Increased gravitational force reveals the mechanical, resonant nature of physiological tremor

Human physiological hand tremor has a resonant component. Proof of this is that its frequency can be modified by adding mass. However, adding mass also increases the load which must be supported. The necessary force requires muscular contraction which will change motor output and is likely to increase limb stiffness. The increased stiffness will partly offset the effect of the increased mass and this can lead to the erroneous conclusion that factors other than resonance are involved in determining tremor frequency. Using a human centrifuge to increase head-to-foot gravitational field strength, we were able to control for the increased effort by increasing force without changing mass. This revealed that the peak frequency of human hand tremor is 99% predictable on the basis of a resonant mechanism. We ask what, if anything, the peak frequency of physiological tremor can reveal about the operation of the nervous system.This work was funded by a BBSRC Industry Interchange Award to J.P.R.S. and R.F.R. C.J.O. was funded by BBSRC grant BB/I00579X/1. C.A.V. was funded by A∗Midex (Aix-Marseille Initiative of Excellence

Intrinsic ankle stiffness is associated with paradoxical calf muscle movement but not postural sway or age

Due to Achilles tendon compliance, passive ankle stiffness is insufficient to stabilise the body when standing. This results in ‘paradoxical’ muscle movement, whereby calf muscles tend to shorten during forward body sway. Natural variation in stiffness may affect this movement. This may have consequences for postural control, with compliant ankles placing greater reliance upon active neural control rather than stretch reflexes. Previous research also suggests ageing reduces ankle stiffness, possibly contributing to reduced postural stability. Here we determine the relationship between ankle stiffness and calf muscle movement during standing, and whether this is associated with postural stability or age. Passive ankle stiffness was measured during quiet stance in 40 healthy volunteers ranging from 18 to 88 years of age. Medial gastrocnemius muscle length was also recorded using ultrasound. We found a significant inverse relationship between ankle stiffness and paradoxical muscle movement, that is, more compliant ankles were associated with greater muscle shortening during forward sway (r ≥ 0.33). This was seen during both quiet stance as well as voluntary sway. However, we found no significant effects of age upon stiffness, paradoxical motion or postural sway. Furthermore, neither paradoxical muscle motion nor ankle stiffness was associated with postural sway. These results show that natural variation in ankle stiffness alters the extent of paradoxical calf muscle movement during stance. However, the absence of a clear relationship to postural sway suggests that neural control mechanisms are more than capable of compensating for a lack of inherent joint stiffness

Visuo-manual tracking: does intermittent control with aperiodic sampling explain linear power and non-linear remnant without sensorimotor noise?

© 2017 The Authors. The Journal of Physiology published by John Wiley & Sons Ltd on behalf of The Physiological Society Key points: A human controlling an external system is described most easily and conventionally as linearly and continuously translating sensory input to motor output, with the inevitable output remnant, non-linearly related to the input, attributed to sensorimotor noise. Recent experiments show sustained manual tracking involves repeated refractoriness (insensitivity to sensory information for a certain duration), with the temporary 200–500 ms periods of irresponsiveness to sensory input making the control process intrinsically non-linear. This evidence calls for re-examination of the extent to which random sensorimotor noise is required to explain the non-linear remnant. This investigation of manual tracking shows how the full motor output (linear component and remnant) can be explained mechanistically by aperiodic sampling triggered by prediction error thresholds. Whereas broadband physiological noise is general to all processes, aperiodic sampling is associated with sensorimotor decision making within specific frontal, striatal and parietal networks; we conclude that manual tracking utilises such slow serial decision making pathways up to several times per second. Abstract: The human operator is described adequately by linear translation of sensory input to motor output. Motor output also always includes a non-linear remnant resulting from random sensorimotor noise from multiple sources, and non-linear input transformations, for example thresholds or refractory periods. Recent evidence showed that manual tracking incurs substantial, serial, refractoriness (insensitivity to sensory information of 350 and 550 ms for 1st and 2nd order systems respectively). Our two questions are: (i) What are the comparative merits of explaining the non-linear remnant using noise or non-linear transformations? (ii) Can non-linear transformations represent serial motor decision making within the sensorimotor feedback loop intrinsic to tracking? Twelve participants (instructed to act in three prescribed ways) manually controlled two systems (1st and 2nd order) subject to a periodic multi-sine disturbance. Joystick power was analysed using three models, continuous-linear-control (CC), continuous-linear-control with calculated noise spectrum (CCN), and intermittent control with aperiodic sampling triggered by prediction error thresholds (IC). Unlike the linear mechanism, the intermittent control mechanism explained the majority of total power (linear and remnant) (77–87% vs. 8–48%, IC vs. CC). Between conditions, IC used thresholds and distributions of open loop intervals consistent with, respectively, instructions and previous measured, model independent values; whereas CCN required changes in noise spectrum deviating from broadband, signal dependent noise. We conclude that manual tracking uses open loop predictive control with aperiodic sampling. Because aperiodic sampling is inherent to serial decision making within previously identified, specific frontal, striatal and parietal networks we suggest that these structures are intimately involved in visuo-manual tracking

