Temporomandibular joint (TMJ) diseases such as osteoarthritis and disc displacement have no permanent treatment options, but lubrication therapies, used in other joints, could be an effective alternative. However, the healthy TMJ contains fibrocartilage, not hyaline cartilage as is found in other joints. As such, the effect of lubrication therapies in the TMJ is unknown. Additionally, only a few studies have characterized the friction coefficient of the healthy TMJ. Like other cartilaginous tissues, the mandibular condyles and discs are subject to changes in friction coefficient due to fluid pressurization. In addition, the friction coefficients of the inferior joint space of the TMJ are affected by the sliding direction and anatomic location. However, these previous findings have not been able to identify how all three of these parameters (anatomic location, sliding direction, and fluid pressurization) influence changes in friction coefficient. This study used Stribeck curves to identify differences in the friction coefficients of mandibular condyles and discs based on anatomic location, sliding direction, and amount of fluid pressurization (friction mode). Friction coefficients were measured using a cartilage on glass tribometer. Both mandibular condyle and disc friction coefficients were well described by Stribeck curves (R 2 range 0.87–0.97; p < 0.0001). These curves changed based on anatomic location (Δ μ ∼ 0.05), but very few differences in friction coefficients were observed based on sliding direction. Mandibular condyles had similar boundary mode and elastoviscous mode friction coefficients to the TMJ disc ( μ min ∼ 0.009 to 0.19) and both were lower than hyaline cartilage in other joints (e.g., knee, ankle, etc.). The observed differences here indicate that the surface characteristics of each anatomic region cause differences in friction coefficients.

The sixth temporomandibular joint (TMJ) Bioengineering Conference (TMJBC) was held on June 14–15 2018, in Redondo Beach, California, 12 years after the first TMJBC. Speakers gave 30 presentations and came from the United States, Europe, Asia, and Australia. The goal of the conference has remained to foster a continuing forum for bioengineers, scientists, and surgeons and veterinarians to advance technology related to TMJ disorders. These collective multidisciplinary interactions over the past decade have made large strides in moving the field of TMJ research forward. Over the past 12 years, in vivo approaches for tissue engineering have emerged, along with a wide variety of degeneration models, as well as with models occurring in nature. Furthermore, biomechanical tools have become more sensitive and new biologic interventions for disease are being developed. Clinical directives have evolved for specific diagnoses, along with patient-specific biological and immunological responses to TMJ replacement devices alloplastic and/or bioengineered devices. The sixth TMJBC heralded many opportunities for funding agencies to advance the field: (1) initiatives on TMJ that go beyond pain research, (2) more training grants focused on graduate students and fellows, (3) partnership funding with government agencies to translate TMJ solutions, and (4) the recruitment of a critical mass of TMJ experts to participate on grant review panels. The TMJ research community continues to grow and has become a pillar of dental and craniofacial research, and together we share the unified vision to ultimately improve diagnoses and treatment outcomes in patients affected by TMJ disorders.

Total disc arthroplasty (TDA) using an artificial disc (AD) is an attractive surgical technique for the treatment of spinal disorders, since it can maintain or restore spinal motion (unlike interbody fusion). However, adverse surgical outcomes of contemporary lumbar TDAs have been reported. We previously proposed a new mobile-bearing AD design concept featuring a biconcave ultra-high-molecular-weight polyethylene mobile core. The objective of this study was to develop an artificial neural network based multiobjective optimization framework to refine the biconcave-core AD design considering multiple TDA performance metrics, simultaneously. We hypothesized that there is a trade-off relationship between the performance metrics in terms of range of motion, facet joint force, and polyethylene contact pressure. By searching the resulting three-dimensional Pareto frontier after multiobjective optimization, it was found that there was a "best-trade-off" AD design, which could balance all the three metrics, without excessively sacrificing each metric. However, for each single-objective optimum AD design, only one metric was optimal, and distinct sacrifices were observed in the other two metrics. For a commercially available biconvex-core AD design, the metrics were even worse than the poorest outcomes of the single-objective optimum AD designs. Therefore, multiobjective design optimization could be useful for achieving native lumbar segment biomechanics and minimal polyethylene contact pressures, as well as for improving existing lumbar motion-preserving surgical treatments.

