Ankle muscles drive mediolateral center of pressure control to ensure stable steady state gait.

During steady-state walking, mediolateral gait stability can be maintained by controlling the center of pressure (CoP). The CoP modulates the moment of the ground reaction force, which brakes and reverses movement of the center of mass (CoM) towards the lateral border of the base of support. In addition to foot placement, ankle moments serve to control the CoP. We hypothesized that, during steady-state walking, single stance ankle moments establish a CoP shift to correct for errors in foot placement. We expected ankle muscle activity to be associated with this complementary CoP shift. During treadmill walking, full-body kinematics, ground reaction forces and electromyography were recorded in thirty healthy participants. We found a negative relationship between preceding foot placement error and CoP displacement during single stance; steps that were too medial were compensated for by a lateral CoP shift and vice versa, steps that were too lateral were compensated for by a medial CoP shift. Peroneus longus, soleus and tibialis anterior activity correlated with these CoP shifts. As such, we identified an (active) ankle strategy during steady-state walking. As expected, absolute explained CoP variance by foot placement error decreased when walking with shoes constraining ankle moments. Yet, contrary to our expectations that ankle moment control would compensate for constrained foot placement, the absolute explained CoP variance by foot placement error did not increase when foot placement was constrained. We argue that this lack of compensation reflects the interdependent nature of ankle moment and foot placement control. We suggest that single stance ankle moments do not only compensate for preceding foot placement errors, but also assist control of the subsequent foot placement. Foot placement and ankle moment control are 'caught' in a circular relationship, in which constraints imposed on one will also influence the other.

Active foot placement control ensures stable gait: Effect of constraints on foot placement and ankle moments.

Step-by-step foot placement control, relative to the center of mass (CoM) kinematic state, is generally considered a dominant mechanism for maintenance of gait stability. By adequate (mediolateral) positioning of the center of pressure with respect to the CoM, the ground reaction force generates a moment that prevents falling. In healthy individuals, foot placement is complemented mainly by ankle moment control ensuring stability. To evaluate possible compensatory relationships between step-by-step foot placement and complementary ankle moments, we investigated the degree of (active) foot placement control during steady-state walking, and under either foot placement-, or ankle moment constraints. Thirty healthy participants walked on a treadmill, while full-body kinematics, ground reaction forces and EMG activities were recorded. As a replication of earlier findings, we first showed step-by-step foot placement is associated with preceding CoM state and hip ab-/adductor activity during steady-state walking. Tight control of foot placement appears to be important at normal walking speed because there was a limited change in the degree of foot placement control despite the presence of a foot placement constraint. At slow speed, the degree of foot placement control decreased substantially, suggesting that tight control of foot placement is less essential when walking slowly. Step-by-step foot placement control was not tightened to compensate for constrained ankle moments. Instead compensation was achieved through increases in step width and stride frequency.

The Pelvic Girdle Pain deadlock: 2. Topics that, so far, have remained out of focus.

INTRODUCTION: In our preceding paper, we concluded that Pelvic Girdle Pain (PGP) should be taken seriously. Still, we do not know its causes. Literature reviews on treatment fail to reveal a consistent pattern, and there are patients who do not respond well to treatment. We designated the lack of progress in research and in the clinic as 'deadlock', and proposed a 'deconstruction' of PGP, that is to say, taking PGP apart into its relevant dimensions. PURPOSE: We examine the proposition that PGP may emerge as local inflammation. Inflammation would be a new dimension to be taken into account, between biomechanics and psychology. To explore the consequences of this idea, we present four different topics that, so far, have remained out of focus. One: The importance of microtrauma. Two: Ways to counteract chronification. Three: The importance of sickness behaviour when systemic inflammation turns into neuroinflammation of the brain. And Four: The mainly emotional and cognitive nature of chronic pain, and how aberrant neuroinflammation may render chronic pain intractable. For intractable pain, sleep and stress management are promising treatment options. IMPLICATIONS: The authors hope that the present paper helps to stimulate the flexible creativity that is required to deal with the biological and psychological impact of PGP. Measuring inflammatory mediators in PGP should be a research priority. It should be understood that the boundaries between biology and psychology are becoming blurred. Clinicians must frequently monitor pain, disability, and mood, and be ready to switch treatment whenever the patient does not improve.

The addition of the MEP amplitude of finger extension muscles to clinical predictors of hand function after stroke: A prospective cohort study.

