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1.  Slowed Reaction Time During Exercise: What is the Mechanism? 
doi:10.1097/JES.0b013e31828aade3
PMCID: PMC3661292  PMID: 23392281
2.  Increased Force Variability in Chronic Stroke: Contributions of Force Modulation below 1 Hz 
PLoS ONE  2013;8(12):e83468.
Increased force variability constitutes a hallmark of arm disabilities following stroke. Force variability is related to the modulation of force below 1 Hz in healthy young and older adults. However, whether the increased force variability observed post stroke is related to the modulation of force below 1 Hz remains unknown. Thus, the purpose of this study was to compare force modulation below 1 Hz in chronic stroke and age-matched healthy individuals. Both stroke and control individuals (N = 26) performed an isometric grip task to submaximal force levels. Coefficient of variation quantified force variability, and power spectrum density of force quantified force modulation below 1 Hz with a high resolution (0.07 Hz). Analyses indicated that force variability was greater for the stroke group compared with to healthy controls and for the paretic hand compared with the non-paretic hand. Force modulation below 1 Hz differentiated the stroke individuals and healthy controls, as well as the paretic and non-paretic hands. Specifically, stroke individuals (paretic hand) exhibited greater power ∼0.2 Hz (0.07–0.35 Hz) and lesser power ∼0.6 Hz (0.49–0.77 Hz) compared to healthy controls (non-dominant hand). Similarly, the paretic hand exhibited greater power ∼0.2 Hz, and lesser power ∼0.6 Hz than the non-paretic hand. Moreover, variability of force was strongly predicted from the modulation of specific frequencies below 1 Hz (R2 = 0.80). Together, these findings indicate that the modulation of force below 1 Hz provides significant insight into changes in motor control after stroke.
doi:10.1371/journal.pone.0083468
PMCID: PMC3873339  PMID: 24386208
3.  Discharge rate modulation of trapezius motor units differs for voluntary contractions and instructed muscle rest 
This study examined discharge rate modulation at respiratory (0–0.5 Hz) and beta (16–32 Hz) frequencies in trapezius motor units active during voluntary contractions and during periods of instructed rest under conditions of low and high psychosocial stress. In separate sessions, single motor unit activity was recorded from the trapezius muscle of healthy women during low-intensity voluntary contractions and during periods of instructed muscle rest that followed voluntary contractions. The level of psychosocial stress during periods of instructed muscle rest was manipulated using a verbal math task combined with social evaluative threat which increased perceived anxiety, heart rate, and blood pressure (P ≤ 0.002). Discharge rate modulation was quantified by the mean power of motor unit discharge rate profiles within frequency bands of interest. Under low stress conditions, motor units active during instructed rest had greater power at 0–0.5 Hz (P = 0.002) and less power at 16–32 Hz (P = 0.009) compared to those active during voluntary contraction. Exposure to the stressor increased the amount of motor unit activity during instructed rest (P = 0.021) but did not alter the power of discharge rate modulation at 0–0.5 Hz (P = 0.391) or 16–32 Hz (P = 0.089). These results indicate that sustained motor unit activity during periods of instructed muscle rest has a lesser contribution from inputs at beta frequencies and a greater contribution from inputs at respiratory frequencies than present during low-intensity voluntary contractions. Furthermore, increases in motor unit activity when exposed to stressors during periods of instructed rest are not caused by changes in inputs at respiratory or beta frequencies.
doi:10.1007/s00221-010-2471-4
PMCID: PMC3764997  PMID: 21063691
Motor unit; Trapezius; Beta oscillation; Respiration; Stress; Electromyography
4.  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
5.  INCREASED VOLUNTARY DRIVE IS ASSOCIATED WITH CHANGES IN COMMON OSCILLATIONS FROM 13 TO 60 HZ OF INTERFERENCE BUT NOT RECTIFIED ELECTROMYOGRAPHY 
Muscle & nerve  2010;42(3):348-354.
The purpose of this study was to compare the capability of interference and rectified electromyography (EMG) to detect changes in the beta (13–30-HZ) and Piper (30–60-HZ) bands when voluntary force is increased. Twenty adults exerted a constant force abduction of the index finger at 15% and 50% of maximum. The common oscillations at various frequency bands (0–500 HZ) were estimated from the first dorsal interosseous muscle using cross wavelets of interference and rectified EMG. For the interference EMG signals, normalized power significantly (P < 0.01) increased with force in the beta (9.0 ± 0.9 vs. 15.5 ± 2.1%) and Piper (13.6 ± 0.9 vs. 21 ± 1.7%) bands. For rectified EMG signals, however, the beta and Piper bands remained unchanged (P > 0.4). Although rectified EMG is used in many clinical studies to identify changes in the oscillatory drive to the muscle, our findings suggest that only interference EMG can accurately capture the increase in oscillatory drive from 13 to 60 HZ with voluntary force.
doi:10.1002/mus.21687
PMCID: PMC3631571  PMID: 20589885
beta band; cross-wavelet; oscillations; Piper band; rectification
6.  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
7.  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
8.  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
9.  Aging and Variability of Voluntary Contractions 
Older adults exhibit greater motor variability, which impairs their accuracy and function, compared with young adults. Low-intensity training that emphasizes muscle coordination reduces variability in older adults. Furthermore, low amount of visual feedback minimizes age-associated differences in variability. We hypothesize that an intervention that combines muscle coordination and reduced visual feedback would be advantageous to improve motor control in older adults.
doi:10.1097/JES.0b013e31820b85ab
PMCID: PMC3631580  PMID: 21206281
neuromuscular control; force variability; movement control; aging; intervention; training
10.  Modulation of Force below 1 Hz: Age-Associated Differences and the Effect of Magnified Visual Feedback 
PLoS ONE  2013;8(2):e55970.