Fibrocartilage is found in the knee meniscus, the temporomandibular joint (TMJ) disc, the pubic symphysis, the annulus fibrosus of intervertebral disc, tendons, and ligaments. These tissues are notoriously difficult to repair due to their avascularity, and limited clinical repair and replacement options exist. Tissue engineering has been proposed as a route to repair and replace fibrocartilages. Using the knee meniscus and TMJ disc as examples, this review describes how fibrocartilages can be engineered toward translation to clinical use. Presented are fibrocartilage anatomy, function, epidemiology, pathology, and current clinical treatments because they inform design criteria for tissue engineered fibrocartilages. Methods for how native tissues are characterized histomorphologically, biochemically, and mechanically to set gold standards are described. Then provided is a review of fibrocartilage-specific tissue engineering strategies, including the selection of cell sources, scaffold or scaffold-free methods, and biochemical and mechanical stimuli. In closing, the Food and Drug Administration (FDA) paradigm is discussed to inform researchers of both the guidance that exists and the questions that remain to be answered with regard to bringing a tissue engineered fibrocartilage product to the clinic.

Aging and degeneration of the intervertebral disk are noted by changes in tissue composition and geometry, including a decrease in nucleus pulposus (NP) area. The NP centroid is positioned slightly posterior of the disk's centroid, but the effect of NP size and location on disk joint mechanics is not well understood. We evaluated the effect of NP size and centroid location on disk joint mechanics under dual-loading modalities (i.e., compression in combination with axial rotation or bending). A finite element model (FEM) was developed to vary the relative NP area (NP:Disk area ratio range = 0.21–0.60). We also evaluated the effect of NP position by shifting the NP centroid anteriorly and posteriorly. Our results showed that compressive stiffness and average first principal strains increased with NP size. Under axial compression, stresses are distributed from the NP to the annulus, and stresses were redistributed toward the NP with axial rotation. Moreover, peak stresses were greater for disks with a smaller NP area. NP centroid location had a greater impact on intradiscal pressure during flexion and extension, where peak pressures in the posterior annulus under extension was greater for disks with a more posteriorly situated NP. In conclusion, the findings from this study highlight the importance of closely mimicking NP size and location in computational models that aim to understand stress/strain distribution during complex loading and for developing repair strategies that aim to recapitulate the mechanical behavior of healthy disks.

Predicting the mechanical behavior of the intervertebral disk (IVD) in health and in disease requires accurate spatial mapping of its compressive mechanical properties. Previous studies confirmed that residual strains in the annulus fibrosus (AF) of the IVD, which result from nonuniform extracellular matrix deposition in response to in vivo loads, vary by anatomical regions (anterior, posterior, and lateral) and zones (inner, middle, and outer). We hypothesized that as the AF is composed of a nonlinear, anisotropic, viscoelastic material, the state of residual strain in the transverse plane would influence the apparent values of axial compressive properties. To test this hypothesis, axial creep indentation tests were performed, using a 1.6 mm spherical probe, at nine different anatomical locations on bovine caudal AFs in both the intact (residual strain present) and strain relieved states. The results showed a shift toward increased spatial homogeneity in all measured parameters, particularly instantaneous strain. This shift was not observed in control AFs, which were tested twice in the intact state. Our results confirm that time-dependent axial compressive properties of the AF are sensitive to the state of residual strain in the transverse plane, to a degree that is likely to affect whole disk behavior.

