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1.  Magnified visual feedback exacerbates positional variability in older adults due to altered modulation of the primary agonist muscle 
The purpose of this study was to determine whether magnified visual feedback during position-holding contractions exacerbates the age-associated differences in motor output variability due to changes in the neural activation of the agonist muscle in the upper and lower limb. Twelve young (18–35 years) and ten older adults (65–85 years) were instructed to accurately match a target position at 5° of index finger abduction and ankle dorsiflexion while lifting 10 % of their 1 repetition maximum (1RM) load. Position was maintained at three different visual angles (0.1°, 1°, and 4°) that varied across trials. Each trial lasted 25 s and visual feedback of position was removed from 15 to 25 s. Positional error was quantified as the root mean square error (RMSE) of the subject’s performance from the target. Positional variability was quantified as the standard deviation of the position data. The neural activation of the first dorsal interosseus and tibialis anterior was measured with surface electromyography (EMG). Older adults were less accurate compared with young adults and the RMSE decreased significantly with an increase in visual gain. As expected, and independent of limb, older adults exhibited significantly greater positional variability compared with young adults that was exacerbated with magnification of visual feedback (1° and 4°). This increase in variability at the highest magnification of visual feedback was predicted by a decrease in power from 12 to 30 Hz of the agonist EMG signal. These findings demonstrate that motor control in older adults is impaired by magnified visual feedback during positional tasks.
doi:10.1007/s00221-012-3219-0
PMCID: PMC3631577  PMID: 22948735
Aging; Visual feedback; Visual gain; EMG; Motor control
2.  Removal of visual feedback alters muscle activity and reduces force variability during constant isometric contractions 
The purpose of this study was to compare force accuracy, force variability and muscle activity during constant isometric contractions at different force levels with and without visual feedback and at different feedback gains. In experiment 1, subjects were instructed to accurately match the target force at 2, 15, 30, 50, and 70% of their maximal isometric force with abduction of the index finger and maintain their force even in the absence of visual feedback. Each trial lasted 22 s and visual feedback was removed from 8–12 to 16–20 s. Each subject performed 6 trials at each target force, half with visual gain of 51.2 pixels/N and the rest with a visual gain of 12.8 pixels/N. Force error was calculated as the root mean square error of the force trace from the target line. Force variability was quantified as the standard deviation and coefficient of variation (CVF) of the force trace. The EMG activity of the agonist (first dorsal interosseus; FDI) was measured with bipolar surface electrodes placed distal to the innervation zone. Independent of visual gain and force level, subjects exhibited lower force error with the visual feedback condition (2.53 ± 2.95 vs. 2.71 ± 2.97 N; P < 0.01); whereas, force variability was lower when visual feedback was removed (CVF: 4.06 ± 3.11 vs. 4.47 ± 3.14, P < 0.01). The EMG activity of the FDI muscle was higher during the visual feedback condition and this difference increased especially at higher force levels (70%: 370 ± 149 vs. 350 ± 143 μV, P < 0.01). Experiment 2 examined whether the findings of experiment 1 were driven by the higher force levels and proximity in the gain of visual feedback. Subjects performed constant isometric contractions with the abduction of the index finger at an absolute force of 2 N, with two distinct feedback gains of 15 and 3,000 pixels/N. In agreement with the findings of experiment 1, subjects exhibited lower force error in the presence of visual feedback especially when the feedback gain was high (0.057 ± 0.03 vs. 0.095 ± 0.05 N). However, force variability was not affected by the vastly distinct feedback gains at this force, which supported and extended the findings from experiment 1. Our findings demonstrate that although removal of visual feedback amplifies force error, it can reduce force variability during constant isometric contractions due to an altered activation of the primary agonist muscle most likely at moderate force levels in young adults.
doi:10.1007/s00221-009-1883-5
PMCID: PMC3631578  PMID: 19544059
3.  The interaction of respiration and visual feedback on the control of force and neural activation of the agonist muscle 
Human movement science  2011;30(6):1022-1038.
The purpose of this study was to compare force variability and the neural activation of the agonist muscle during constant isometric contractions at different force levels when the amplitude of respiration and visual feedback were varied. Twenty young adults (20–32 years, 10 men and 10 women) were instructed to accurately match a target force at 15 and 50% of their maximal voluntary contraction (MVC) with abduction of the index finger while controlling their respiration at different amplitudes (85, 100 and 125% normal) in the presence and absence of visual feedback. Each trial lasted 22 s and visual feedback was removed from 8–12 to 16–20 s. Each subject performed 3 trials with each respiratory condition at each force level. Force variability was quantified as the standard deviation of the detrended force data. The neural activation of the first dorsal interosseus (FDI) was measured with bipolar surface electrodes placed distal to the innervation zone. Relative to normal respiration, force variability increased significantly only during high-amplitude respiration (~63%). The increase in force variability from normal- to high-amplitude respiration was strongly associated with amplified force oscillations from 0–3 Hz (R2 ranged from .68 – .84; p < .001). Furthermore, the increase in force variability was exacerbated in the presence of visual feedback at 50% MVC (vision vs. no-vision: .97 vs. .87 N) and was strongly associated with amplified force oscillations from 0–1 Hz (R2 = .82) and weakly associated with greater power from 12–30 Hz (R2 = .24) in the EMG of the agonist muscle. Our findings demonstrate that high-amplitude respiration and visual feedback of force interact and amplify force variability in young adults during moderate levels of effort.