Predictive feedback control and Fitts' law

Fitts’ law is a well established empirical formula, known for encapsulating the “speed-accuracy trade-off”. For discrete, manual movements from a starting location to a target, Fitts’ law relates movement duration to the distance moved and target size. The widespread empirical success of the formula is suggestive of underlying principles of human movement control. There have been previous attempts to relate Fitts’ law to engineering-type control hypotheses and it has been shown that the law is exactly consistent with the closed-loop step-response of a time-delayed, first-order system. Assuming only the operation of closed-loop feedback, either continuous or intermittent, this paper asks whether such feedback should be predictive or not predictive to be consistent with Fitts law. Since Fitts’ law is equivalent to a time delay separated from a first-order system, known control theory implies that the controller must be predictive. A predictive controller moves the time-delay outside the feedback loop such that the closed-loop response can be separated into a time delay and rational function whereas a non- predictive controller retains a state delay within feedback loop which is not consistent with Fitts’ law. Using sufficient parameters, a high-order non-predictive controller could approximately reproduce Fitts’ law. However, such high-order, “non-parametric” controllers are essentially empirical in nature, without physical meaning, and therefore are conceptually inferior to the predictive controller. It is a new insight that using closed-loop feedback, prediction is required to physically explain Fitts’ law. The implication is that prediction is an inherent part of the “speed-accuracy trade-off”

Intermittent control models of human standing: similarities and differences

Two architectures of intermittent control are compared and contrasted in the context of the single inverted pendulum model often used for describing standing in humans. The architectures are similar insofar as they use periods of open-loop control punctuated by switching events when crossing a switching surface to keep the system state trajectories close to trajectories leading to equilibrium. The architectures differ in two significant ways. Firstly, in one case, the open-loop control trajectory is generated by a system-matched hold, and in the other case, the open-loop control signal is zero. Secondly, prediction is used in one case but not the other. The former difference is examined in this paper. The zero control alternative leads to periodic oscillations associated with limit cycles; whereas the system-matched control alternative gives trajectories (including homoclinic orbits) which contain the equilibrium point and do not have oscillatory behaviour. Despite this difference in behaviour, it is further shown that behaviour can appear similar when either the system is perturbed by additive noise or the system-matched trajectory generation is perturbed. The purpose of the research is to come to a common approach for understanding the theoretical properties of the two alternatives with the twin aims of choosing which provides the best explanation of current experimental data (which may not, by itself, distinguish beween the two alternatives) and suggesting future experiments to distinguish between the two alternatives

Adaptation to altered balance conditions in unilateral amputees due to atherosclerosis: a randomized controlled study