BACKGROUND: Within the first 72 hours after stroke, active finger extension is a strong predictor of long-term dexterity. Transcranial magnetic stimulation may add prognostic value to clinical assessment, which is especially relevant for patients unable to follow instructions. OBJECTIVE: The current prospective cohort study aims at determining whether amplitude of motor evoked potentials of the extensor digitorum communis (EDC) can improve clinical prediction after stroke when added to clinical tests. METHODS: the amplitude of motor evoked potentials of the affected EDC muscle at rest was measured in 18 participants within 4 weeks after stroke, as were the ability to perform finger extension and the Fugl-Meyer Motor Assessment of the upper extremity (FMA_UE). These three determinants were related to the FMA_UE at 26 weeks after stroke (FMA_UE26), both directly, and via the proportional recovery prediction model. The relation between amplitude of the motor evoked potentials and FMA_UE26 was evaluated for EDC. For comparison, also the MEP amplitudes of biceps brachii and adductor digiti minimi muscles were recorded. RESULTS: Patients' ability to voluntarily extend the fingers was strongly related to FMA_UE26, in our cohort there were no false negative results for this predictor. Our data revealed that the relation between amplitude of motor evoked potential of EDC and FMA_UE26 was significant, but moderate (rs = 0.58) without added clinical value. The other tested muscles did not correlate significantly to FMA_UE26. CONCLUSIONS: Our study demonstrates no additional value of motor evoked potential amplitude of the affected EDC muscle to the clinical test of finger extension, the latter being more strongly related to FMA_UE26.

How does upper extremity Fugl-Meyer motor score relate to resting-state EEG in chronic stroke? A power spectral density analysis.

OBJECTIVE: We investigated the potential added value of high-density resting-state EEG by addressing differences with healthy individuals and associations with Fugl-Meyer motor assessment of the upper extremity (FM-UE) scores in chronic stroke. METHODS: Twenty-one chronic stroke survivors with initial upper limb paresis and eleven matched controls were included. Group differences regarding resting-state EEG parameters (Delta Alpha ratio (DAR) and pairwise-derived Brain Symmetry Index (BSI)) and associations with FM-UE were investigated, as well as lateralization of BSI and the value of different frequency bands. RESULTS: Chronic stroke survivors showed higher BSI compared to controls (p < 0.001), most pronounced in delta and theta frequency bands (p < 0.0001; p < 0.001). In the delta and theta band, BSI was significantly negatively associated with FM-UE (both p = 0.008) corrected for confounding factors. DAR showed no differences between groups nor association with FM-UE. Directional BSI showed increased power in the affected versus the unaffected hemisphere. CONCLUSIONS: Asymmetry in spectral power between hemispheres was present in chronic stroke, most pronounced in low frequencies and related to upper extremity motor function deficit. SIGNIFICANCE: BSI is related to motor impairment and higher in chronic stroke patients compared to healthy controls, suggesting that BSI may be a marker of selective motor control.

Involvement of superior temporal areas in audiovisual and audiomotor speech integration.

Perception of speech sounds is affected by observing facial motion. Incongruence between speech sounds and watching somebody articulating may influence the perception of auditory syllable, referred to as the McGurk effect. We tested the degree to which silent articulation of a syllable also affects speech perception and searched for its neural correlates. Listeners were instructed to identify the auditory syllables /pa/ and /ta/ while silently articulating congruent/incongruent syllables or observing videos of a speaker's face articulating them. As a baseline, we included an auditory-only condition without competing visual or sensorimotor input. As expected, perception of sounds degraded when incongruent syllables were observed, and also when they were silently articulated, albeit to a lesser extent. This degrading was accompanied by significant amplitude modulations in the beta frequency band in right superior temporal areas. In these areas, the event-related beta activity during congruent conditions was phase-locked to responses evoked during the auditory-only condition. We conclude that proper temporal alignment of different input streams in right superior temporal areas is mandatory for both audiovisual and audiomotor speech integration.

Coordinative task difficulty and behavioural errors are associated with increased long-range beta band synchronization.

The neural network and the task-dependence of (local) activity changes involved in bimanual coordination are well documented. However, much less is known about the functional connectivity within this neural network and its modulation according to manipulations of task complexity. Here, we assessed neural activity via high-density electroencephalography, focussing on changes of activity in the beta frequency band (~15-30Hz) across the motor network in 26 young adult participants (19-29 years old). We investigated how network connectivity was modulated with task difficulty and errors of performance during a bimanual visuomotor movement consisting of dial rotation according to three different ratios of speed: an isofrequency movement (1:1), a non-isofrequency movement with the right hand keeping the fast pace (1:3), and the converse ratio with the left hand keeping the fast pace (3:1). To quantify functional coupling, we determined neural synchronization which might be key for the timing of the activity within brain regions during task execution. Individual source activity with realistic head models was reconstructed at seven regions of interest including frontal and parietal areas, among which we estimated phase-based connectivity. Partial least squares analysis revealed a significant modulation of connectivity with task difficulty, and significant correlations between connectivity and errors in performance, in particular between sensorimotor cortices. Our findings suggest that modulation of long-range synchronization is instrumental for coping with increasing task demands in bimanual coordination.

