|dc.description.abstract||Modern human shoulder function is affected by the evolutionary adaptations that have occurred to ensure survival and prosperity of the species. Robust examination of behavioral shoulder performance and injury risk can be holistically improved through an interdisciplinary approach that integrates anthropology and biomechanics. Anthropological research methods have attempted to resolve gaps in human shoulder evolution, while biomechanics research has attempted to explain the musculoskeletal function of the modern human shoulder. Coordination of these two fields can allow different perspectives to contribute to a more complete interpretation of, not only how the modern human shoulder is susceptible to specific injuries, but also why.
How the modern human shoulder arose from a, likely, weight-bearing, arboreal past to its modern form, and what this has meant for modern behaviors, is not well understood. Despite a weight-bearing, locomotor ancestral usage, the modern human upper extremity is highly fatigable in repetitive, low to moderate force tasks, such as overhead reaching. The closest living human relative, modern chimpanzees, has retained an arboreal, locomotor upper extremity. Interdisciplinary comparative research on humans and chimpanzees could lead to greater insight on modern human shoulder function. The purpose of this research was to explore the modern human capacity for ancestral, brachiating behaviors and resultant injury mechanisms through comparative experimental, computational modeling and probabilistic modeling approaches with chimpanzees.
The first study experimentally explored the modern human ability to perform a horizontal bimanual arm-suspensory climbing task. EMG of 12 upper extremity muscles and motion capture of the arm and thorax were monitored in experienced and inexperienced climbers. Results were also compared to previously published or collected data on chimpanzees performing an analogous task. While all human climbers used a high proportion of their muscular reserve to perform the task, experienced climbers had moderately reduced muscle activity in most muscles, particularly during phasic shifts of the climb cycle and moderately more efficient kinematics. Climbing kinematics and muscle activity were very similar between humans and chimpanzees. However, chimpanzees appear to have a different utility of the posterior deltoid, suggesting an influence of their habitual arboreal behaviors.
The second and third studies describe the development of a deterministic chimpanzee musculoskeletal glenohumeral model. Study 2 developed geometric parameters of chimpanzee shoulder rhythm and glenoid stability ratios for the construction of a chimpanzee glenohumeral model. The chimpanzee glenohumeral model of Study 3 was designed to parallel an existing human glenohumeral model, enabling comparative analyses. The chimpanzee glenohumeral model consists of three modules – an external torque module, musculoskeletal geometric module, and an internal muscle force prediction module. Together, these modules use postural kinematics, subject specific anthropometrics and hand forces to estimate joint reaction forces and moments, subacromial space dimensions, and muscle and tissue forces. Using static postural data from Study 1, predicted muscle forces and subacromial space were compared between chimpanzees and humans during an overhead, weight-bearing climbing task. Compared to chimpanzees, the human model predicted a 2mm narrower subacromial space, deltoid muscle forces that were often double those of chimpanzees and a strong reliance on infraspinatus and teres minor (60-100% maximal force) over other rotator cuff muscles.
Finally, the deterministic chimpanzee and human glenohumeral models were expanded in Study 4 to a probabilistic analysis of rotator cuff function between species. Using probabilistic software and the same postural climbing inputs, both models had anthropologically relevant musculoskeletal features perturbed in a series of Monte Carlo simulations – muscle origins and insertions, glenoid inclination and glenoid stability – to determine if rotator cuff muscle force prediction distributions would converge between species. Human rotator cuff muscle behavior did not converge with chimpanzees using probabilistic simulation. The human model continued to predict strong dependence on infraspinatus and teres minor, with 99% confidence intervals of [0-100]% maximal force, over supraspinatus and subscapularis, with confidence intervals of [0-5]% maximal force. Chimpanzee rotator cuff confidence intervals were typically between [0-40]% maximal force, with median force for all four rotator cuff muscles typically 5-20% maximal force. While perturbation of muscle origins and insertions had the greatest effect on muscle force output distributions, no musculoskeletal variation notably modified human climbing performance.
Structural musculoskeletal differences between species dictated differences in glenohumeral function. The results from all studies indicate susceptibility for the fatigue-induced initiation of subacromial impingement syndrome and rotator cuff pathology in modern humans during overhead and repetitive tasks. Lower muscle absolute PCSA in humans, combined with a laterally oriented glenohumeral joint and laterally projecting acromion reduced the capacity for overhead postures and weight-bearing postures. These evolutionary differences may have been vestigial consequences, concurrent with necessary adaptions for important, evolutionary human-centric behaviors such as throwing. However, they have influenced the high rates of rotator cuff pathology in humans compared to closely related primates.
The present work represents an important first step toward a broad scope of future research. Interdisciplinary computational modeling offers an evolving and improving alternative to traditional methods to study human evolution and function. Computational and probabilistic simulations can be expanded to numerous other biomechanical and evolutionary queries. The results of this thesis are a promising initial step to examining the evolutionary structural connection to biomechanical human function through comparative computational modeling.||en