the generation and control of force amplitude, change and duration. Here we tested if activity in human basal ganglia is associated with submovements generally, or was specific to a condition where the submovements only occurred in the face of unexpected proprioceptive error. Submovements were induced by introducing unexpected and variable viscous loads (augmenting the need for trial specific grip forces) or by reducing target size (augmenting the need for visually guided on-line control) in a 1-D target capture task. In both cases, subjects compensated for the increased task difficulty by generating corrective submovements, which were closely matched in frequency and type. Activity in the internal segment of the globus pallidus and subthalamic nucleus correlated strongly with the number of submovements during the viscous challenge but not with the target challenge. The effects could not be explained by kinematic differences, i.e. movement amplitude or average number of submovements. The results support a specific role for the basal ganglia in error correction under conditions of variable load where there is a need for the dynamic control of force within an ongoing movement.
doi:10.1523/JNEUROSCI.3301-10.2011
PMCID: PMC3037279  PMID: 21289168
Motor control; on-line correction; error; adaptation; basal ganglia; human
3.  Submovement Composition of Head Movement 
PLoS ONE  2012;7(11):e47565.
Limb movement is smooth and corrections of movement trajectory and amplitude are barely noticeable midflight. This suggests that skeletomuscular motor commands are smooth in transition, such that the rate of change of acceleration (or jerk) is minimized. Here we applied the methodology of minimum-jerk submovement decomposition to a member of the skeletomuscular family, the head movement. We examined the submovement composition of three types of horizontal head movements generated by nonhuman primates: head-alone tracking, head-gaze pursuit, and eye-head combined gaze shifts. The first two types of head movements tracked a moving target, whereas the last type oriented the head with rapid gaze shifts toward a target fixed in space. During head tracking, the head movement was composed of a series of episodes, each consisting of a distinct, bell-shaped velocity profile (submovement) that rarely overlapped with each other. There was no specific magnitude order in the peak velocities of these submovements. In contrast, during eye-head combined gaze shifts, the head movement was often comprised of overlapping submovements, in which the peak velocity of the primary submovement was always higher than that of the subsequent submovement, consistent with the two-component strategy observed in goal-directed limb movements. These results extend the previous submovement composition studies from limb to head movements, suggesting that submovement composition provides a biologically plausible approach to characterizing the head motor recruitment that can vary depending on task demand.
doi:10.1371/journal.pone.0047565
PMCID: PMC3489904  PMID: 23139749
4.  Characterization of Information-Based Learning Benefits with Submovement Dynamics and Muscular Rhythmicity 
PLoS ONE  2013;8(12):e82920.
For skill advancement, motor variability must be optimized based on target information during practice sessions. This study investigated structural changes in kinematic variability by characterizing submovement dynamics and muscular oscillations after practice with visuomotor tracking under different target conditions. Thirty-six participants were randomly assigned to one of three groups (simple, complex, and random). Each group practiced tracking visual targets with trajectories of varying complexity. The velocity trajectory of tracking was decomposed into 1) a primary contraction spectrally identical to the target rate and 2) an intermittent submovement profile. The learning benefits and submovement dynamics were conditional upon experimental manipulation of the target information. Only the simple and complex groups improved their skills with practice. The size of the submovements was most greatly reduced by practice with the least target information (simple > complex > random). Submovement complexity changed in parallel with learning benefits, with the most remarkable increase in practice under a moderate amount of target information (complex > simple > random). In the simple and complex protocols, skill improvements were associated with a significant decline in alpha (8–12 Hz) muscular oscillation but a potentiation of gamma (35–50 Hz) muscular oscillation. However, the random group showed no significant change in tracking skill or submovement dynamics, except that alpha muscular oscillation was reduced. In conclusion, submovement and gamma muscular oscillation are biological markers of learning benefits. Effective learning with an appropriate amount of target information reduces the size of submovements. In accordance with the challenge point hypothesis, changes in submovement complexity in response to target information had an inverted-U function, pertaining to an abundant trajectory-tuning strategy with target exactness.
doi:10.1371/journal.pone.0082920
PMCID: PMC3867443  PMID: 24367568
5.  Age-related changes in speed and accuracy during rapid targeted center of pressure movements near the posterior limit of the base of support 
Background
Backward falls are often associated with injury, particularly among older women. An age-related increase occurs in center of pressure variability when standing and leaning. So, we hypothesized that, in comparison to young women, older women would display a disproportionate decrease of speed and accuracy in the primary center of pressure submovements as movement amplitude increases.