doi:10.1016/j.humov.2010.09.007
PMCID: PMC3202062  PMID: 21546109
4.  Greater amount of visual feedback decreases force variability by reducing force oscillations from 0–1 and 3–7 Hz 
The purpose was to determine the relation between visual feedback gain and variability in force and whether visual gain-induced changes in force variability were associated with frequency-specific force oscillations and changes in the neural activation of the agonist muscle. Fourteen young adults (19–29 years) were instructed to accurately match the target force at 2 and 10% of their maximal voluntary contraction with abduction of the index finger. Force was maintained at specific visual feedback gain levels that varied across trials. Each trial lasted 20 s and the amount of visual feedback was varied by changing the visual gain from 0.5 to 1,474 pixels/N (13 levels; equals ~0.001–4.57°). Force variability was quantified as the standard deviation of the detrended force data. The neural activation of the first dorsal interosseus (FDI) was measured with surface electromyography. The mean force did not vary significantly with the amount of visual feedback. In contrast, force variability decreased from low gains compared to moderate gains (0.5–4 pixels/N: 0.09 ± 0.04 vs. 64–1,424 pixels/N: 0.06 ± 0.02 N). The decrease in variability was predicted by a decrease in the power of force oscillations from 0–1 Hz (~50%) and 3–7 Hz (~20%). The activity of the FDI muscle did not vary across the visual feedback gains. These findings demonstrate that in young adults force variability can be decreased with increased visual feedback gain (>64 pixels/N vs. 0.5–4 pixels/N) due to a decrease in the power of oscillations in the force from 0–1 and 3–7 Hz.
doi:10.1007/s00421-009-1301-5
PMCID: PMC2863099  PMID: 19953262
Visual feedback; Visual gain; Force variability; EMG; Force oscillations; First dorsal interosseus
5.  Greater amount of visual information exacerbates force control in older adults during constant isometric contractions 
The purpose of this study was to compare control of force and modulation of agonist muscle activity of young and older adults when the amount of visual feedback was varied at two different force levels. Ten young adults (25 years ± 4 years, 5 men and 5 women) and ten older adults (71 years ± 5 years, 4 men and 6 women) were instructed to accurately match a constant target force at 2 and 30% of their maximal isometric force with abduction of the index finger. Each trial lasted 35 s, and the amount of visual feedback was varied by changing the visual angle at 0.05, 0.5, and 1.5°. Each subject performed three trials for each visual angle condition. Force variability was quantified as the standard deviation and coefficient of variation (CV) of force. Modulation of the agonist muscle activity was quantified as the normalized power spectrum density of the EMG signal recorded from two pairs of bipolar electrodes placed on the first dorsal interosseus muscle. The frequency bands of interest were between 5 and 100 Hz. There were significant age-associated differences in force control with changes in the amount of visual feedback. The CV of force did not change with visual angle for young adults, whereas it increased for older adults. Although older adults exhibited similar CV of force to young adults at 0.05° (5.95 ± 0.67 vs. 5.47 ± 0.5), older adults exhibited greater CV of force than young adults at 0.5° (8.49 ± 1.34 vs. 5.05 ± 0.5) and 1.5° (8.23 ± 1.12 vs. 5.49 ± 0.6). In addition, there were age-associated differences in the modulation of the agonist muscle activity. Young adults increased normalized power in the EMG signal from 13 to 60 Hz with an increase in visual angle, whereas older adults did not. These findings suggest that greater amount of visual information may be detrimental to the control of a constant isometric contraction in older adults, and this impairment may be due to their inability to effectively modulate the motor neuron pool of the agonist muscle.
doi:10.1007/s00221-011-2777-x
PMCID: PMC3631570  PMID: 21800256
Visual gain; EMG; Aging; Force variability
6.  Timing variability and not force variability predicts the endpoint accuracy of fast and slow isometric contractions 
The purpose of the study was to determine the contributions of endpoint variance and trajectory variability to the endpoint accuracy of goal-directed isometric contractions when the target force and contraction speed were varied. Thirteen young adults (25 ± 6 years) performed blocks of 15 trials at each of 2 contraction speeds and 4 target forces. Subjects were instructed to match the peak of a parabolic force trajectory to a target force by controlling the abduction force exerted by the index finger. The time to peak force was either 150 ms (fast) or 1 s (slow). The target forces were 20, 40, 60, and 80% of the maximal force that could be achieved in 150 ms during an MVC. The same absolute forces were required for both contraction speeds. Endpoint accuracy and variability in force and time along with intramuscular EMG activity of the agonist (first dorsal interosseus) and antagonist (second palmar interosseus) muscles were quantified for each block of trials. The principal dependent variables were endpoint error (shortest distance between the coordinates of the target and the peak force), endpoint variance (sum of the variance in peak force and time to peak force), trial-to-trial variability (SD of peak force and time to peak force), SD of the force trajectory (SD of the detrended force from force onset to peak force), normalized peak EMG amplitude, and the SD of normalized peak EMG amplitude. Stepwise multiple linear regression models were used to determine the EMG activity parameters that could explain the differences observed in endpoint error and endpoint variance. Endpoint error increased with target force for the fast contractions, but not for the slow contractions. In contrast, endpoint variance was greatest at the lowest force and was not associated with endpoint error at either contraction speed. Furthermore, force trajectory SD was not associated with endpoint error or endpoint variance for either contraction speed. Only the trial-to-trial variability of the timing predicted endpoint accuracy for fast and slow contractions. These findings indicate that endpoint error in tasks that require force and timing accuracy is minimized by controlling timing variability but not force variability, and that endpoint error is not related to the amplitude of the activation signal.