A spine is proven to be subjected to a follower load which is a compressive load of physiologic magnitude acting on the whole spine. The path of the follower load approximates the tangent to the curve of the spine in in vivo neutral standing posture. However, the specific path location of the follower load is still unclear. The aim of this study is to find out the most realistic location of the follower load path (FLP) for a lumbar spine in standing. A three-dimensional (3D) nonlinear finite element model (FEM) of lumbosacral vertebrae (L1-S1) with consideration of the calibrated material properties was established and validated by comparing with the experimental data. We show that the shape of the lumbosacral spine is strongly affected by the location of FLP. An evident nonlinear relationship between the FLP location and the kinematic response of the L1-S1 lumbosacral spine exists. The FLP at about 4 and 3 mm posterior to the curve connecting the center of the vertebral bodies delivers the most realistic location in standing for healthy people and patients having low back pains (LPBs), respectively. Moreover, the “sweeping” method introduced in this study can be applicable to all individualized FEM to determine the location of FLP.

Intervertebral disk (IVD) degeneration is commonly described by loss of height and hydration. However, in the first stage of IVD degeneration, this loss has not yet occurred. In the current study, we use an ex vivo degeneration model to analyze the changes in IVDs mechanical behavior in the first phase of degeneration. We characterize these changes by stretched-exponential fitting, and suggest the fitted parameters as markers for early degeneration. Enzymatic degeneration of healthy lumbar caprine IVDs was induced by injecting 100 μ L of Chondroïtinase ABC (Cabc) into the nucleus. A no-intervention and phosphate buffered saline (PBS) injected group were used as controls. IVDs were cultured in a bioreactor for 20 days under diurnal, simulated-physiological loading (SPL) conditions. Disk deformation was continuously monitored. Changes in disk height recovery behavior were quantified using stretched-exponential fitting. Disk height, histological sections, and water- and glycosaminoglycan (GAG)-content measurements were used as gold standards for the degenerative state. Cabc injection caused significant GAG loss from the nucleus and had detrimental effects on poro-elastic mechanical properties of the IVDs. These were progressive over time, with a propensity toward more linear recovery behavior. On histological sections, both PBS and Cabc injected IVDs showed moderate degeneration. A small GAG loss yields changes in IVD recovery behavior, which can be quantified with stretched-exponential fitting. Parameters changed significantly compared to control. Studies on disk degeneration and biomaterial engineering for degenerative disk disease (DDD) could benefit from focusing on IVD biomechanical behavior rather than height and water-content, as a marker for early disk degeneration.

A subset of temporomandibular joint (TMJ) disorders is attributed to joint degeneration. The pig has been considered the preferred in vivo model for the evaluation of potential therapies for TMJ disorders, and practical considerations such as cost and husbandry issues have favored the use of young, skeletally immature animals. However, the effect of growth on the biochemical and biomechanical properties of the TMJ disk and articulating cartilage has not been examined. The present study investigates the effect of age on the biochemical and biomechanical properties of healthy porcine TMJs at 3, 6, and 9 months of age. DNA, hydroxyproline, and glycosaminoglycan (GAG) content were determined and the disks and condyles were tested in uniaxial unconfined stress relaxation compression from 10% to 30% strain. TMJ disks were further assessed with a tensile test to failure technique, which included the ability to test multiple samples from the same region of an individual disk to minimize the intraspecimen variation. No differences in biochemical properties for the disk or compressive properties at 30% stress relaxation in the disk and condylar cartilage were found. In tension, no differences were observed for peak stress and tensile modulus. The collagen content of the condyle was higher at 9 months than 3 months (p < 0.05), and the GAG content was higher at 9 months than 6 months (p < 0.05). There was a trend of increased compressive instantaneous modulus with age. As such, age-matched controls for growing pigs are probably appropriate for most parameters measured.