Methods
Ground reaction forces were recorded from thirteen healthy young and twelve older women while performing rapid, targeted, center of pressure movements of small and large amplitude in upright stance. Measures included center of pressure speed, the number of center of pressure submovements, and the incidence rate of primary center of pressure submovements undershooting the target.
Findings
In comparison to young women, older women used slower primary submovements, particularly as movement amplitude increased (P < 0.01). Even though older women achieved similar endpoint accuracy, they demonstrated a 2 to 5-fold increase in the incidence of primary submovement undershooting for large-amplitude movements (P < 0.01). Overall, posterior center of pressure movements of older women were 41% slower and exhibited 43% more secondary submovements than in young women (P < 0.01).
Interpretations
We conclude that the increased primary submovement undershoots and secondary center of pressure submovements in the older women reflect the use of a conservative control strategy near the posterior limit of their base of support.
doi:10.1016/j.clinbiomech.2012.06.007
PMCID: PMC3444664  PMID: 22770467
Balance; Center of pressure submovement; Postural control; Aging
6.  Movement structure in young and elderly adults during goal-directed movements of the left and right arm 
Brain and cognition  2008;69(1):30-38.
Elderly adults often exhibit performance deficits during goal-directed movements of the dominant arm compared with young adults. Recent studies involving hemispheric lateralization have provided evidence that the dominant and non-dominant hemisphere-arm systems are specialized for controlling different movement parameters and that hemispheric specialization may be reduced during normal aging. The purpose was to examine age-related differences in the movement structure for the dominant (right) and non-dominant (left) during goal-directed movements. Young and elderly adults performed 72 aiming movements as fast and as accurately as possible to visual targets with both arms. The findings suggest that previous research utilizing the dominant arm can be generalized to the non-dominant arm because performance was similar for the two arms. However, as expected, the elderly adults showed shorter relative primary submovement lengths and longer relative primary submovement durations, reaction times, movement durations, and normalized jerk scores compared to the young adults.
doi:10.1016/j.bandc.2008.05.002
PMCID: PMC2663567  PMID: 18556103
Aging; Feedback; Hand; Laterality; Submovement
7.  Dynamic Primitives of Motor Behavior 
Biological cybernetics  2012;106(0):727-739.
We present in outline a theory of sensorimotor control based on dynamic primitives, which we define as attractors. To account for the broad class of human interactive behaviors—especially tool use—we propose three distinct primitives: submovements, oscillations and mechanical impedances, the latter necessary for interaction with objects. Due to fundamental features of the neuromuscular system, most notably its slow response, we argue that encoding in terms of parameterized primitives may be an essential simplification required for learning, performance, and retention of complex skills. Primitives may simultaneously and sequentially be combined to produce observable forces and motions. This may be achieved by defining a virtual trajectory composed of submovements and/or oscillations interacting with impedances. Identifying primitives requires care: in principle, overlapping submovements would be sufficient to compose all observed movements but biological evidence shows that oscillations are a distinct primitive. Conversely, we suggest that kinematic synergies, frequently discussed as primitives of complex actions, may be an emergent consequence of neuromuscular impedance. To illustrate how these dynamic primitives may account for complex actions, we briefly review three types of interactive behaviors: constrained motion, impact tasks, and manipulation of dynamic objects.
doi:10.1007/s00422-012-0527-1
PMCID: PMC3735361  PMID: 23124919
Discrete; submovement; rhythmic; oscillation; impedance; primitive
8.  The effect of age, movement direction, and target size on the maximum speed of targeted COP movements in healthy women 
Human movement science  2012;31(5):1213-1223.
