A MECHANISTIC APPROACH TO POSTURAL DEVELOPMENT IN CHILDREN

Upright standing is intrinsically unstable and requires active control. The central nervous system's feedback process is the active control that integrates multi-sensory information to generate appropriate motor commands to control the plant (the body with its musculotendon actuators). Maintaining standing balance is not trivial for a developing child because the feedback and the plant are both developing and the sensory inputs used for feedback are continually changing. Knowledge gaps exist in characterizing the critical ability of adaptive multi-sensory reweighting for standing balance control in children. Furthermore, the separate contributions of the plant and feedback and their relationship are poorly understood in children, especially when considering that the body is multi-jointed and feedback is multi-sensory. The purposes of this dissertation are to use a mechanistic approach to study multi-sensory abilities of typically developing (TD) children and children with Developmental Coordination Disorder (DCD). The specific aims are: 1) to characterize postural control under different multi-sensory conditions in TD children and children with DCD; 2) to characterize the development of adaptive multi-sensory reweighting in TD children and children with DCD; and, 3) to identify the plant and feedback for postural control in TD children and how they change in response to visual reweighting. In the first experiment (Aim 1), TD children, adults, and 7-year-old children with DCD are tested under four sensory conditions (no touch/no vision, with touch/no vision, no touch/with vision, and with touch/with vision). We found that touch robustly attenuated standing sway in all age groups. Children with DCD used touch less effectively than their TD peers and they also benefited from using vision to reduce sway. In the second experiment (Aim 2), TD children (4- to 10-year-old) and children with DCD (6- to 11-year-old) were presented with simultaneous small-amplitude touch bar and visual scene movement at 0.28 and 0.2 Hz, respectively, within five conditions that independently varied the amplitude of the stimuli. We found that TD children can reweight to both touch and vision from 4 years on and the amount of reweighting increased with age. However, multi-sensory fusion (i.e., inter-modal reweighting) was only observed in the older children. Children with DCD reweight to both touch and vision at a later age (10.8 years) than their TD peers. Even older children with DCD do not show advanced multisensory fusion. Two signature deficits of multisensory reweighting are a weak vision reweighting and a general phase lag to both sensory modalities. The final aim involves closed-loop system identification of the plant and feedback using electromyography (EMG) and kinematic responses to a high- or low-amplitude visual perturbation and two mechanical perturbations in children ages six and ten years and adults. We found that the plant is different between children and adults. Children demonstrate a smaller phase difference between trunk and leg than adults at higher frequencies. Feedback in children is qualitatively similar to adults. Quantitatively, children show less phase advance at the peak of the feedback curve which may be due to a longer time delay. Under the high and low visual amplitude conditions, children show less gain change (interpreted as reweighting) than adults in the kinematic and EMG responses. The observed kinematic and EMG reweighting are mainly due to the different use of visual information by the central nervous system as measured by the open-loop mapping from visual scene angle to EMG activity. The plant and the feedback do not contribute to reweighting