There is lack of investigation capturing the complex mechanical interaction of tissue-engineered intervertebral disk (IVD) constructs in physiologically relevant environmental conditions. In this study, mechanical characterization of anisotropic electrospinning (ES) substrates made of polycaprolactone (PCL) was carried out in wet and dry conditions and viability of human bone marrow derived mesenchymal stem cells (hMSCs) seeded within double layers of ES PCL were also studied. Cyclic compression of IVD-like constructs composed of an agarose core confined by ES PCL double layers was implemented using a bioreactor and the cellular response to the mechanical stimulation was evaluated. Tensile tests showed decrease of elastic modulus of ES PCL as the angle of stretching increased, and at 60 deg stretching angle in wet, the maximum ultimate tensile strength (UTS) was observed. Based on the configuration of IVD-like constructs, the calculated circumferential stress experienced by the ES PCL double layers was 40 times of the vertical compressive stress. Confined compression of IVD-like constructs at 5% and 10% displacement dramatically reduced cell viability, particularly at 10%, although cell presence in small and isolated area can still be observed after mechanical conditioning. Hence, material mechanical properties of tissue-engineered scaffolds, composed of fibril structure of polymer with low melting point, are affected by the testing condition. Circumferential stress induced by axial compressive stimulation, conveyed to the ES PCL double layer wrapped around an agarose core, can affect the viability of cells seeded at the interface, depending on the mechanical configuration and magnitude of the load.

Intervertebral disc degeneration is a prevalent phenomenon associated with back pain. It is of critical clinical interest to discriminate disc health and identify early stages of degeneration. Traditional clinical T2-weighted magnetic resonance imaging (MRI), assessed using the Pfirrmann classification system, is subjective and fails to adequately capture initial degenerative changes. Emerging quantitative MRI techniques offer a solution. Specifically, T2* mapping images water mobility in the macromolecular network, and our preliminary ex vivo work shows high predictability of the disc's glycosaminoglycan content (s-GAG) and residual mechanics. The present study expands upon this work to predict the biochemical and biomechanical properties in vivo and assess their relationship with both age and Pfirrmann grade. Eleven asymptomatic subjects (range: 18–62 yrs) were enrolled and imaged using a 3T MRI scanner. T2-weighted images (Pfirrmann grade) and quantitative T2* maps (predict s-GAG and residual stress) were acquired. Surface maps based on the distribution of these properties were generated and integrated to quantify the surface volume. Correlational analyses were conducted to establish the relationship between each metric of disc health derived from the quantitative T2* maps with both age and Pfirrmann grade, where an inverse trend was observed. Furthermore, the nucleus pulposus (NP) signal in conjunction with volumetric surface maps provided the ability to discern differences during initial stages of disc degeneration. This study highlights the ability of T2* mapping to noninvasively assess the s-GAG content, residual stress, and distributions throughout the entire disc, which may provide a powerful diagnostic tool for disc health assessment.

Advanced computational human body models (HBM) enabling enhanced safety require verification and validation at different levels or scales. Specifically, the motion segments, which are the building blocks of a detailed neck model, must be validated with representative experimental data to have confidence in segment and, ultimately, full neck model response. In this study, we introduce detailed finite element motion segment models and assess the models for quasi-static and dynamic loading scenarios. Finite element segment models at all levels in the lower human cervical spine were developed from scans of a 26-yr old male subject. Material properties were derived from the in vitro experimental data. The segment models were simulated in quasi-static loading in flexion, extension, lateral bending and axial rotation, and at dynamic rates in flexion and extension in comparison to previous experimental studies and new dynamic experimental data introduced in this study. Single-valued experimental data did not provide adequate information to assess the model biofidelity, while application of traditional corridor methods highlighted that data sets with higher variability could lead to an incorrect conclusion of improved model biofidelity. Data sets with continuous or multiple moment–rotation measurements enabled the use of cross-correlation for an objective assessment of the model and highlighted the importance of assessing all motion segments of the lower cervical spine to evaluate the model biofidelity. The presented new segment models of the lower cervical spine, assessed for range of motion and dynamic/traumatic loading scenarios, provide a foundation to construct a biofidelic model of the spine and neck, which can be used to understand and mitigate injury for improved human safety in the future.