Rapid center of pressure (COP) movements are often required to avoid falls. Little is known about the effect of age on rapid and accurate volitional COP movements. We hypothesized that COP movements to a target would be slower and exhibit more submovements in older versus younger adults, particularly in posterior versus anterior movements. Healthy older (N = 12, mean age = 76 years) and young women (N = 13, mean age = 23 years) performed anterior and posterior lean movements while standing on a force plate, and were instructed to move their COP ‘as fast and as accurately as possible’ using visual feedback. The results show that rapid posterior COP movements were slower and had an increased number of submovements and ratio of peak-to-average velocity, in comparison to anterior movements (p < .005). Moreover, older compared to younger adults were 27% slower and utilized nearly twice as many compensatory submovements (p < .005), particularly when moving posteriorly (p < .05). Older women also had higher ratios of peak-to-average COP velocity than young (p < .05). Thus, despite moving more slowly, older women needed to take more frequent submovements to maintain COP accuracy, particularly posteriorly, thereby providing evidence of a compensatory strategy that may be used for preventing backward falls.
doi:10.1016/j.humov.2011.11.002
PMCID: PMC3330159  PMID: 22225924
Balance; Center of pressure; Speed-accuracy trade-offs; Gerontology
9.  Dynamic primitives in the control of locomotion 
Humans achieve locomotor dexterity that far exceeds the capability of modern robots, yet this is achieved despite slower actuators, imprecise sensors, and vastly slower communication. We propose that this spectacular performance arises from encoding motor commands in terms of dynamic primitives. We propose three primitives as a foundation for a comprehensive theoretical framework that can embrace a wide range of upper- and lower-limb behaviors. Building on previous work that suggested discrete and rhythmic movements as elementary dynamic behaviors, we define submovements and oscillations: as discrete movements cannot be combined with sufficient flexibility, we argue that suitably-defined submovements are primitives. As the term “rhythmic” may be ambiguous, we define oscillations as the corresponding class of primitives. We further propose mechanical impedances as a third class of dynamic primitives, necessary for interaction with the physical environment. Combination of these three classes of primitive requires care. One approach is through a generalized equivalent network: a virtual trajectory composed of simultaneous and/or sequential submovements and/or oscillations that interacts with mechanical impedances to produce observable forces and motions. Reliable experimental identification of these dynamic primitives presents challenges: identification of mechanical impedances is exquisitely sensitive to assumptions about their dynamic structure; identification of submovements and oscillations is sensitive to their assumed form and to details of the algorithm used to extract them. Some methods to address these challenges are presented. Some implications of this theoretical framework for locomotor rehabilitation are considered.
doi:10.3389/fncom.2013.00071
PMCID: PMC3689288  PMID: 23801959
discrete; submovement; rhythmic; oscillation; impedance; primitive; locomotion; rehabilitation
10.  Action selection and refinement in subcortical loops through basal ganglia and cerebellum 
Subcortical loops through the basal ganglia and the cerebellum form computationally powerful distributed processing modules (DPMs). This paper relates the computational features of a DPM's loop through the basal ganglia to experimental results for two kinds of natural action selection. First, functional imaging during a serial order recall task was used to study human brain activity during the selection of sequential actions from working memory. Second, microelectrode recordings from monkeys trained in a step-tracking task were used to study the natural selection of corrective submovements. Our DPM-based model assisted in the interpretation of puzzling data from both of these experiments. We come to posit that the many loops through the basal ganglia each regulate the embodiment of pattern formation in a given area of cerebral cortex. This operation serves to instantiate different kinds of action (or thought) mediated by different areas of cerebral cortex. We then use our findings to formulate a model of the aetiology of schizophrenia.
doi:10.1098/rstb.2007.2063
PMCID: PMC2440782  PMID: 17428771
modularity; serial order; pattern classification; error correction; schizophrenia; presynaptic inhibition
11.  CO-REGULATION OF MOVEMENT SPEED AND ACCURACY BY CHILDREN WITH HEAVY PRENATAL ALCOHOL EXPOSURE1, 2 
Perceptual and motor skills  2011;112(1):172-182.