doi:10.1007/s00221-009-2126-5
PMCID: PMC3631579  PMID: 20033680
Hand; First dorsal interosseus; Force control; Neural noise
7.  Aging interferes central control mechanism for eccentric muscle contraction 
Previous studies report greater activation in the cortical motor network in controlling eccentric contraction (EC) than concentric contraction (CC) despite lower muscle activation level associated with EC vs. CC in healthy, young individuals. It is unknown, however, whether elderly people exhibiting increased difficulties in performing EC than CC possess this unique cortical control mechanism for EC movements. To address this question, we examined functional magnetic resonance imaging (fMRI) data acquired during EC and CC of the first dorsal interosseous (FDI) muscle in 11 young (20–32 years) and 9 old (67–73 years) individuals. During the fMRI experiment, all subjects performed 20 CC and 20 EC of the right FDI with the same angular distance and velocity. The major findings from the behavioral and fMRI data analysis were that (1) movement stability was poorer in EC than CC in the old but not the young group; (2) similar to previous electrophysiological and fMRI reports, the EC resulted in significantly stronger activation in the motor control network consisting of primary, secondary and association motor cortices than CC in the young and old groups; (3) the biased stronger activation towards EC was significantly greater in the old than the young group especially in the secondary and association cortices such as supplementary and premotor motor areas and anterior cingulate cortex; and (4) in the primary motor and sensory cortices, the biased activation towards EC was significantly greater in the young than the old group. Greater activation in higher-order cortical fields for controlling EC movement by elderly adults may reflect activities in these regions to compensate for aging-related impairments in the ability to control complex EC movements. Our finding is useful for potentially guiding the development of targeted therapies to counteract age-related movement deficits and to prevent injury.
doi:10.3389/fnagi.2014.00086
PMCID: PMC4023019  PMID: 24847261
brain activation; concentric contraction; eccentric contraction; fMRI; movement stability
8.  Multi-digit coordination during lifting a horizontally oriented object: synergies control with referent configurations 
We explored digit coordination during the acceleration phase of a quick lifting movement of a hand-held horizontal object. We tested three hypotheses related to: (1) the scaling of mechanical variables produced by the hand with changes in the external load, torque, and moment of inertia; (2) changes in the safety margin for the thumb with both the loading conditions and acceleration; and (3) changes in the indices of synergies. The subjects held a horizontal handle with a prismatic grasp (the thumb acted on top of the handle) and performed series of “very quick” lifting movements to a visual target. Multi-digit synergies were quantified as co-variation indices among elemental variables (forces and moments produced by individual digits). The resultant force scaled with the external load but not torque, while the grip force scaled with the external torque but not load. The safety margin dropped with an increase in acceleration; it also showed changes with the external torque and moment of inertia. Total moment of force was primarily produced by the tangential forces (over 80 %) across all movement phases and loading conditions. The index and little fingers produced close to zero moment with their normal forces, while the middle and ring fingers produced consistent moments due to the reproducible shifts of their centers of pressure. Synergy indices at the upper level of the assumed hierarchy (the task is shared between the thumb and virtual finger—an imagined digit with the action equal to that of the four fingers combined) did not drop with acceleration for the three force vector components and one of the moment vector components. They did drop with acceleration at the lower level (virtual finger action is shared among the four fingers). There was a trade-off between synergy indices computed at the two levels for the three force vector components, but not for the moment of force components. We confirmed specialization of different fingers with respect to different task components in quick manipulation tasks. The findings have implications for hypotheses on the control of voluntary movements involving redundant sets of effectors. Within the referent configuration hypothesis, components of a referent configuration may be adjusted to task mechanical characteristics using simple scaling rules. The neural organization of multi-digit synergies in a hierarchal system is able to selectively protect synergies related to stabilization of some performance variables from detrimental effects of the rate of change of those variables. A large number of apparently redundant elemental variables are not the source of additional computational problems but may be beneficial, allowing the central nervous system to facilitate synergies at both levels of the hierarchy.
doi:10.1007/s00221-012-3215-4
PMCID: PMC3465706  PMID: 22910900
Prehension; Hand; Synergy; Safety margin
9.  Hand Function is Altered in Individuals with a History of Illicit Stimulant Use 
PLoS ONE  2014;9(12):e115771.