Phase Dynamics in Human Visuomotor Control - Health & Disease

In this thesis, comprised of four publications, I investigated phase dynamics of visuomotor control in humans during upright stance in response to an oscillatory visual drive. For this purpose, I applied different versions of a ‘moving room’ paradigm in virtual reality while stimulating human participants with anterior-posterior motion of their visual surround and analyzed their bodily responses. Human balance control constitutes a complex interplay of interdependent processes. The main sensory contributors include vision, vestibular input, and proprioception, with a dominant role attributed to vision. The purpose of the balance control system is to keep the body’s center of mass (COM) within a certain spatial range around the current base of support. Ever-changing environmental circumstances along with sensory noise cause the body to permanently sway around its point of equilibrium. Considering this sway, the human body can be modelled as a (multi-link) inverted pendulum. To maintain balance while being exposed to perturbations of the visual environment, humans adjust their sway to counteract the perceived motion of their bodies. Neurodegenerative diseases like Parkinson’s impair balance control and thus are likely to affect these mechanisms. Hence, investigation of bodily responses to a visual drive gives insight into visuomotor control in health and disease. In my first study, I introduced inter-trial phase coherence (ITPC) as a novel method to investigate postural responses to periodical visual stimulation. I found that human participants phase-locked the motion of their center of pressure (COP) to a 3-D dot cloud which oscillated in the anterior-posterior direction. This effect was equally strong for a low frequency of visual stimulation at 0.2 Hz and a high frequency of 1.5 Hz, the latter exceeding the previously assumed frequency range associated with coherent postural sway responses to periodical oscillations of the visual environment (moving room). Moreover, I was able to show that ITPC reliably captured responses in almost all participants, thereby addressing the common problem of inter-subject variability in body sway research. Based on the results of my first study, I concluded phase locking to be an essential feature in human postural control. For the second study, I introduced a mobile and cost-effective setup to apply a visual paradigm consisting of a virtual tunnel which stretched in the anterior-posterior direction and oscillated back and forth at three distinct frequencies (0.2 Hz, 0.8 Hz, and 1.2 Hz). Because tracking of the COP alone neglects crucial information about how COM shifts are arranged across the body, I included additional full-body motion tracking here to evaluate sway of individual body segments. Using a modified measure of phase locking, the phase locking value (PLV), allowed me to find participants phase-locking not only their COP, but also additional segments of their body to the visual drive. While their COP exhibited a strong phase locking to all frequencies of visual stimulation, distribution of phase locking across the body underwent a shift as the frequency of the visual stimulation increased. For the lowest frequency of 0.2 Hz, participants phase-locked almost their entire body to the stimulus. At higher frequencies, this phase locking shifted towards the lower torso and hip, with subjects almost exclusively phase-locking their hip to the visual drive at the highest frequency of 1.2 Hz. Having introduced a novel and reliable measurement along with a mobile setup, these results allowed me to empirically confirm shifts in postural strategies previously proposed in the literature. In the third study, a collaboration with the neurology department of the Universitätsklinikum Gießen und Marburg (UKGM), I used the same setup and paradigm as in the previous study and additionally derived the trajectory of the COM from a weighted combination of certain body segments. The aim was to investigate phase locking of body sway in a group of patients suffering from Parkinson’s disease (PD) to find potential means for an early diagnosis of the illness. For this purpose, I recruited a group of PD patients, an age-matched control group, and a group of young healthy adults. Even though the sway amplitude of PD patients was significantly larger than that of both other groups, they phase-locked their COP and COM in a similar manner as the control groups. However, considering individual body segments, the shift in PLV distribution differed between groups. While young healthy adults, analogous to the participants in the second study, exhibited a shift towards exclusive phase locking of their hips as frequency of the stimulation increased, both PD patients and age-matched controls maintained a rather homogeneous phase locking across their body. This suggested increased body stiffness, although being an effect of age rather than disease. Overall, I concluded that patients of early-to-mid stage PD exhibit impaired motor control, reflected in their increased sway amplitude, but intact visuomotor processing, indicated by their ability to phase-lock the motion of their body to a visual drive. The fourth study, to which I contributed as second author, used experimental data collected from an additional visual condition in the course of the third study. This condition consisted of unpredictable back and forward motion of the simulated tunnel. Here, we investigated the velocity profiles of the COP and COM in response to the unpredictable visual motion and a baseline condition at which the tunnel remained static. We found PD patients to exhibit larger velocities of their COP and COM under both conditions when compared to the control groups. When examining the net increase that unpredictable motion had on the velocity of both parameters, we found a significantly higher increase in COP velocity for both PD patients and age-matched controls, but no increase in COM velocity in any of the groups. These results suggested that all groups successfully maintained their balance under unpredictable visual perturbations, but that PD patients and older adults required more effort to accomplish this task, as reflected by the increased velocity of their COP. Again, these results indicated an effect of age rather than disease on the observed postural responses. In summary, using innovative phase-locking techniques and simultaneously tracking multiple body sway parameters, I was able to provide novel insight into visuomotor control in humans. First, I overcame previous issues of inconsistent sway parameters in groups of participants; Second, I found phase-locking to be an essential feature of visuomotor processing, which also allowed me to empirically confirm previously established theories of postural control; Third, through studies in collaboration with the neurology department of the UKGM, I was able to uncover new aspects of visuomotor processing in Parkinson’s, contributing to a better understanding of the sensorimotor aspects of the disease

Development of a theoretical model for upright postural control in lower limb prosthesis users