Summary
The study investigated how children with heavy prenatal alcohol exposure regulate movement speed and accuracy during goal-directed movements. 16 children ages 7 to 17 years with confirmed histories of heavy in utero alcohol exposure, and 21 nonalcohol-exposed control children completed a series of reciprocal tapping movements between two spatial targets. 5 different targets sets were presented, representing a range of task difficulty between 2 and 6 bits of information. Estimates of percent error rate, movement time, slope, and linear fit of the resulting curve confirmed that for goal-directed, reciprocal tapping responses, performance of the group with prenatal alcohol exposure was described by a linear function, as predicted by Fitts’ law, by sacrificing movement accuracy. The index of performance was the same for the two groups: it initially increased, then leveled off for more difficult movements.
PMCID: PMC3838298  PMID: 21466090
12.  Recognizing Biological Motion and Emotions from Point-Light Displays in Autism Spectrum Disorders 
PLoS ONE  2012;7(9):e44473.
One of the main characteristics of Autism Spectrum Disorder (ASD) are problems with social interaction and communication. Here, we explored ASD-related alterations in ‘reading’ body language of other humans. Accuracy and reaction times were assessed from two observational tasks involving the recognition of ‘biological motion’ and ‘emotions’ from point-light displays (PLDs). Eye movements were recorded during the completion of the tests. Results indicated that typically developed-participants were more accurate than ASD-subjects in recognizing biological motion or emotions from PLDs. No accuracy differences were revealed on two control-tasks (involving the indication of color-changes in the moving point-lights). Group differences in reaction times existed on all tasks, but effect sizes were higher for the biological and emotion recognition tasks. Biological motion recognition abilities were related to a person’s ability to recognize emotions from PLDs. However, ASD-related atypicalities in emotion recognition could not entirely be attributed to more basic deficits in biological motion recognition, suggesting an additional ASD-specific deficit in recognizing the emotional dimension of the point light displays. Eye movements were assessed during the completion of tasks and results indicated that ASD-participants generally produced more saccades and shorter fixation-durations compared to the control-group. However, especially for emotion recognition, these altered eye movements were associated with reductions in task-performance.
doi:10.1371/journal.pone.0044473
PMCID: PMC3435310  PMID: 22970227
13.  Motor imagery of tool use: relationship to actual use and adherence to Fitts’ law across tasks 
A Fitts’ task was used to investigate how tools are incorporated into the internal representations that underlie pointing movements, and whether such knowledge can be generalized across tasks. We measured the speed-accuracy trade-offs that occurred as target width was varied for both real and imagined movements. The dynamics of the pointing tool used in the task were manipulated—regular pen, top-heavy tool, and bottom-heavy tool—to test the fidelity of internal representations of movements involving the use of novel tools. To test if such representations can be generalized, the orientation of the pointing task was also manipulated (horizontal vs. vertical). In all conditions, both real and imagined performances conformed to the speed-accuracy relationship described by Fitts’ law. We found significant differences in imagined MTs for the two weighted tools compared to the regular pen, but not between the weighted tools. By contrast, real movement durations differed between all tools. These results indicate that even relatively brief experience using novel tools is sufficient to influence the internal representation of the dynamics of the tool-limb system. However, in the absence of feedback, these representations do not make explicit differences in performances resulting from the unique dynamics of these weighted tools.
doi:10.1007/s00221-012-3004-0
PMCID: PMC3351569  PMID: 22294026
Tool use; Imagery; Visually guided movement; Sensorimotor control; Fitts’ law; Motor learning
14.  A sensory source for motor variation 
Nature  2005;437(7057):10.1038/nature03961.