Use of illicit stimulant drugs such as methamphetamine, cocaine, and ecstasy are a significant worldwide problem. However, little is known about the effect of these drugs on movement. The aim of the current study was to investigate hand function in adults with a history of illicit stimulant use. We hypothesized that prior use of illicit stimulant drugs is associated with abnormal manipulation of objects. The study involved 22 subjects with a history of illicit stimulant use (aged 29±8 yrs; time since last use: 1.8±4.0 yrs) and two control groups comprising 27 non-drug users (aged 25±8 yrs) and 17 cannabis users with no history of stimulant use (aged 22±5 yrs). Each subject completed screening tests (neuropsychological assessment, medical history questionnaire, lifetime drug history questionnaire, and urine drug screen) prior to gripping and lifting a light-weight object with the dominant right hand. Horizontal grip force, vertical lift force, acceleration, and first dorsal interosseus electromyographic (EMG) activity were recorded during three trials. In trial one, peak grip force was significantly greater in the stimulant group (12.8±3.9 N) than in the control groups (non-drug: 10.3±4.6 N; cannabis: 9.4±2.9 N, P<0.022). However, peak grip force did not differ between groups in trials two and three. The results suggest that individuals with a history of stimulant use overestimate the grip force required to manipulate a novel object but, are able to adapt grip force in subsequent lifts. The results suggest that movement dysfunction may be an unrecognized consequence of illicit stimulant use.
doi:10.1371/journal.pone.0115771
PMCID: PMC4278704  PMID: 25545892
10.  Safety margins in older adults increase with improved control of a dynamic object 
Older adults face decreasing motor capabilities due to pervasive neuromuscular degradations. As a consequence, errors in movement control increase. Thus, older individuals should maintain larger safety margins than younger adults. While this has been shown for object manipulation tasks, several reports on whole-body activities, such as posture and locomotion, demonstrate age-related reductions in safety margins. This is despite increased costs for control errors, such as a fall. We posit that this paradox could be explained by the dynamic challenge presented by the body or also an external object, and that age-related reductions in safety margins are in part due to a decreased ability to control dynamics. To test this conjecture we used a virtual ball-in-cup task that had challenging dynamics, yet afforded an explicit rendering of the physics and safety margin. The hypotheses were: (1) When manipulating an object with challenging dynamics, older adults have smaller safety margins than younger adults. (2) Older adults increase their safety margins with practice. Nine young and 10 healthy older adults practiced moving the virtual ball-in-cup to a target location in exactly 2 s. The accuracy and precision of the timing error quantified skill, and the ball energy relative to an escape threshold quantified the safety margin. Compared to the young adults, older adults had increased timing errors, greater variability, and decreased safety margins. With practice, both young and older adults improved their ability to control the object with decreased timing errors and variability, and increased their safety margins. These results suggest that safety margins are related to the ability to control dynamics, and may explain why in tasks with simple dynamics older adults use adequate safety margins, but in more complex tasks, safety margins may be inadequate. Further, the results indicate that task-specific training may improve safety margins in older adults.
doi:10.3389/fnagi.2014.00158
PMCID: PMC4094928  PMID: 25071566
aging; safety margin; human; dynamics; object manipulation; motor control; motor learning
11.  Time to Task Failure and Motor Cortical Activity Depend on the Type of Feedback in Visuomotor Tasks 
PLoS ONE  2012;7(3):e32433.
The present study aimed to elucidate whether the type of feedback influences the performance and the motor cortical activity when executing identical visuomotor tasks. For this purpose, time to task failure was measured during position- and force-controlled muscular contractions. Subjects received either visual feedback about the force produced by pressing a force transducer or about the actual position between thumb and index finger. Participants were instructed to either match the force level of 30% MVC or the finger position corresponding to the thumb and index finger angle at this contraction intensity. Subjects demonstrated a shorter time to task failure when they were provided with feedback about their joint position (11.5±6.2 min) instead of force feedback (19.2±12.8 min; P = 0.01). To test differences in motor cortical activity between position- and force-controlled contractions, subthreshold transcranial magnetic stimulation (subTMS) was applied while executing submaximal (20% MVC) contractions. SubTMS resulted in a suppression of the first dorsal interosseus muscle (FDI) EMG in both tasks. However, the mean suppression for the position-controlled task was significantly greater (18.6±9.4% vs. 13.3±7.5%; P = 0.025) and lasted longer (13.9±7.5 ms vs. 9.3±4.3 ms; P = 0.024) compared to the force-controlled task. The FDI background EMG obtained without stimulation was comparable in all conditions. The present results demonstrate that the presentation of different feedback modalities influences the time to task failure as well as the cortical activity. As only the feedback was altered but not the mechanics of the task, the present results add to the body of evidence that suggests that the central nervous system processes force and position information in different ways.
doi:10.1371/journal.pone.0032433
PMCID: PMC3302870  PMID: 22427836
12.  Age-related differences in corticospinal excitability during a choice reaction time task 
Age  2012;35(5):1705-1719.