Methods used to assess quiet standing in unilateral prosthesis users often assume validity of an inverted pendulum model despite this being shown as invalid in some instances. The aim of the current study was to evaluate the validity of a proposed unilaterally-constrained pin-controller model in explaining postural control in unilateral prosthesis users. Prosthesis users were contrasted against the theoretical model as were able-bodied controls that stood on a platform which unilaterally constrained movement of the CoP. All participants completed bouts of quiet standing with eyes open, eyes closed and with feedback on inter-limb weight bearing asymmetry. Correlation coefficients were used to infer inverted pendulum behavior in both the anteroposterior and mediolateral directions and were derived from both kinematic (body attached markers) and kinetic (centre of pressure) experimental data. Larger, negative correlation coefficients reflected better model adherence, whilst low or no correlation reflected poorer model adherence. Inverted pendulum behavior derived from kinematic data, indicated coefficients of high magnitude in both mediolateral (all cases range 0.71–0.78) and anteroposterior (0.88–0.91) directions, irrespective of groups. Inverted pendulum behavior derived from kinetic data in the anteroposterior direction indicated validity of the model with large negative coefficients associated with the unconstrained/intact limbs (prosthesis users: − 0.45 to − 0.65, control group: − 0.43 to − 0.72), small coefficients in constrained/prosthetic limbs (prosthesis users: − 0.02 to 0.07, control group: 0.13–0.26) and large negative coefficients in combined conditions (prosthesis users: − 0.36 to − 0.56, control group: − 0.71 to − 0.82). For the mediolateral direction, coefficients were negligible for individual limbs (0.03–0.17) and moderate to large negative correlations, irrespective of group (− 0.31 to − 0.73). Data suggested both prosthesis users’ and able-bodied individuals’ postural control conforms well to that predicted by a unilaterally-constrained pin-controller model, which has implications for the fundamental control of posture in transtibial prosthesis users

Sensory-Related Changes in Two-Segment Dynamics on a Sway-Referenced Support Surface

In its simplest form, the human postural control system can be described as a closed-loop control system consisting of a plant (body segments and musculotendon actuators) and feedback. Previous efforts to understand the contributions of plant and feedback employed techniques to "open the loop" which is problematic with the study of posture because the plant is unstable without feedback. In the present experiment, a closed-loop system identification method was used to "open the loop" without removal of sensory feedback. Subjects stood on a movable platform facing a visual scene, both of which were capable of rotation about an axis coaxial with the subject's ankles. The visual stimulus (present all trials) consisted of a 10-frequency sum-of-sines while movement of the support surface consisted of the following conditions: 1. Stationary; 2. Sway-referenced to the subject's body sway; 3. 10-frequency sum-of-sines; 4. Combined sway-referenced and sum-of-sines. Closed-loop frequency response functions were calculated for visual stimulus to EMG and visual stimulus to body sway angle. The open loop frequency response function for the plant was determined by dividing the frequency response functions, mathematically canceling the effects of feedback. With respect to the visual stimulus, gains for the leg segment showed no differences between the four platform conditions. Phase for the stationary condition was lower at the higher stimulus driving frequencies than for any of the moving platform conditions. In contrast, trunk segment gains were lower for the sway-referenced conditions at lower stimulus frequencies than for the stationary and sum-of-sines conditions. Phase showed a slight lead of the legs over the trunk for the sway-referenced conditions. The phase relationship between the trunk and leg segments, typically in-phase below ~1 Hz and anti-phase above ~1 Hz, showed a gradual transition at a lower frequency for the sway-referenced conditions than for the stationary or sum-of-sines conditions. Complex coherence showed a "legs-leading" coordinative relationship at the phase mode transition for the two sway-referenced conditions. Differences in the frequency response functions demonstrate that the plant changes with platform condition requiring different postural control strategies to maintain stability