Suppose that the variability in our movements1–9 is caused not by noise in the motor system itself, nor by fluctuations in our intentions or plans, but rather by errors in our sensory estimates of the external parameters that define the appropriate action. For tasks in which precision is at a premium, performance would be optimal if no noise were added in movement planning and execution: motor output would be as accurate as possible given the quality of sensory inputs. Here we use visually guided smooth-pursuit eye movements in primates10 as a testing ground for this notion of optimality. In response to repeated presentations of identical target motions, nearly 92% of the variance in eye trajectory can be accounted for as a consequence of errors in sensory estimates of the speed, direction and timing of target motion, plus a small background noise that is observed both during eye movements and during fixations. The magnitudes of the inferred sensory errors agree with the observed thresholds for sensory discrimination by perceptual systems, suggesting that the very different neural processes of perception and action are limited by the same sources of noise.
doi:10.1038/nature03961
PMCID: PMC2551316  PMID: 16163357
15.  Distinguishing highly confident accurate and inaccurate memory: insights about relevant and irrelevant influences on memory confidence 
Memory (Hove, England)  2012;20(1):48-62.
It is generally believed that accuracy and confidence in one’s memory are related, but there are many instances when they diverge. Accordingly, it is important to disentangle the factors which contribute to memory accuracy and confidence, especially those factors that contribute to confidence, but not accuracy. We used eye movements to separately measure fluent cue processing, the target recognition experience, and relative evidence assessment on recognition confidence and accuracy. Eye movements were monitored during a face-scene associative recognition task, in which participants first saw a scene cue, followed by a forced-choice recognition test for the associated face, with confidence ratings. Eye movement indices of the target recognition experience were largely indicative of accuracy, and showed a relationship to confidence for accurate decisions. In contrast, eye movements during the scene cue raised the possibility that more fluent cue processing was related to higher confidence for both accurate and inaccurate recognition decisions. In a second experiment, we manipulated cue familiarity, and therefore cue fluency. Participants showed higher confidence for cue-target associations for when the cue was more familiar, especially for incorrect responses. These results suggest that over-reliance on cue familiarity and under-reliance on the target recognition experience may lead to erroneous confidence.
doi:10.1080/09658211.2011.633919
PMCID: PMC3269835  PMID: 22171810
16.  Both Lexical and Non-Lexical Characters Are Processed during Saccadic Eye Movements 
PLoS ONE  2012;7(9):e46383.
On average our eyes make 3–5 saccadic movements per second when we read, although their neural mechanism is still unclear. It is generally thought that saccades help redirect the retinal fovea to specific characters and words but that actual discrimination of information only occurs during periods of fixation. Indeed, it has been proposed that there is active and selective suppression of information processing during saccades to avoid experience of blurring due to the high-speed movement. Here, using a paradigm where a string of either lexical (Chinese) or non-lexical (alphabetic) characters are triggered by saccadic eye movements, we show that subjects can discriminate both while making saccadic eye movement. Moreover, discrimination accuracy is significantly better for characters scanned during the saccadic movement to a fixation point than those not scanned beyond it. Our results showed that character information can be processed during the saccade, therefore saccades during reading not only function to redirect the fovea to fixate the next character or word but allow pre-processing of information from the ones adjacent to the fixation locations to help target the next most salient one. In this way saccades can not only promote continuity in reading words but also actively facilitate reading comprehension.
doi:10.1371/journal.pone.0046383
PMCID: PMC3460897  PMID: 23029504
17.  Relationship Between Motor Variability, Accuracy, and Ball Speed in the Tennis Serve 
Journal of Human Kinetics  2012;33:45-53.