Age-related declines in central processing may affect corticospinal (CS) excitability that underlies the emergence of voluntary responses to external stimuli. We used single-pulse transcranial magnetic stimulation (TMS) over the primary motor cortex to explore the evolution of CS excitability in 14 young and ten elderly healthy right-handed participants. Motor-evoked potentials (MEPs) were elicited in the right or left first dorsal interosseus (FDI) during the preparatory and premotor periods of a choice reaction time (CRT) task, which required selection of left or right index finger responses. Both age groups showed significant suppression of CS excitability in the preparatory period. However, suppression was generally less pronounced in older than in young adults. Moreover, our data indicated that a reduced suppression in the right FDI during the preparatory period was associated with longer reaction times (RTs) in older adults only. In the premotor period, both age groups demonstrated comparable facilitation levels towards movement onset. Our findings indicate that increased RTs among older individuals could be directly associated with declines in preparatory processes.
doi:10.1007/s11357-012-9471-1
PMCID: PMC3776102  PMID: 23007962
Aging; Choice reaction time; Response preparation; Transcranial magnetic stimulation; TMS
13.  Grasping the Changes Seen in Older Adults When Reaching for Objects of Varied Texture 
PLoS ONE  2013;8(7):e69040.
Old age is associated with reduced mobility of the hand. To investigate age related decline when reaching-to-lift an object we used sophisticated kinematic apparatus to record reaches carried out by healthy older and younger participants. Three objects of different widths were placed at three different distances, with objects having either a high or low friction surface (i.e. rough or slippery). Older participants showed quantitative differences to their younger counterparts – movements were slower and peak speed did not scale with object distance. There were also qualitative differences with older adults showing a greater propensity to stop the hand and adjust finger position before lifting objects. The older participants particularly struggled to lift wide slippery objects, apparently due to an inability to manipulate their grasp to provide the level of precision necessary to functionally enclose the object. These data shed light on the nature of age related changes in reaching-to-grasp movements and establish a powerful technique for exploring how different product designs will impact on prehensile behavior.
doi:10.1371/journal.pone.0069040
PMCID: PMC3729566  PMID: 23935918
14.  Prehension synergies and control with referent hand configurations 
We used the framework of the equilibrium-point hypothesis (in its updated form based on the notion of referent configuration) to investigate the multi-digit synergies at two levels of a hypothetical hierarchy involved in prehensile actions. Synergies were analyzed at the thumb–virtual finger (VF) level (VF is an imaginary digit with the mechanical action equivalent to that of the four actual fingers) and at the individual finger level. The subjects performed very quick vertical movements of a handle into a target. A load could be attached off-center to provide a pronation or supination torque. In a few trials, the handle was unexpectedly fixed to the table and the digits slipped off the sensors. In such trials, the hand stopped at a higher vertical position and rotated into pronation or supination depending on the expected torque. The aperture showed non-monotonic changes with a large, fast decrease and further increase, ending up with a smaller distance between the thumb and the fingers as compared to unperturbed trials. Multi-digit synergies were quantified using indices of co-variation between digit forces and moments of force across unperturbed trials. Prior to the lifting action, high synergy indices were observed at the individual finger level while modest indices were observed at the thumb–VF level. During the lifting action, the synergies at the individual finger level disappeared while the synergy indices became higher at the thumb–VF level. The results support the basic premise that, within a given task, setting a referent configuration may be described with a few referent values of variables that influence the equilibrium state, to which the system is attracted. Moreover, the referent configuration hypothesis can help interpret the data related to the trade-off between synergies at different hierarchical levels.
doi:10.1007/s00221-009-2128-3
PMCID: PMC2846225  PMID: 20033397
Prehension; Synergy; Referent configuration; Grip
15.  Movement trajectory smoothness is not associated with the endpoint accuracy of rapid multi-joint arm movements in young and older adults 
Acta psychologica  2013;143(2):157-167.
The minimum variance theory proposes that motor commands are corrupted by signal-dependent noise and smooth trajectories with low noise levels are selected to minimize endpoint error and endpoint variability. The purpose of the study was to determine the contribution of trajectory smoothness to the endpoint accuracy and endpoint variability of rapid multi-joint arm movements. Young and older adults performed arm movements (4 blocks of 25 trials) as fast and as accurately as possible to a target with the right (dominant) arm. Endpoint accuracy and endpoint variability along with trajectory smoothness and error were quantified for each block of trials. Endpoint error and endpoint variance were greater in older adults compared with young adults, but decreased at a similar rate with practice for the two age groups. The greater endpoint error and endpoint variance exhibited by older adults were primarily due to impairments in movement extent control and not movement direction control. The normalized jerk was similar for the two age groups, but was not strongly associated with endpoint error or endpoint variance for either group. However, endpoint variance was strongly associated with endpoint error for both the young and older adults. Finally, trajectory error was similar for both groups and was weakly associated with endpoint error for the older adults. The findings are not consistent with the predictions of the minimum variance theory, but support and extend previous observations that movement trajectories and endpoints are planned independently.
doi:10.1016/j.actpsy.2013.02.011
PMCID: PMC3677705  PMID: 23584101
aging; neural noise; skill; practice
16.  Reflecting on Mirror Mechanisms: Motor Resonance Effects during Action Observation Only Present with Low-Intensity Transcranial Magnetic Stimulation 
PLoS ONE  2013;8(5):e64911.
