Patellofemoral pain (PFP) is believed to be associated with deficits in coordination between the different heads of the quadriceps muscle; however, considerable debate exists in the literature regarding the presence of such a deficit. Discrepancies between studies may be explained by differences in experimental tasks, such as whether the task is performed with open (OKC) or closed kinetic chain (CKC), or whether the activity is voluntary or triggered. Particular interest has been directed toward the function of the vastus medialis obliquus (VMO), which is a short muscle with limited ability to exert torque across the knee joint, but probably has a particular role in controlling patellofemoral joint position. Another short muscle that may influence knee joint position control is popliteus (POP), which is located in the back of the knee. This thesis investigates task specific activity of quadriceps in CKC versus OKC and studies the relative activity between the four heads of the quadriceps in PFP subjects compared to controls without knee pain in voluntary activity (CKC and OKC) and postural responses to balance perturbations. In addition, this thesis investigates the presumed function of POP for control of joint position in postural tasks in healthy individuals. All subjects were of normal weight and height and between 18 and 40 years. Quadriceps activity was tested for isometric with identical joint configuration in CKC and OKC, and it was performed as a reaction time task. Balance perturbations were elicited by unpredictable anterior and posterior translations of the support surface. Function of POP was investigated in unpredictable support surface translations and in self induced provocations to balance by moving the arms. Muscle activity was recorded with electromyography (EMG). Optic kinematic analysis was used to obtain specific movement responses to perturbations of balance. The quadriceps muscles were activated differently in CKC and OKC. VMO was activated earlier and to a greater degree in CKC. Rectus femoris was activated earlier and to a greater degree in OKC. PFP subjects reacted slower in both CKC and OKC, but there was no difference between groups in the relative activity between the different heads of the quadriceps. In the unpredictable support surface translation in the anterior direction, PFP subjects responded with earlier onset of VMO and with greater trunk and hip flexion in the anterior translation. POP activation in response to support surface translations in both directions occurred before all other muscles measured. In the self-initiated provocations of balance, POP was activated after the initiation of the balance provocation. This thesis concludes that quadriceps activity was task specific. The lack of difference between groups in OKC and CKC, and the difference between groups in postural responses suggest that variations in motor behaviour may occur only in tasks habitually performed. Differences in muscle activation patterns may be related to compensatory strategies to unload the quadriceps muscles and the patellofemoral joint. Furthermore, this thesis suggests that POP muscle may have a particular role in active control of the knee joint.

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.

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The issue of the laterality of control during unimanual and bimanual coordination was addressed in this thesis. Two tasks were used throughout: a repetitive discrete response task (finger tapping) and a continuous task (circle-drawing). Different mechanisms have been implicated in the temporal control of repetitive discrete movements and continuous movements. The tasks also differ in the degree of spatiotemporal coordination required which might have important implications in the question of laterality of control. The first section of the thesis examined between-hand differences in the dynamics of performance during unimanual and bimanual coordination. During tapping, the dominant hand was faster and less temporally variable than the nondominant hand. During circle drawing the dominant hand was faster, more accurate, less temporally and spatially variable, and produced smoother trajectories than the nondominant hand. During bimanual coordination, several of these asymmetries were attenuated: the rate of movement of the two hands became equivalent (the hands became temporally coupled), the asymmetry in temporal variability during tapping was reduced, and the asymmetry in trajectory smoothness during circle drawing was reduced. The second section of the thesis examined the effects of disrupting motor processes with transcranial magnetic stimulation (TMS) over the left or right primary motor cortex (M1) on the ongoing performance of the hands. In the first study, TMS over left or right M1 during unimanual tapping caused large disruptions to tapping with the contralateral hand but had little effect on the ipsilateral hand. In contrast, for a subset of trials during bimanual tapping, two lateralized effects of stimulation were seen: the effect of TMS on the contralateral hand was greater after stimulation over left M1 than after stimulation over right M1, and prolonged changes in inter-tap interval were observed in the left hand regardless of the side of stimulation. In the second study, TMS over left M1 during circle drawing decreased the accuracy of drawing with both the contralateral and ipsilateral hand, whereas TMS over right M1 decreased accuracy of drawing only with the contralateral hand. This lateralized effect was not limited to the bimanual case, but was also apparent during unimanual drawing. The final chapter addressed issues in bimanual motor control after unilateral stroke. Performance of the affected limb was examined during unimanual and bimanual coordination in a group of stroke patients with varying levels of impairment. The results indicated an improvement in the performance of the affected limb for some patients with mild to moderate, but not severe upper limb motor deficits during bimanual movement. The improvements were limited to the patients who showed evidence of temporal coupling between the hands. These findings support the hypothesis that the dominant motor cortex has a role in the control of both hands during bimanual coordination. In addition, the dominant hemisphere appears to play a role in controlling both hands during unimanual movements which require a greater degree of spatiotemporal coordination. The final study suggests that temporal coupling between the limbs is crucial for the facilitation of performance of the affected limb during bimanual coordination, which has both theoretical and practical implications.