The main objective of this study was to analyze the motor variability in the performance of the tennis serve and its relationship to performance outcome. Seventeen male tennis players took part in the research, and they performed 20 serves. Linear and non-linear variability during the hand movement was measured by 3D Motion Tracking. Ball speed was recorded with a sports radar gun and the ball bounces were video recorded to calculate accuracy. The results showed a relationship between the amount of variability and its non-linear structure found in performance of movement and the outcome of the serve. The study also found that movement predictability correlates with performance. An increase in the amount of movement variability could affect the tennis serve performance in a negative way by reducing speed and accuracy of the ball.
doi:10.2478/v10078-012-0043-3
PMCID: PMC3588677  PMID: 23486998
variability; tennis serve; performance
18.  Flexible, task-dependent use of sensory feedback to control hand movements 
We tested whether changing accuracy demands for simple pointing movements leads humans to adjust the feedback control laws that map sensory signals from the moving hand to motor commands. Subjects made repeated pointing movements in a virtual environment to touch a button whose shape varied randomly from trial-to-trial – between squares, rectangles oriented perpendicular to the movement path and rectangles oriented parallel to the movement path. Subjects performed the task on a horizontal table, but saw the target configuration and a virtual rendering of their pointing finger through a mirror mounted between a monitor and the table. On a one-third of trials, the position of the virtual finger was perturbed by ±1 cm either in the movement direction or perpendicular to the movement direction when the finger passed behind an occluder. Subjects corrected quickly for the perturbations despite not consciously noticing them; however, they corrected almost twice as much for perturbations aligned with the narrow dimension of a target than for perturbations aligned with the long dimension. These changes in apparent feedback gain appeared in the kinematic trajectories soon after the time of the perturbations, indicating that they reflect differences in the feedback control law used throughout the duration of movements. The results indicate that the brain adjusts its feedback control law for individual movements “on-demand” to fit task demands. Simulations of optimal control laws for a two-joint arm show that accuracy demands alone, coupled with signal dependent noise lead to qualitatively the same behavior.
doi:10.1523/JNEUROSCI.3522-09.2011
PMCID: PMC3047484  PMID: 21273407
Feedback; optimal control; motor control; pointing; online control
19.  Motor equivalence and self-motion induced by different movement speeds 
This study investigated pointing movements in 3D asking two questions: (1) Is goal-directed reaching accompanied by self-motion, a component of the joint velocity vector that leaves the hand’s movement unaffected? (2) Are differences in the terminal joint configurations among different speeds of reaching motor equivalent (i.e., terminal joint configurations differ more in directions of joint space that do not produce different pointer-tip positions than in directions that do) or non-motor equivalent (i.e., terminal joint configurations differ equally or more in directions of joint space that lead to different pointer-tip positions than in directions that do not affect the pointer-tip position). Subjects reached from an identical starting joint configuration and pointer-tip location to targets at slow, moderate, and fast speeds. Ten degrees of freedom of joint motion of the arm were recorded. The relationship between changes in the joint configuration and the three-dimensional pointer-tip position was expressed by a standard kinematic model, and the range- and null subspaces were computed from the associated Jacobian matrix. (1) The joint velocity vector and (2) the difference vector between terminal joint configurations from pairs of speed conditions were projected into the two subspaces. The relative length of the two components was used to quantify the amount of self-motion and the presence of motor equivalence, respectively. Results revealed that reaches were accompanied by a significant amount of self-motion at all reaching speeds. Self-motion scaled with movement speed. In addition, the difference in the terminal joint configuration between pairs of different reaching speeds revealed motor equivalence. The results are consistent with a control system that takes advantage of motor redundancy, allowing for flexibility in the face of perturbations, here induced by different movement speeds.
doi:10.1007/s00221-011-2541-2
PMCID: PMC3066062  PMID: 21287157
Reaching; Motor Control; Motor equivalence; Movement velocity
20.  Imagined Self-Motion Differs from Perceived Self-Motion: Evidence from a Novel Continuous Pointing Method 
PLoS ONE  2009;4(11):e7793.