On the role of stability in animal morphology and neural control

Mechanical stability is vital for the fitness and survival of animals and is a crucial aspect of robot design and control. Stability depends on multiple factors, including the body\u27s intrinsic mechanical response and feedback control. But feedback control is more fragile than the body\u27s innate mechanical response or open-loop control strategies because of sensory noise and time-delays in feedback. This thesis examines the overarching hypothesis that stability demands have played a crucial role in how animal form and function arise through natural selection and motor learning. In two examples, finger contact and overall body stability, we investigated the relationship between morphology, open-loop control, and stability. By studying the stability of the internal degrees of freedom of a finger when pushing on a hard surface, we find that stability limits the force that we can produce and is a dominant aspect of the neural control of the finger\u27s muscles. In our study on whole body lateral stability during locomotion in terrestrial animals, we find that the overall body aspect ratio has evolved to ensure passive lateral stability on the uneven terrain of natural environments. Precisely gripping an object with the fingertips is a hallmark of human hand dexterity. In Chapter 2, we show how human fingers are intrinsically prone to a buckling-type postural instability and how humans use careful neural orchestration of our muscles so that the elastic response of our muscles can suppress the intrinsic instability. In Chapter 3, we extend these findings further to examine the nature of neuromuscular variability and how the nervous system deals with the need for muscle-induced stability. We find that there is structure to neuromuscular variability so that most of the variability lies within the subspace that does not affect stability. Inspired by the open-loop stable control of our index fingers, in Chapter 4, we derive open-loop stability conditions for a general mechanical linkage with arbitrary joint torques subjected to holonomic constraints. The solution that we derive is physically realizable as cable-driven active mechanical linkages. With a user-prescribed cable layout, we pose the problem of actuating the system to maintain stability while subject to goals like energy minimization as a convex optimization problem. We are thus able to use efficient optimization methods available for convex problems and demonstrate numerical solutions in examples inspired by the finger. Chapter 5 presents a general formulation of the stability criteria for active mechanical linkages subject to Pfaffian holonomic and non-holonomic constraints. Active mechanical linkages subject to multiple constraints represent the mechanics of systems spanning many domains and length scales, such as limbs and digits in animals and robots, and elastic networks like actin meshes in microscopic systems. We show that a constrained mechanical linkage with regular stiffness and damping, and circulation-free feedback, can only destabilize by static buckling when subject to holonomic constraints. In contrast, the same mechanical linkage, subject to a non-holonomic constraint, such as a skate contact, can exhibit either static buckling or flutter instability. Chapter 6 moves away from neural control and studies the shape of animal bodies and their relationship to stability in locomotion. We investigate why small land animals tend to have a crouched or sprawled posture, whereas larger animals are generally more upright. We propose a new hypothesis that the scaling of body aspect ratio with size is driven by the scale-dependent unevenness of natural terrain. We show that the scaling law arising from the need for stability on rough natural terrain correctly predicts the frontal aspect ratio scaling law across 335 terrestrial vertebrates and invertebrates, spanning eight orders of magnitude in mass so that smaller animals have a wider aspect ratio. We also carry out statistical analyses that consider the phylogenetic relationship among the species in our dataset to show that the scaling is not due to gradual changes of the traits over time. Thus, stability demands on natural terrain may have driven the macroevolution of body aspect ratio across terrestrial animals. Interrogating unstable and marginally stable behaviors has helped us identify the morphological and control features that allow animals to perform robustly in noisy environments where perfect sensory feedback cannot be assumed. Although the thesis identifies the `what\u27 and `why,\u27 further studies are needed to understand `how\u27 mechanics and development intertwine to give rise to control and form in growing and adapting biological organisms

Robust muscle synergies for postural control

The musculoskeletal structure of the human and animal body provides multiple solutions for performing any single motor behavior. The long-term goal of the work presented here is to determine the neuromechanical strategies used by the nervous system to appropriately coordinate muscles in order to achieve the performance of daily motor tasks. The overall hypothesis is that the nervous system simplifies muscle coordination by the flexible activation of muscle synergies, defined as a group of muscles activated as a unit, that perform task-level biomechanical functions. To test this hypothesis we investigated whether muscle synergies can be robustly used as building blocks for constructing the spatiotemporal muscle coordination patterns in human and feline postural control under a variety of biomechanical contexts. We demonstrated the generality and robustness of muscle synergies as a simplification strategy for both human and animal postural control. A few robust muscle synergies were able to reproduce the spatial and temporal variability in human and cat postural responses, regardless of stance configuration and perturbation type. In addition inter-trial variability in human postural responses was also accounted for by these muscle synergies. Finally, the activation of each muscle synergy in cat produced a specific stabilizing force vector, suggesting that muscle synergies control task-level variables. The identified muscle synergies may represent general modules of motor output underlying muscle coordination in posture that can be activated in different sensory contexts to achieve different postural goals. Therefore muscle synergies represents a simplifying mechanism for muscle coordination in natural behaviors not only because it is a strategy for reducing the number of variables to be controlled, but because it represents a mechanism for simply controlling multi-segmental task-level variables.Ph.D.Committee Chair: Ting, Lena H.; Committee Member: Chang, Young-Hui; Committee Member: Lee, Robert H.; Committee Member: Nichols, T. Richard; Committee Member: Wolf, Steve L

Identification of Chronic Postural Stability Impairments Associated With History of Concussion