Neuromuscular electrical stimulation (NMES) has become an increasingly popular rehabilitative treatment approach for swallowing disorders (dysphagia). However, its precise effects on swallowing biomechanics and measures of swallowing neurophysiology are unclear. Clearly defined NMES treatment protocols that have been corroborated by thorough empirical research are lacking. The primary objective of this research programme was therefore to establish optimal NMES treatment parameters for the anterior hyo-mandibular (submental) musculature, a muscle group that is critically involved in the oral and pharyngeal phases of swallowing. Based on previous research, the primary hypothesis was that various NMES treatment protocols would have differential effects of either enhancing or inhibiting the excitability of corticobulbar projections to this muscle group. The research paradigm used to test this hypothesis was an evaluation of MEP amplitude and onset latency, recorded in the functional context of volitional contraction of the submental musculature (VC) and contraction of this muscle group during the pharyngeal phase of volitional swallowing (VPS, volitional pharyngeal swallow). Outcome measures were recorded before and at several time points after each NMES treatment trial. This methodology is similar to, but improved upon, research paradigms previously reported. Changes in corticobulbar excitability in response to various NMES treatment protocols were recorded in a series of experiments. Ten healthy research participants were recruited into a study that evaluated the effects of event-related NMES, whereas 15 healthy research participants were enrolled in a study that investigated the effects of non-event-related NMES. In a third cohort of 35 healthy research participants, task-dependent differences in corticobulbar excitability were evaluated during three conditions of submental muscle contraction: VC, VPS and submental muscle contraction during the pharyngeal phase of reflexive swallowing (RPS, reflexive pharyngeal swallowing). Event-related NMES induced frequency-depended changes in corticobulbar excitability. NMES administered at 80 Hz facilitated MEP amplitude, whereas NMES at 5 Hz and 20 Hz inhibited MEP amplitude. No changes were observed after NMES at 40 Hz. Maximal excitatory or inhibitory changes occurred 60 min post-treatment. Changes in MEP amplitude in response to event-related NMES were only observed when MEPs were recorded during the VC condition, whereas MEPs recorded during the VPS condition remained unaffected. Non-event-related NMES did not affect MEP amplitude in either of the muscle contraction conditions. Similarly, MEP onset latencies remained unchanged across all comparisons. MEPs were detected most consistently during the VC contraction condition. They were less frequently detected and were smaller in amplitude for the VPS condition and they were infrequently detected during pre-activation by RPS. The documented results indicate that event-related NMES has a more substantial impact on MEP amplitude than non-event-related NMES, producing excitatory and inhibitory effects. Comparison of MEPs recorded during VC, VPS and RPS suggests that different neural networks may govern the motor control of submental muscle activation during these tasks. This research programme is the first to investigate the effects of various NMES treatment protocols on the excitability of submental corticobulbar projections. It provides important new information for the use of NMES in clinical rehabilitation practices and our understanding of the neural networks governing swallowing motor control.