Effects of handrail hold and light touch on energetics, step parameters, and neuromuscular activity during walking after stroke.

BACKGROUND: Holding a handrail or using a cane may decrease the energy cost of walking in stroke survivors. However, the factors underlying this decrease have not yet been previously identified. The purpose of the current study was to fill this void by investigating the effect of physical support (through handrail hold) and/or somatosensory input (through light touch contact with a handrail) on energy cost and accompanying changes in both step parameters and neuromuscular activity. Elucidating these aspects may provide useful insights into gait recovery post stroke. METHODS: Fifteen stroke survivors participated in this study. Participants walked on a treadmill under three conditions: no handrail contact, light touch of the handrail, and firm handrail hold. During the trials we recorded oxygen consumption, center of pressure profiles, and bilateral activation of eight lower limb muscles. Effects of the three conditions on energy cost, step parameters and neuromuscular activation were compared statistically using conventional ANOVAs with repeated measures. In order to examine to which extent energy cost and step parameters/muscle activity are associated, we further employed a partial least squares regression analysis. RESULTS: Handrail hold resulted in a significant reduction in energy cost, whereas light touch contact did not. With handrail hold subjects took longer steps with smaller step width and improved step length symmetry, whereas light touch contact only resulted in a small but significant decrease in step width. The EMG analysis indicated a global drop in muscle activity, accompanied by an increased constancy in the timing of this activity, and a decreased co-activation with handrail hold, but not with light touch. The regression analysis revealed that increased stride time and length, improved step length symmetry, and decreased muscle activity were closely associated with the decreased energy cost during handrail hold. CONCLUSION: Handrail hold, but not light touch, altered step parameters and was accompanied by a global reduction in muscle activity, with improved timing constancy. This suggests that the use of a handrail allows for a more economic step pattern that requires less muscular activation without resulting in substantial neuromuscular re-organization. Handrail use may thus have beneficial effects on gait economy after stroke, which cannot be accomplished through enhanced somatosensory input alone.

Understanding bimanual coordination across small time scales from an electrophysiological perspective.

Bimanual movement involves a variety of coordinated functions, ranging from elementary patterns that are performed automatically to complex patterns that require practice to be performed skillfully. The neural dynamics accompanying these coordination patterns are complex and rapid. By means of electro- and magneto-encephalographic approaches, it has been possible to examine these dynamics during bimanual coordination with excellent temporal resolution, which complements other neuroimaging modalities with superb spatial resolution. This review focuses on EEG/MEG studies that unravel the processes involved in movement planning and execution, motor learning, and executive functions involved in task switching and dual tasking. Evidence is presented for a spatio-temporal reorganization of the neural networks within and between hemispheres to meet increased task difficulty demands, induced or spontaneous switches in coordination mode, or training-induced neuroplastic modulation in coordination dynamics. Future theoretical developments will benefit from the integration of research techniques unraveling neural activity at different time scales. Ultimately this work will contribute to a better understanding of how the human brain orchestrates complex behavior via the implementation of inter- and intra-hemispheric coordination networks.

A benchmark test of accuracy and precision in estimating dynamical systems characteristics from a time series.

Characteristics of dynamical systems are often estimated to describe physiological processes. For instance, Lyapunov exponents have been determined to assess the stability of the cardio-vascular system, respiration, and, more recently, human gait and posture. However, the systematic evaluation of the accuracy and precision of these estimates is problematic because the proper values of the characteristics are typically unknown. We fill this void with a set of standardized time series with well-defined dynamical characteristics that serve as a benchmark. Estimates ought to match these characteristics, at least to good approximation. We outline a procedure to employ this generic benchmark test and illustrate its capacity by examining methods for estimating the maximum Lyapunov exponent. In particular, we discuss algorithms by Wolf and co-workers and by Rosenstein and co-workers and evaluate their performances as a function of signal length and signal-to-noise ratio. In all scenarios, the precision of Rosenstein's algorithm was found to be equal to or greater than Wolf's algorithm. The latter, however, appeared more accurate if reasonably large signal lengths are available and noise levels are sufficiently low. Due to its modularity, the presented benchmark test can be used to evaluate and tune any estimation method to perform optimally for arbitrary experimental data.

Nonlinear coupling between occipital and motor cortex during motor imagery: a dynamic causal modeling study.