While abnormal loading is widely believed to cause cervical spine disc diseases, in vivo cervical disc deformation during dynamic neck motion has not been well delineated. This study investigated the range of cervical disc deformation during an in vivo functional flexion–extension of the neck. Ten asymptomatic human subjects were tested using a combined dual fluoroscopic imaging system (DFIS) and magnetic resonance imaging (MRI)-based three-dimensional (3D) modeling technique. Overall disc deformation was determined using the changes of the space geometry between upper and lower endplates of each intervertebral segment (C3/4, C4/5, C5/6, and C6/7). Five points (anterior, center, posterior, left, and right) of each disc were analyzed to examine the disc deformation distributions. The data indicated that between the functional maximum flexion and extension of the neck, the anterior points of the discs experienced large changes of distraction/compression deformation and shear deformation. The higher level discs experienced higher ranges of disc deformation. No significant difference was found in deformation ranges at posterior points of all the discs. The data indicated that the range of disc deformation is disc level dependent and the anterior region experienced larger changes of deformation than the center and posterior regions, except for the C6/7 disc. The data obtained from this study could serve as baseline knowledge for the understanding of the cervical spine disc biomechanics and for investigation of the biomechanical etiology of disc diseases. These data could also provide insights for development of motion preservation surgeries for cervical spine.

Knowledge of spinal loads in neighboring disks after interbody fusion plays an important role in the clinical decision of this treatment as well as in the elucidation of its effect. However, controversial findings are still noted in the literature. Moreover, there are no existing models for efficient prediction of intervertebral disk stresses within annulus fibrosus (AF) and nucleus pulposus (NP) regions. In this present study, a new hybrid rigid-deformable modeling workflow was established to quantify the mechanical stress behaviors within AF and NP regions of the L1–2, L2–3, and L4–5 disks after interbody fusion at L3–4 level. The changes in spinal loads were compared with results of the intact model without interbody fusion. The fusion outcomes revealed maximal stress changes (10%) in AF region of L1–2 disk and in NP region of L2–3 disk. The minimal stress change (1%) is noted at the NP region of the L1–2 disk. The validation of simulation outcomes of fused and intact lumbar spine models against those of other computational models and in vivo measurements showed good agreements. Thus, this present study may be used as a novel design guideline for a specific implant and surgical scenario of the lumbar spine disorders.

Standing balanced reach is a fundamental task involved in many activities of daily living that has not been well analyzed quantitatively to assess and characterize the multisegmental nature of the body's movements. We developed a dynamic balanced reach test (BRT) to analyze performance in this activity; in which a standing subject is required to maintain balance while reaching and pointing to a target disk moving across a large projection screen according to a sum-of-sines function. This tracking and balance task is made progressively more difficult by increasing the disk's overall excursion amplitude. Using kinematic and ground reaction force data from 32 young healthy subjects, we investigated how the motions of the tracking finger and whole-body center of mass (CoM) varied in response to the motion of the disk across five overall disk excursion amplitudes. Group representative performance statistics for the cohort revealed a monotonically increasing root mean squared (RMS) tracking error (RMSE) and RMS deviation (RMSD) between whole-body CoM (projected onto the ground plane) and the center of the base of support (BoS) with increasing amplitude (p < 0.03). Tracking and CoM response delays remained constant, however, at 0.5 s and 1.0 s, respectively. We also performed detailed spectral analyses of group-representative response data for each of the five overall excursion amplitudes. We derived empirical and analytical transfer functions between the motion of the disk and that of the tracking finger and CoM, computed tracking and CoM responses to a step input, and RMSE and RMSD as functions of disk frequency. We found that for frequencies less than 1.0 Hz, RMSE generally decreased, while RMSE normalized to disk motion amplitude generally increased. RMSD, on the other hand, decreased monotonically. These findings quantitatively characterize the amplitude- and frequency-dependent nature of young healthy tracking and balance in this task. The BRT is not subject to floor or ceiling effects, overcoming an important deficiency associated with most research and clinical instruments used to assess balance. This makes a comprehensive quantification of young healthy balance performance possible. The results of such analyses could be used in work space design and in fall-prevention instructional materials, for both the home and work place. Young healthy performance represents “exemplar” performance and can also be used as a reference against which to compare the performance of aging and other clinical populations at risk for falling.