Background
The extent to which actual movements and imagined movements maintain a shared internal representation has been a matter of much scientific debate. Of the studies examining such questions, few have directly compared actual full-body movements to imagined movements through space. Here we used a novel continuous pointing method to a) provide a more detailed characterization of self-motion perception during actual walking and b) compare the pattern of responding during actual walking to that which occurs during imagined walking.
Methodology/Principal Findings
This continuous pointing method requires participants to view a target and continuously point towards it as they walk, or imagine walking past it along a straight, forward trajectory. By measuring changes in the pointing direction of the arm, we were able to determine participants' perceived/imagined location at each moment during the trajectory and, hence, perceived/imagined self-velocity during the entire movement. The specific pattern of pointing behaviour that was revealed during sighted walking was also observed during blind walking. Specifically, a peak in arm azimuth velocity was observed upon target passage and a strong correlation was observed between arm azimuth velocity and pointing elevation. Importantly, this characteristic pattern of pointing was not consistently observed during imagined self-motion.
Conclusions/Significance
Overall, the spatial updating processes that occur during actual self-motion were not evidenced during imagined movement. Because of the rich description of self-motion perception afforded by continuous pointing, this method is expected to have significant implications for several research areas, including those related to motor imagery and spatial cognition and to applied fields for which mental practice techniques are common (e.g. rehabilitation and athletics).
doi:10.1371/journal.pone.0007793
PMCID: PMC2771354  PMID: 19907655
21.  Relative performance of the two hands in simple and choice reaction time tasks 
There is evidence that the left hemisphere is more competent for motor control than the right hemisphere. This study investigated whether this hemispheric asymmetry is expressed in the latency/duration of sequential responses performed by the left and/or right hands. Thirty-two right-handed young adults (16 males, 16 females; 18-25 years old) were tested in a simple or choice reaction time task. They responded to a left and/or right visual target by moving their left and/or right middle fingers between two keys on each side of the midline. Right hand reaction time did not differ from left hand reaction time. Submovement times were longer for the right hand than the left hand when the response was bilateral. Pause times were shorter for the right hand than the left hand, both when the responses were unilateral or bilateral. Reaction time results indicate that the putatively more efficient response preparation by the left hemisphere motor mechanisms is not expressed behaviorally. Submovement time and pause time results indicate that the putatively more efficient response execution by the left hemisphere motor mechanisms is expressed behaviorally. In the case of the submovements, the less efficient motor control of the left hand would be compensated by a more intense attention to this hand.
doi:10.1590/1414-431X20132932
PMCID: PMC3932976  PMID: 24345871
Motor lateral asymmetry; Reaction time; Movement time; Response complexity; Unilateral response; Bilateral response
22.  Measurement and Correction of Microscopic Head Motion during Magnetic Resonance Imaging of the Brain 
PLoS ONE  2012;7(11):e48088.
Magnetic resonance imaging (MRI) is a widely used method for non-invasive study of the structure and function of the human brain. Increasing magnetic field strengths enable higher resolution imaging; however, long scan times and high motion sensitivity mean that image quality is often limited by the involuntary motion of the subject. Prospective motion correction is a technique that addresses this problem by tracking head motion and continuously updating the imaging pulse sequence, locking the imaging volume position and orientation relative to the moving brain. The accuracy and precision of current MR-compatible tracking systems and navigator methods allows the quantification and correction of large-scale motion, but not the correction of very small involuntary movements in six degrees of freedom. In this work, we present an MR-compatible tracking system comprising a single camera and a single 15 mm marker that provides tracking precision in the order of 10 m and 0.01 degrees. We show preliminary results, which indicate that when used for prospective motion correction, the system enables improvement in image quality at both 3 T and 7 T, even in experienced and cooperative subjects trained to remain motionless during imaging. We also report direct observation and quantification of the mechanical ballistocardiogram (BCG) during simultaneous MR imaging. This is particularly apparent in the head-feet direction, with a peak-to-peak displacement of 140 m.
doi:10.1371/journal.pone.0048088
PMCID: PMC3492340  PMID: 23144848
23.  NOMAD-Ref: visualization, deformation and refinement of macromolecular structures based on all-atom normal mode analysis 
Nucleic Acids Research  2006;34(Web Server issue):W52-W56.