Concussion is the most common form of traumatic brain injury (TBI). However, there is a disproportionate level of understanding between the acute and chronic impairments associated with traumatic brain injury. Specifically, problems maintaining balance during standing and walking are cardinal signs of acute concussion, but the temporal extent to which postural control deficits remain following the initial injury are not well defined or understood. The purpose of the projects composing this dissertation was to examine the long-term effects of a prior history of concussion on static (i.e. standing) and dynamic (i.e. gait) postural control. To address this, healthy adults aged 18-45 reporting a prior history of concussion(s) as well as age-matched controls with no documented concussion history were recruited to participate. Static postural control was assessed using a force plate system to track each participant’s center-of-pressure during standing. Spatiotemporal parameters as well as head stability during gait were assessed using a pressure-sensitive walkway and accelerometers placed at the head, neck, and lower trunk, respectively. The findings of these projects indicate that concussion has detrimental effects on both static and dynamic postural stability years after the initial injury and clinical determination of recovery. Specifically, individuals with a prior history of concussion demonstrated greater postural sway displacement and reduced sway regularity under dual-task conditions compared to the control group. In addition, previously concussed individuals demonstrated less variability in their gait cadence and step length, which suggests a reduction in the complexity of the neural networks contributing to postural control. Lastly, individuals with a history of concussion demonstrated greater triaxial accelerations at the head during gait, indicating a reduced ability to attenuate gait-related oscillations and stabilize the head. Collectively, these findings indicate that concussion is associated with impaired postural control that persists for years after the initial injury and well beyond the point where clinical testing protocols can identify deficits in maintaining balance. Future efforts should be directed toward incorporating more sophisticated measures and analyses of postural stability in concussion screenings to improve clinicians’ abilities to identify the scope in which concussion negatively impacts the function of the central nervous system

Neuromuscular Control Strategy during Object Transport while Walking: Adaptive Integration of Upper and Lower Limb Movements

When carrying an object while walking, a significant challenge for the central nervous system (CNS) is to preserve the object’s stability against the inter-segmental interaction torques and ground reaction forces. Studies documented several strategies used by the CNS: modulation of grip force (GF), alterations in upper limb kinematics, and gait adaptations. However, the question of how the CNS organizes the multi-segmental joint and muscle coordination patterns to deal with gait-induced perturbations remains poorly understood. This dissertation aimed to explore the neuromuscular control strategy utilized by the CNS to transport an object during walking successfully. Study 1 examined the inter-limb coordination patterns of the upper limbs when carrying a cylinder-shaped object while walking on a treadmill. It was predicted that transporting an object in one hand would affect the movement pattern of the contralateral arm to maintain the overall angular momentum. The results showed that transporting an object caused a decreased anti-phase coordination, but it did not induce significant kinematic and muscle activation changes in the unconstrained arm. Study 2 examined muscle synergy patterns for upper limb damping behavior by using non-negative matrix factorization (NNMF) method. Four synergies were identified, showing a proximal-to-distal pattern of activation preceding heel contacts. Study 3 examined the effect of different precision demands (carrying a cup with or without a ball) and altered visual information (looking forward vs. looking at an object) on the upper limb damping behavior and muscle synergies. Increasing precision demand induced stronger damping behavior and increased the electromyography (EMG) activation of wrist/hand flexors and extensors. The NNMF results replicated Study 2 in that the stabilization of proximal joints occurred before the distal joints. The results indicated that the damping incorporates tonic and phasic muscle activation to ensure object stabilization. Overall, three experiments showed that the CNS adopts a similar synergy pattern regardless of task constraint or altered gaze direction while modulating the amount of muscle activation for object stabilization. Kinematic changes can differ depending on the different levels of constraint, as shown in the smaller movement amplitude of the shoulder joint in the transverse plane during the task with higher precision demand

Sports Performance and Health

Sports performance is primarily associated with elite sport, however, recreational athletes are increasingly attempting to emulate elite athletes. Performance optimization is distinctly multidisciplinary. Optimized training concepts and the use of state-of-the-art technologies are crucial for improving performance. However, sports performance enhancement is in constant conflict with the protection of athletes’ health. Notwithstanding the known positive effects of physical activity on health, the prevention and management of sports injuries remain major challenges to be addressed. Accordingly, this Special Issue on "Sports Performance and Health" consists of 17 original research papers, one review paper, and one commentary, and covers a wide range of topics related to fatigue, movement asymmetries, optimization of sports performance by training, technique, and/or tactics enhancements, prevention and management of sports injuries, optimization of sports equipment to increase performance and/or decrease the risk of injury, and innovations for sports performance, health, and load monitoring. As this Special Issue offers several new insights and multidisciplinary perspectives on sports performance and health, readers from around the world who work in these areas are expected to benefit from this Special Issue collection