We demonstrate the capacity of dynamic causal modeling to characterize the nonlinear coupling among cortical sources that underlie time-frequency modulations in MEG data. Our experimental task involved the mental rotation of hand drawings that ten subjects used to decide if it was a right or left hand. Reaction times were shorter when the stimuli were presented with a small rotation angle (fast responses) compared to a large rotation angle (slow responses). The grand-averaged data showed that in both cases performance was accompanied by a marked increase in gamma activity in occipital areas and a concomitant decrease in alpha and beta power in occipital and motor regions. Modeling directed (cross) frequency interactions between the two regions revealed that after the stimulus induced a gamma increase and beta decrease in occipital regions, interactions with the motor area served to attenuate these modulations. The difference between fast and slow behavioral responses was manifest as an altered coupling strength in both forward and backward connections, which led to a less pronounced attenuation for more difficult (slow reaction time) trials. This was mediated by a (backwards) beta to gamma coupling from motor till occipital sources, whereas other interactions were mainly within the same frequency. Results are consistent with the theory of predictive coding and suggest that during motor imagery, the influence of motor areas on activity in occipital cortex co-determines performance. Our study illustrates the benefit of modeling experimental responses in terms of a generative model that can disentangle the contributions of intra-areal vis-à-vis inter-areal connections to time-frequency modulations during task performance.

Differential modulations of ipsilateral and contralateral beta (de)synchronization during unimanual force production.

Unilateral movement is usually accompanied by ipsilateral activity in the primary motor cortex (M1). It is still largely unclear whether this activity reflects interhemispheric 'cross-talk' of contralateral M1 that facilitates movement, or results from processes that inhibit motor output. We investigated the role of beta power in ipsilateral M1 during unimanual force production. Significant ipsilateral beta desynchronization occurred during continuous dynamic but not during static force production. Moreover, event-related time-frequency analysis revealed bilateral desynchronization patterns, whereas post-movement synchronization was confined to the contralateral hemisphere. Our findings indicate that ipsilateral activation is not merely the result of interhemispheric cross-talk but involves additional processes. Given observations of differential blood oxygen level-dependent responses in ipsilateral and contralateral M1, and the correlation between beta desynchronization and the firing rate of pyramidal tract neurons in contralateral M1 during movement, we speculate that beta desynchronization in contra- and ipsilateral M1 arises from distinct neural activation patterns.

The relationship between structural and functional connectivity: graph theoretical analysis of an EEG neural mass model.

We investigated the relationship between structural network properties and both synchronization strength and functional characteristics in a combined neural mass and graph theoretical model of the electroencephalogram (EEG). Thirty-two neural mass models (NMMs), each representing the lump activity of reasonably large groups of interacting excitatory and inhibitory neurons, were reciprocally and excitatory coupled using random rewiring as described by Watts and Strogatz. Numerical analysis of the network revealed an abrupt transition towards a synchronized state as a function of increasing coupling strength alpha. Synchronization increased with increasing degree and decreasing regularity of the network. Parameters of the functional network showed a diverse dependency on structural connectivity: normalized clustering coefficient gamma and path length lambda increased with increasing alpha. For sufficiently large alpha, however, gamma decreased with increasing rewiring probability p, while lambda increased. Hence, a structured functional network exists despite the randomness of the underlying structural network. That is, patterns of functional connectivity are influenced by patterns of the corresponding structural level but do not necessarily agree with those.

Cortico-spinal synchronization reflects changes in performance when learning a complex bimanual task.

Motor performance is accompanied by neural activity in various cortical and sub-cortical areas. This intricate network has to be delicately orchestrated. We analyzed the role of beta synchronization in motor learning using magneto-encephalography combined with electromyography. Cortico-spinal synchronization in the beta band was found to be of particular importance in establishing bimanual movement patterns in the context of a 3:2 polyrhythmic (isometric) force production task. Its dynamics correlated highly with the learning of this complex bimanual motor skill. We submit that the cortical dynamics entrains the spinal motor system by which cortico-spinal beta synchrony serves higher-level motor control functions as primary means of information transfer along the neural axis.

Asymmetry in pulse-coupled oscillators with delay.

We studied the dynamics of synchronization in asymmetrically coupled neural oscillators with time delay. Stability analysis revealed that symmetric excitatory coupling results in synchrony at multiple phase relations. Asymmetry yields two saddle-node bifurcations of the stable states when coupling is asymmetric. By contrast, with inhibitory coupling only in phase or antiphase is stable as long as coupling is symmetric. Otherwise, these stable states shift or even vanish. The reduced bistability range suggests the beneficial role of asymmetric coupling for reliable neural information transfer.