L’apprentissage du langage parlé nécessite un contrôle musculaire très précis des différents organes intervenant dans la production de la parole. La production de sons voisés, qui résulte de l’auto-oscillation des cordes vocales, est notamment influencée par l’ensemble du système phonatoire, du diaphragme jusqu’aux lèvres. Les travaux réalisés dans le cadre de cette thèse s’inscrivent dans un contexte de modélisation physique de la parole. Les objectifs s’articulent autour de la compréhension des phénomènes physiques gouvernant la production de sons voisés. Les études sont appliquées à des cas pour lesquels le contrôle de la production est fortement altéré, lorsque le locuteur souffre de pertes auditives importantes. Dans ce cas de figure, les interactions physiques peuvent jouer un rôle important dans l’apparition de troubles de la production. La démarche adoptée consiste alors dans un premier temps à observer les phénomènes étudiés au moyen de mesures in vivo puis à proposer des modèles théoriques mécanique, aérodynamique et acoustique permettant de les décrire. Ensuite, les modèles sont validés en comparant avec des mesures réalisées sur une maquette de l’appareil phonatoire. Finalement,des simulations numériques temporelles basées sur un modèle à deux masses pour décrire le comportement mécanique des cordes vocales, permettent de tester les modèles physiques pour des cas concrets de production. La première étude se concentre sur les sauts de fréquence fondamentale qui accompagnent les transitions involontaires entre deux mécanismes laryngés lors de la production d’une voyelle. Les travaux expérimentaux et numériques montrent qu’une transition de mécanisme laryngé est la manifestation d’une bifurcation du système laryngé et que ces bifurcations se produisent lors d’une variation de la raideur des cordes vocales, de la pression sous-glottique, de l’aire glottique initiale ou bien de la longueur des résonateurs acoustiques. Les modèles théoriques permettent de reproduire les sauts de fréquence fondamentale observés expérimentalement. Ils sont utilisés pour étudier les différentes stratégies motrices responsables de ces sauts de fréquence. La deuxième étude porte sur la production de consonnes plosives, en particulier sur les effets de la réalisation d’une occlusion du conduit vocal sur l’arrêt puis l’apparition de l’oscillation des cordes vocales. Les simulations de séquences voyelle - plosive bilabiale non voisée (/p/)-voyelle effectuées montrent que l’expansion passive de la cavité supraglottique est à l’origine du maintien de l’auto-oscillation des cordes vocales après la fermeture du conduit vocal et que l’augmentation de la longueur du conduit vocal a pour effet de réduire le délai entre le relâchement de l’occlusion et l’apparition de l’oscillation des cordes vocales. Ces résultats impliquent que l’articulation joue un rôle considérable sur le mode de voisement (voisée ou non voisée) de la consonne et sur la valeur du Voice Onset Time pour une consonne plosive non voisée. / Language learning requires specific muscle control of all organs that contribute to speech production. Voiced sounds production, which results from vocal folds self oscillation, is especially influenced by the whole phonatory apparatus, from diaphragm to lips. The general background of this thesis is the physical modeling of speech production and the objectives are motivated by a better comprehension of physical phenomena occurring in voiced sounds production. In the frame of this work, studies are focused on cases where speech production control is impaired, for example when the speaker suffers from an important hearing loss. In this situation, physical interactions can play an important role in speech production disorders emergence. The approach adopted here is first to observe the studied phenomena thanks to invivo measurements then to describe them thanks to theoretical models. Thereafter, the models are validated by comparing theoretical results with measurements performed on a replica of the phonatory apparatus. Finally, numerical simulations in the time domain, based on a two-mass model, allow to apply physical models to specific speech production occurrences.The first study deals with fundamental frequency jumps that are observed during an unvoluntary transition between two different laryngeal mechanisms in case of vowel production.Experimental and numerical results highlight that a transition between two different laryngeal mechanisms is a symptom of the laryngeal system bifurcation, and that such bifurcation occurs during a variation of the vocal folds stiffness, the subglottal pressure,the prephonatory glottal area or the acoustic resonators length. The theoretical models allow to simulate the fundamental frequency jumps that are observed experimentally. They are used to study the different motor strategies responsible for these frequency jumps.The second study deals with plosive consonants production, and in particular with the effectsof a vocal tract occlusion on voicing offset and onset. Simulations of vowel – voiceless plosive - vowel production highlight that passive expansion of the supraglottal cavity is responsible for the voicing extension after vocal tract closure, and that increase of the vocal tract length leads to a shorter delay between the vocal tract occlusion release and the voicing onset. These results highlight that the articulation plays an important role in voicing (voiced or voiceless) and in voice-onset-time value for a voiceless plosive.

abstract: Anticipatory planning of digit positions and forces is critical for successful dexterous object manipulation. Anticipatory (feedforward) planning bypasses the inherent delays in reflex responses and sensorimotor integration associated with reactive (feedback) control. It has been suggested that feedforward and feedback strategies can be distinguished based on the profile of grip and load force rates during the period between initial contact with the object and object lift. However, this has not been validated in tasks that do not constrain digit placement. The purposes of this thesis were (1) to validate the hypothesis that force rate profiles are indicative of the control strategy used for object manipulation and (2) to test this hypothesis by comparing manipulation tasks performed with and without digit placement constraints. The first objective comprised two studies. In the first study an additional light or heavy mass was added to the base of the object. In the second study a mass was added, altering the object's center of mass (CM) location. In each experiment digit force rates were calculated between the times of initial digit contact and object lift. Digit force rates were fit to a Gaussian bell curve and the goodness of fit was compared across predictable and unpredictable mass and CM conditions. For both experiments, a predictable object mass and CM elicited bell shaped force rate profiles, indicative of feedforward control. For the second objective, a comparison of performance between subjects who performed the grasp task with either constrained or unconstrained digit contact locations was conducted. When digit location was unconstrained and CM was predictable, force rates were well fit to a bell shaped curve. However, the goodness of fit of the force rate profiles to the bell shaped curve was weaker for the constrained than the unconstrained digit placement condition. These findings seem to indicate that brain can generate an appropriate feedforward control strategy even when digit placement is unconstrained and an infinite combination of digit placement and force solutions exists to lift the object successfully. Future work is needed that investigates the role digit positioning and tactile feedback has on anticipatory control of object manipulation. / Dissertation/Thesis / M.S. Bioengineering 2011