<p>Abstract</p> <p>Background</p> <p>Amputation impairs the ability to balance. We examined adaptation strategies in balance following dysvascularity-induced unilateral tibial amputation in skilled prosthetic users (SPU) and first fitted amputees (FFA) (N = 28).</p> <p>Methods</p> <p>Excursions of center of pressure (COP) were determined during 20 s quiet standing using a stabilometry system with eyes-open on both legs or on the non-affected leg(s). Main measures: COP trajectories and time functions; distribution of reaction forces between the two legs; inclination angles obtained through second order regression analysis using stabilogram data.</p> <p>Results</p> <p>FFA vs SPU demonstrated 27.8% greater postural sway in bilateral stance (p = 0.0004). Postural sway area was smaller in FFA standing on the non-affected leg compared with SPU (p = 0.028). The slope of the regression line indicating postural stability was nearly identical in FFA and SPU and the direction of regression line was opposite for the left and right leg amputees.</p> <p>Conclusion</p> <p>Of the two adaptation strategies in balance, the first appears before amputation due to pain and fatigue in the affected leg. This strategy appears in the form of reduced postural sway while standing on the non-affected leg. The second adaptation occurs during rehabilitation and regular use of the prosthesis resulting in normal weightbearing associated with reduced postural sway on two legs and return to the normal postural stability on one leg.</p

Synchrony of hand-foot coupled movements: is it attained by mutual feedback entrainment or by independent linkage of each limb to a common rhythm generator?

BACKGROUND: Synchrony of coupled oscillations of ipsilateral hand and foot may be achieved by controlling the interlimb phase difference through a crossed kinaesthetic feedback between the two limbs, or by an independent linkage of each limb cycle to a common clock signal. These alternative models may be experimentally challenged by comparing the behaviour of the two limbs when they oscillate following an external time giver, either alone or coupled together. RESULTS: Ten subjects oscillated their right hand and foot both alone and coupled (iso- or antidirectionally), paced by a metronome. Wrist and ankle angular position and Electromyograms (EMG) from the respective flexor and extensor muscles were recorded. Three phase delays were measured: i) the clk-mov delay, between the clock (metronome beat) and the oscillation peak; ii) the neur (neural) delay, between the clock and the motoneurone excitatory input, as inferred from the EMG onset; and iii) the mech (mechanical) delay between the EMG onset and the corresponding point of the limb oscillation. During uncoupled oscillations (0.4 Hz to 3.0 Hz), the mech delay increased from -7° to -111° (hand) and from -4° to -83° (foot). In contrast, the clk-mov delay remained constant and close to zero in either limb since a progressive advance of the motoneurone activation on the pacing beat (neur advance) compensated for the increasing mech delay. Adding an inertial load to either extremity induced a frequency dependent increase of the limb mechanical delay that could not be completely compensated by the increase of the neural phase advance, resulting in a frequency dependent increment of clk-mov delay of the hampered limb. When limb oscillations were iso- or antidirectionally coupled, either in the loaded or unloaded condition, the three delays did not significantly change with respect to values measured when limbs were moved separately. CONCLUSION: The absence of any significant effect of limb coupling on the measured delays suggests that during hand-foot oscillations, both iso- and antidirectionally coupled, each limb is synchronised to the common rhythm generator by a "private" position control, with no need for a crossed feedback interaction between limbs

There is no simple temporal relationship between the initiation of rapid reactive hand movements and the phase of an enhanced physiological tremor in man

A postural hand tremor of enhanced size was induced in eleven subjects. There was rhythmic activity in the posturally active extensor muscles synchronised to the tremor oscillation, which implied an oscillatory modulation of motor neurones.The subjects executed single rapid wrist flexion movements in response to a flash of light. The light flash was presented at an instant when the wrist was spontaneously moving in the flexion direction, extension direction or at random. The time taken to generate a movement was not significantly different or more consistent in any of the conditions.Inspection of individual and averaged acceleration and EMG records strongly suggests that the movement is made at a time that is independent of the tremor cycle at the time of stimulus presentation or at the moment of movement initiation.These observations suggest that the central or spinal mechanism generating tremor does not gate the central mechanism that produces voluntary movement