Oscillations in force output change in specific frequency bins and have important implications for understanding aging and pathological motor control. Although previous studies have demonstrated that oscillations from 0–1 Hz can be influenced by aging and visuomotor processing, these studies have averaged power within this bandwidth and not examined power in specific frequencies below 1 Hz. The purpose was to determine whether a differential modulation of force below 1 Hz contributes to changes in force control related to manipulation of visual feedback and aging. Ten young adults (25±4 yrs, 5 men) and ten older adults (71±5 yrs, 4 men) were instructed to accurately match a target force at 2% of their maximal isometric force for 35 s with abduction of the index finger. Visual feedback was manipulated by changing the visual angle (0.05°, 0.5°, 1.5°) or removing it after 15 s. Modulation of force below 1 Hz was quantified by examining the absolute and normalized power in seven frequency bins. Removal of visual feedback increased normalized power from 0–0.33 Hz and decreased normalized power from 0.66–1.0 Hz. In contrast, magnification of visual feedback (visual angles of 0.5° and 1.5°) decreased normalized power from 0–0.16 Hz and increased normalized power from 0.66–1.0 Hz. Older adults demonstrated a greater increase in the variability of force with magnification of visual feedback compared with young adults (P = 0.05). Furthermore, older adults exhibited differential force modulation of frequencies below 1 Hz compared with young adults (P<0.05). Specifically, older adults exhibited greater normalized power from 0–0.16 Hz and lesser normalized power from 0.66–0.83 Hz. The changes in force modulation predicted the changes in the variability of force with magnification of visual feedback (R2 = 0.80). Our findings indicate that force oscillations below 1 Hz are associated with force control and are modified by aging and visual feedback.
doi:10.1371/journal.pone.0055970
PMCID: PMC3569433  PMID: 23409099
11.  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
12.  Aging and movement errors when lifting and lowering light loads 
Age  2010;33(3):393-407.
The purpose was to determine the influence of movement variability and level of muscle activation on the accuracy of targeted movements performed with the index finger by young and older adults. Twelve young (27.4 ± 4.4 years) and 12 older adults (74.5 ± 8.9 years) attempted to match the end position of an index finger movement to a target position when lifting and lowering a light load (10% of the maximum). Visual feedback was provided after each trial. Movement error was calculated as the absolute distance from the target. Movement variability was quantified as the standard deviation of finger acceleration and the variability of end position across trials. The EMG activity of first dorsal interosseus (FDI) and second palmar interosseus (SPI) muscles was measured with intramuscular electrodes. Older adults exhibited greater spatial and temporal errors and greater variability in finger acceleration and end position during both the lifting and lowering tasks. Older adults lifted the load by activating FDI less but SPI the same as young adults, whereas they lowered the load by activating SPI less and FDI the same as young adults. In addition, older adults exhibited lower variability across trials in SPI activation when lifting the load and lower variability for FDI activation when lowering the load. The findings demonstrate that the decrease in spatial and temporal accuracy observed in older adults when lifting and lowering a light load to a target position was due to greater movement variability and differences in antagonistic muscle activity.
doi:10.1007/s11357-010-9190-4
PMCID: PMC3168598  PMID: 20945163
Motor output variability; Older adults; Movement control; EMG; Muscle synergy; Antagonist muscles
13.  Neural control of the lips differs for young and older adults following a perturbation 
Aging impairs the control of many skilled movements including speech. The purpose of this paper was to investigate whether young and older adults adapt to lower lip perturbations during speech differently. Twenty men (10 young, 26 ± 3 years of age; 10 older, 60 ± 9 years of age) were requested to repeat the word (“papa”) 300 times. In 15% of the trials, the subjects experienced a mechanical perturbation on the lower lip. Displacement and neural activation (EMG) of the upper and lower lips were evaluated. Perturbations to the lower lip caused a greater increase in the maximum displacement of the lower lip for older adults compared with young adults (34.7 ± 19% vs. 13.4 ± 17%; P = 0.017). Furthermore, young adults exhibited significantly greater 30–100 Hz normalized EMG power for the lower lip compared to the upper lip (P < 0.005). In young adults, changes from normal to perturbed trials in the 30–50 Hz frequency band of the EMG were negatively correlated to the changes from normal to perturbed trials in the lower lip maximum displacement (R2 = 0.48; P = 0.025). It is concluded that young adults adapt better to lower lip perturbations compared with older adults and that the associated neural activation strategy of the involved muscle is different for the two age groups.
doi:10.1007/s00221-010-2411-3
PMCID: PMC3167083  PMID: 20852991
EMG; Wavelet analysis; Speech; Cortical drive
14.  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

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