Normal mode analysis (NMA) is an efficient way to study collective motions in biomolecules that bypasses the computational costs and many limitations associated with full dynamics simulations. The NOMAD-Ref web server presented here provides tools for online calculation of the normal modes of large molecules (up to 100 000 atoms) maintaining a full all-atom representation of their structures, as well as access to a number of programs that utilize these collective motions for deformation and refinement of biomolecular structures. Applications include the generation of sets of decoys with correct stereochemistry but arbitrary large amplitude movements, the quantification of the overlap between alternative conformations of a molecule, refinement of structures against experimental data, such as X-ray diffraction structure factors or Cryo-EM maps and optimization of docked complexes by modeling receptor/ligand flexibility through normal mode motions. The server can be accessed at the URL .
doi:10.1093/nar/gkl082
PMCID: PMC1538881  PMID: 16845062
24.  Evidence for on-line visual guidance during saccadic gaze shifts. 
Rapid orientating movements of the eyes are believed to be controlled ballistically. The mechanism underlying this control is thought to involve a comparison between the desired displacement of the eye and an estimate of its actual position (obtained from the integration of the eye velocity signal). This study shows, however, that under certain circumstances fast gaze movements may be controlled quite differently and may involve mechanisms which use visual information to guide movements prospectively. Subjects were required to make large gaze shifts in yaw towards a target whose location and motion were unknown prior to movement onset. Six of those tested demonstrated remarkable accuracy when making gaze shifts towards a target that appeared during their ongoing movement. In fact their level of accuracy was not significantly different from that shown when they performed a 'remembered' gaze shift to a known stationary target (F3,15 = 0.15, p > 0.05). The lack of a stereotypical relationship between the skew of the gaze velocity profile and movement duration indicates that on-line modifications were being made. It is suggested that a fast route from the retina to the superior colliculus could account for this behaviour and that models of oculomotor control need to be updated.
PMCID: PMC1690195  PMID: 10518326
25.  Motion correction in optical coherence tomography volumes on a per A-scan basis using orthogonal scan patterns 
Biomedical Optics Express  2012;3(6):1182-1199.
High speed Optical Coherence Tomography (OCT) has made it possible to rapidly capture densely sampled 3D volume data. One key application is the acquisition of high quality in vivo volumetric data sets of the human retina. Since the volume is acquired in a few seconds, eye movement during the scan process leads to distortion, which limits the accuracy of quantitative measurements using 3D OCT data. In this paper, we present a novel software based method to correct motion artifacts in OCT raster scans. Motion compensation is performed retrospectively using image registration algorithms on the OCT data sets themselves. Multiple, successively acquired volume scans with orthogonal fast scan directions are registered retrospectively in order to estimate and correct eye motion. Registration is performed by optimizing a large scale numerical problem as given by a global objective function using one dense displacement field for each input volume and special regularization based on the time structure of the acquisition process. After optimization, each volume is undistorted and a single merged volume is constructed that has superior signal quality compared to the input volumes. Experiments were performed using 3D OCT data from the macula and optic nerve head acquired with a high-speed ultra-high resolution 850 nm spectral OCT as well as wide field data acquired with a 1050 nm swept source OCT instrument. Evaluation of registration performance and result stability as well as visual inspection shows that the algorithm can correct for motion in all three dimensions and on a per A-scan basis. Corrected volumes do not show visible motion artifacts. In addition, merging multiple motion corrected and registered volumes leads to improved signal quality. These results demonstrate that motion correction and merging improves image quality and should also improve morphometric measurement accuracy from volumetric OCT data.
doi:10.1364/BOE.3.001182
PMCID: PMC3370961  PMID: 22741067
(170.4500) Optical coherence tomography; (170.4470) Ophthalmology; (100.2980) Image enhancement; (100.5010) Pattern recognition

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