Transcranial magnetic stimulation (TMS) studies indicate that the observation of other people's actions influences the excitability of the observer's motor system. Motor evoked potential (MEP) amplitudes typically increase in muscles which would be active during the execution of the observed action. This ‘motor resonance’ effect is thought to result from activity in mirror neuron regions, which enhance the excitability of the primary motor cortex (M1) via cortico-cortical pathways. The importance of TMS intensity has not yet been recognised in this area of research. Low-intensity TMS predominately activates corticospinal neurons indirectly, whereas high-intensity TMS can directly activate corticospinal axons. This indicates that motor resonance effects should be more prominent when using low-intensity TMS. A related issue is that TMS is typically applied over a single optimal scalp position (OSP) to simultaneously elicit MEPs from several muscles. Whether this confounds results, due to differences in the manner that TMS activates spatially separate cortical representations, has not yet been explored. In the current study, MEP amplitudes, resulting from single-pulse TMS applied over M1, were recorded from the first dorsal interosseous (FDI) and abductor digiti minimi (ADM) muscles during the observation of simple finger abductions. We tested if the TMS intensity (110% vs. 130% resting motor threshold) or stimulating position (FDI-OSP vs. ADM-OSP) influenced the magnitude of the motor resonance effects. Results showed that the MEP facilitation recorded in the FDI muscle during the observation of index-finger abductions was only detected using low-intensity TMS. In contrast, changes in the OSP had a negligible effect on the presence of motor resonance effects in either the FDI or ADM muscles. These findings support the hypothesis that MN activity enhances M1 excitability via cortico-cortical pathways and highlight a methodological framework by which the neural underpinnings of action observation can be further explored.
doi:10.1371/journal.pone.0064911
PMCID: PMC3665751  PMID: 23724104
17.  Fluctuations in motor output of a hand muscle can be altered by the mechanical properties of the position sensor 
Journal of neuroscience methods  2007;168(1):164-173.
Fluctuations in motor output are typically quantified by the standard deviation (SD) of displacement or acceleration. The aim of the study was to determine the influence of a linear variable-displacement transducer (LVDT) on the SDs and spectral content of displacement and acceleration during steady isometric and anisometric contractions performed with the first dorsal interosseus muscle. Thirteen young adults supported six loads when performing position-holding and position-tracking tasks when the LVDT either was or was not attached to the index finger. The LVDT reduced the magnitude of the SDs in displacement and acceleration and disrupted the load-dependent modulation of the spectral properties of these signals. When the LVDT was not connected to the finger, the displacement SD was greatest during concentric contractions, the acceleration SD was greatest during eccentric contractions, and there were load-dependent changes in the power density spectra. Although the LVDT may be used for assessing relative changes in displacement, its ability to provide absolute measures of fluctuations in motor output is limited. The results provide baseline measures of the fluctuations in motor output during steady contractions with a hand muscle and how the method used to detect displacement alters these measures.
doi:10.1016/j.jneumeth.2007.09.015
PMCID: PMC2253721  PMID: 17964659
First dorsal interosseus muscle; Steadiness; Displacement; Acceleration
18.  Transcranial magnetic stimulation coregistered with MRI: a comparison of a guided versus blind stimulation technique and its effect on evoked compound muscle action potentials 
Introduction and methods
Compound muscle action potentials (CMAPs) elicited by transcranial magnetic stimulation (TMS) are characterized by enormous variability, even when attempts are made to stimulate the same scalp location. This report describes the results of a comparison of the spatial errors in coil placement and resulting CMAP characteristics using a guided and blind TMS stimulation technique. The former uses a coregistration system, which displays the intersection of the peak TMS induced electric field with the cortical surface. The latter consists of the conventional placement of the TMS coil on the optimal scalp position for activation of the first dorsal interossei (FDI) muscle.
Results
Guided stimulation resulted in significantly improved spatial precision for exciting the corticospinal projection to the FDI compared to blind stimulation. This improved precision of coil placement was associated with a significantly increased probability of eliciting FDI responses. Although these responses tended to have larger amplitudes and areas, the coefficient of variation between guided and blind stimulation induced CMAPs did not significantly differ.
Conclusion
The results of this study demonstrate that guided stimulation improves the ability to precisely revisit previously stimulated cortical loci as well as increasing the probability of eliciting TMS induced CMAPs. Response variability, however, is due to factors other than coil placement.
PMCID: PMC2845153  PMID: 11595135
Transcranial magnetic stimulation and coregistration system; Compound muscle action potentials; Variability
19.  Involuntary paretic wrist/finger flexion forces and EMG increase with shoulder abduction load in individuals with chronic stroke 
Objective
Clinical observations of the flexion synergy in individuals with chronic hemiparetic stroke describe coupling of shoulder, elbow, wrist, and finger joints. Yet, experimental quantification of the synergy within a shoulder abduction (SABD) loading paradigm has focused only on shoulder and elbow joints. The paretic wrist and fingers have typically been studied in isolation. Therefore, this study quantified involuntary behavior of paretic wrist and fingers during concurrent activation of shoulder and elbow.