A role of beta oscillatory synchrony in biasing response competition?

Beta-range oscillatory activity measured over the motor cortex and beta synchrony between cortex and spinal cord can be up- or downregulated in anticipation of a postural challenge or the initiation of movement. Based on these properties of beta activity in the preparation for future events, the present investigation addressed whether simultaneous up- and downregulation of beta activity might act as an online mechanism to suppress and select competing responses. Measures of local and long-range beta synchrony were obtained from electroencephalographic and electromyographic signals recorded during a cued choice reaction task. Analyses focused on task-related changes in beta synchrony during a 2-s delay period between cue and response signal. Analyzed separately, none of the beta measures (spectral power, corticospinal coherence, corticospinal phase synchronization) showed simultaneous up- and downregulation over opposite hemispheres controlling the competing responses. However, the combined pattern of beta measures showed beta power desynchronization associated with selection of a response and increased corticospinal coherence and phase synchronization associated with suppression of a response. These results indicate that concurrent up- and downregulation of different components of beta oscillatory activity is likely to have a functional role in response selection, resembling attentional modulation of alpha activity in visual selection.

Heterogeneity of muscle activation in relation to force direction: a multi-channel surface electromyography study on the triceps surae muscle.

Several skeletal muscles can be divided into sub-modules, called neuromuscular compartments (NMCs), which are thought to be controlled independently and to have distinct biomechanical functions. We looked for distinct muscle activation patterns in the triceps surae muscle (TS) using surface electromyography (EMG) during voluntary contraction. Nine subjects performed isometric and isotonic plantar flexions combined with forces along pre-defined directions. Besides the forces under the ball of the foot, multi-channel surface EMG was measured with electrodes homogeneously distributed over the entire TS. Using principal component analysis, common (global) components were omitted from the EMG signals, thereby estimating muscle activity sufficiently accurate to track fine fluctuations of force during an isotonic contraction (r=0.80+/-0.09). A subsequent cluster analysis showed a topographical organization of co-activated parts of the muscle that was different between subjects. Low and negative correlations between the EMG activity within clusters were found, indicating a substantial heterogeneity of TS activation. The correlations between cluster time series and forces at the foot in specific directions differed substantially between clusters, showing that the differentially activated parts of the TS had specific biomechanical functions.

Graph theoretical analysis of magnetoencephalographic functional connectivity in Alzheimer's disease.

In this study we examined changes in the large-scale structure of resting-state brain networks in patients with Alzheimer's disease compared with non-demented controls, using concepts from graph theory. Magneto-encephalograms (MEG) were recorded in 18 Alzheimer's disease patients and 18 non-demented control subjects in a no-task, eyes-closed condition. For the main frequency bands, synchronization between all pairs of MEG channels was assessed using a phase lag index (PLI, a synchronization measure insensitive to volume conduction). PLI-weighted connectivity networks were calculated, and characterized by a mean clustering coefficient and path length. Alzheimer's disease patients showed a decrease of mean PLI in the lower alpha and beta band. In the lower alpha band, the clustering coefficient and path length were both decreased in Alzheimer's disease patients. Network changes in the lower alpha band were better explained by a 'Targeted Attack' model than by a 'Random Failure' model. Thus, Alzheimer's disease patients display a loss of resting-state functional connectivity in lower alpha and beta bands even when a measure insensitive to volume conduction effects is used. Moreover, the large-scale structure of lower alpha band functional networks in Alzheimer's disease is more random. The modelling results suggest that highly connected neural network 'hubs' may be especially at risk in Alzheimer's disease.

Steady and transient coordination structures of walking and running.

We studied multisegmental coordination and stride characteristics in nine participants while walking and running on a treadmill. The study's main aim was to evaluate the coordination patterns of walking and running and their variance as a function of locomotion speed, with a specific focus on gait transitions and accompanying features like hysteresis and critical fluctuations. Stride characteristics changed systematically with speed in a gait-dependent fashion, but exhibited no hysteresis. Multisegmental coordination of walking and running was captured by four principal components, the first two of which were present in both gaits. Locomotion speed had subtle yet systematic differential effects on the relative phasing between the identified components in both walking and running and its variance, in particular in the immediate vicinity of gait transitions. Unlike the stride characteristics, the identified coordination patterns revealed clear evidence of both hysteresis and critical fluctuations around transition points. Overall, the results suggest that walking and running entail similar, albeit speed- and gait-dependent, coordination structures, and that gait transitions bear signatures of nonequilibrium phase transitions. Application of multivariate analyses of whole-body recordings appears crucial to detect these features in a reliable fashion.