Methods
Eight individuals with chronic moderate-to-severe hemiparesis and four controls participated. Isometric wrist/finger and thumb flexion forces and wrist/finger flexor and extensor electromyograms (EMG) were measured at two positions when lifting the arm: in front of the torso and at maximal reaching distance. The task was completed in the ACT3D robotic device with six SABD loads by paretic, non-paretic, and control limbs.
Results
Considerable forces and EMG were generated during lifting of the paretic arm only, and they progressively increased with SABD load. Additionally, the forces were greater at the maximal reach position than at the position front of the torso.
Conclusions
Flexion of paretic wrist and fingers is involuntarily coupled with certain shoulder and elbow movements.
Significance
Activation of the proximal upper limb must be considered when seeking to understand, rehabilitate, or develop devices to assist the paretic hand.
doi:10.1016/j.clinph.2012.01.009
PMCID: PMC3729226  PMID: 22364723
20.  Corticospinal Excitability Modulation During Action Observation 
This study used the transcranial magnetic stimulation/motor evoked potential (TMS/MEP) technique to pinpoint when the automatic tendency to mirror someone else's action becomes anticipatory simulation of a complementary act. TMS was delivered to the left primary motor cortex corresponding to the hand to induce the highest level of MEP activity from the abductor digiti minimi (ADM; the muscle serving little finger abduction) as well as the first dorsal interosseus (FDI; the muscle serving index finger flexion/extension) muscles. A neuronavigation system was used to maintain the position of the TMS coil, and electromyographic (EMG) activity was recorded from the right ADM and FDI muscles. Producing original data with regard to motor resonance, the combined TMS/MEP technique has taken research on the perception-action coupling mechanism a step further. Specifically, it has answered the questions of how and when observing another person's actions produces motor facilitation in an onlooker's corresponding muscles and in what way corticospinal excitability is modulated in social contexts.
doi:10.3791/51001
PMCID: PMC4093910  PMID: 24429584
Behavior; Issue 82; action observation; transcranial magnetic stimulation; motor evoked potentials; corticospinal excitability
21.  Interhemispheric Plasticity in Humans 
Introduction
Chronic unimanual motor practice increases the motor output not only in the trained but also in the non-exercised homologous muscle in the opposite limb. We examined the hypothesis that adaptations in motor cortical excitability of the non-trained primary motor cortex (iM1) and in interhemispheric inhibition (IHI) from the trained to the non-trained M1 mediate this inter-limb cross education.
Methods
Healthy, young volunteers (n = 12) performed 1000 submaximal voluntary contractions (MVC) of the right first dorsal interosseus (FDI) at 80% MVC over 20 sessions.
Results
Trained FDI’s MVC increased 49.9% and the untrained FDI’s MVC increased 28.1%. Although corticospinal excitability in iM1, measured with magnetic brain stimulation (TMS) before and after every 5th session, increased 6% at rest, these changes, as those in intracortical inhibition and facilitation, did not correlate with cross education. When weak and strong TMS of iM1 were, respectively, delivered on a background of a weak and strong muscle contraction of the right FDI, excitability of iM1 increased dramatically after 20 sessions. IHI decreased 8.9% acutely within sessions and 30.9% chronically over 20 sessions and these chronic reductions progressively became more strongly associated with cross education. There were no changes in force or TMS measures in the trained group’s left ADM nor were there changes in a non-exercising control group (n = 8).
Conclusions
The findings provide the first evidence for plasticity of interhemispheric connections to mediate cross education produced by a simple motor task.
doi:10.1249/MSS.0b013e31820a94b8
PMCID: PMC4137570  PMID: 21200340
muscle; force; motor cortex; excitability; motor control
22.  Analysis of squat and stoop dynamic liftings: muscle forces and internal spinal loads 
European Spine Journal  2006;16(5):687-699.
Despite the well-recognized role of lifting in back injuries, the relative biomechanical merits of squat versus stoop lifting remain controversial. In vivo kinematics measurements and model studies are combined to estimate trunk muscle forces and internal spinal loads under dynamic squat and stoop lifts with and without load in hands. Measurements were performed on healthy subjects to collect segmental rotations during lifts needed as input data in subsequent model studies. The model accounted for nonlinear properties of the ligamentous spine, wrapping of thoracic extensor muscles to take curved paths in flexion and trunk dynamic characteristics (inertia and damping) while subject to measured kinematics and gravity/external loads. A dynamic kinematics-driven approach was employed accounting for the spinal synergy by simultaneous consideration of passive structures and muscle forces under given posture and loads. Results satisfied kinematics and dynamic equilibrium conditions at all levels and directions. Net moments, muscle forces at different levels, passive (muscle or ligamentous) forces and internal compression/shear forces were larger in stoop lifts than in squat ones. These were due to significantly larger thorax, lumbar and pelvis rotations in stoop lifts. For the relatively slow lifting tasks performed in this study with the lowering and lifting phases each lasting ∼2 s, the effect of inertia and damping was not, in general, important. Moreover, posterior shift in the position of the external load in stoop lift reaching the same lever arm with respect to the S1 as that in squat lift did not influence the conclusion of this study on the merits of squat lifts over stoop ones. Results, for the tasks considered, advocate squat lifting over stoop lifting as the technique of choice in reducing net moments, muscle forces and internal spinal loads (i.e., moment, compression and shear force).
doi:10.1007/s00586-006-0240-7
PMCID: PMC2213554  PMID: 17103232
Muscle force; Finite element; Dynamic; Kinematics; Lifting technique
23.  Ulnar Nerve Conduction Study of the First Dorsal Interosseous Muscle in Korean Subjects 
Annals of Rehabilitation Medicine  2011;35(5):658-663.
Objective
To derive normative values for ulnar nerve conduction study of the active recording electrode on the first dorsal interosseous muscle (FDI) and the reference electrode on the proximal phalanx of the thumb.
Method
Ulnar nerve motor conduction study with FDI and abductor digiti minimi muscle (ADM) recording was performed in 214 hands of 107 healthy subjects. Ulnar NCS was performed with 2 different recording electrode montages (ADM-base of 5th finger; FDI-thumb) and differences in latency and amplitude were compared. Using this technique, the initial positivity of ulnar compound muscle action potential (CMAP) was not observed in any response with FDI recording.
Results
The maximal values for distal motor latency to the ADM and FDI muscle were 3.8 ms and 4.4 ms, respectively. The maximal difference of distal motor latency between the bilateral FDI recordings was 0.6 ms. The maximal ipsilateral latency difference between ADM and FDI was 1.4 ms.
Conclusion
Placement of the reference electrode on the thumb results in a CMAP without an initial positivity and the normative values obtained may be useful in the diagnosis of ulnar neuropathy at the wrist.
doi:10.5535/arm.2011.35.5.658
PMCID: PMC3309257  PMID: 22506188
Nerve conduction; Reference electrode; Thumb; Ulnar neuropathies; Wrist
24.  Effects of Aging and Tai Chi on a Finger-Pointing Task with a Choice Paradigm 
Background. This cross-sectional study examined the effect of aging on performing finger-pointing tasks involving choices and whether experienced older Tai Chi practitioners perform better than healthy older controls in such tasks. Methods. Thirty students and 30 healthy older controls were compared with 31 Tai Chi practitioners. All the subjects performed a rapid index finger-pointing task. The visual signal appeared randomly under 3 conditions: (1) to touch a black ball as quickly and as accurately as possible, (2) not to touch a white ball, (3) to touch only the white ball when a black and a white ball appeared simultaneously. Reaction time (RT) of anterior deltoid electromyogram, movement time (MT) from electromyogram onset to touching of the target, end-point accuracy from the center of the target, and the number of wrong movements were recorded. Results. Young students displayed significantly faster RT and MT, achieving significantly greater end-point accuracy and fewer wrong movements than older controls. Older Tai Chi practitioners had significantly faster MT than older controls. Conclusion. Finger-pointing tasks with a choice paradigm became slower and less accurate with age. Positive findings suggest that Tai Chi may slow down the aging effect on eye-hand coordination tasks involving choices that require more cognitive progressing.
doi:10.1155/2013/653437
PMCID: PMC3588207  PMID: 23476699
25.  The Effects of Parkinson's Disease and Age on Syncopated Finger Movements 
Brain research  2009;1290:12-20.
In young healthy adults, syncopated finger movements (movements between consecutive beats) are characterized by a frequency-dependent change in phase at movement rates near 2 Hz. A similar frequency-dependent phase transition is observed during bimanual anti-phase (asymmetric) tasks in healthy young adults, but this transition frequency is significantly lowered in both patients with Parkinson's disease (PD) and older adults. To date, no study has examined the transition frequency associated with unimanual syncopated movements in patients with PD or older adults. This study examined the effects of movement frequency on the performance of unconstrained syncopated index finger flexion movements in patients with PD, older adult subjects matched to patients with PD, and young adult subjects. Syncopated movements were paced by an acoustic tone that increased in frequency from 1 to 3 Hz in 0.25 Hz increments. Movement phase was quantified and the movement frequency where subjects transitioned from syncopation to synchronization was compared between groups. The principal finding was a marked impairment in the ability of patients with PD to perform syncopated movements when off medication. Medication did not significantly improve performance. In addition, the transition frequency for older adult subjects was lower than young adult subjects. These findings demonstrate that, similar to bimanual tasks, the coordination dynamics associated with unimanual syncopated finger movements transition from a stable to an unstable pattern at significantly lower frequencies in patients with PD and older adults compared to young adults.
doi:10.1016/j.brainres.2009.06.099
PMCID: PMC2747317  PMID: 19596277
syncopation; aging; Parkinson's disease; movement frequency

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