The vertebral strength and strain can be assessed in vitro by both using isolated vertebrae and sets of three adjacent vertebrae (the central one is loaded through the disks). Our goal was to elucidate if testing single-vertebra-specimens in the elastic regime provides different surface strains to three-vertebrae-segments. Twelve three-vertebrae sets were extracted from thoracolumbar human spines. To measure the principal strains, the central vertebra of each segment was prepared with eight strain-gauges. The sets were tested mechanically, allowing comparison of the surface strains between the two boundary conditions: first when the same vertebra was loaded through the disks (three-vertebrae-segment) and then with the endplates embedded in cement (single-vertebra). They were all subjected to four nondestructive tests (compression, traction, torsion clockwise, and counterclockwise). The magnitude of principal strains differed significantly between the two boundary conditions. For axial loading, the largest principal strains (along vertebral axis) were significantly higher when the same vertebra was tested isolated compared to the three-vertebrae-segment. Conversely, circumferential strains decreased significantly in the single vertebrae compared to the three-vertebrae-segment, with some variations exceeding 100% of the strain magnitude, including changes from tension to compression. For torsion, the differences between boundary conditions were smaller. This study shows that, in the elastic regime, when the vertebra is loaded through a cement pot, the surface strains differ from when it is loaded through the disks. Therefore, when single vertebrae are tested, surface strain should be taken with caution.

To assess how posture affects the transmission of mechanical energy up the spinal column during vibration, 18 porcine functional spinal units (FSUs) were exposed to a sinusoidal force (1500 ± 1200 N) at 5 Hz for 120 min in either a flexed, extended, or neutral posture. Force and FSU height were measured continuously throughout the collection. From these data, specimen height loss, dynamic stiffness, hysteresis, and parameters from a standard linear solid (SLS) model were determined and analyzed for differences between postures. Posture had an influence on all of these parameters. In extension, the FSU had higher dynamic stiffness values than when neutral or flexed (p < 0.0001). In flexion, the FSU had higher hysteresis than both an extended or neutral posture (p < 0.0001). Height loss was greatest in a flexed posture and smallest in an extended posture (p < 0.0001). In extension, the series spring element in the SLS model had a stiffness value higher than both flexed and neutral posture conditions, whereas the stiffness in the parallel spring was the same between extension and neutral (p < 0.01), both higher than in flexion. Viscosity coefficients were highest in extension compared to both flexed and neutral (p < 0.01). Based on these results, it was determined that posture had a significant influence in determining the mechanical properties of the spine when exposed to cyclic compressive loading.

Intervertebral disk (IVD) degeneration is a prevalent health problem that is highly linked to back pain. To understand the disease and tissue response to therapies, ex vivo whole IVD organ culture systems have recently been introduced. The goal of this work was to develop and validate the design of a whole spinal segment culturing system that loads the disk in complex loading similar to the in vivo condition, while preserving the adjacent endplates and vertebral bodies. The complex loading applied to the spinal segment (flexion–extension (FE), bilateral bending, and compression) was achieved with three pneumatic cylinders rigidly attached to a triangular loading platform. A culture container housed the spinal segment and was attached to the loading mechanism, which allowed for loading of the spinal segment. The dynamic bioreactor was able to achieve physiologic loading conditions with 100 N of applied compression and approximately 2–4 N · m of applied torque. The function of the bioreactor was validated through testing of bovine caudal IVDs with intact endplates and vertebral bodies that were isolated within 2 hrs of death and cultured for 14 days. The resulting IVD cell viability following 14 days of loading was much higher than unloaded control IVDs. The loading system accurately mimicked FE, bilateral bending, and compression motions seen during daily activities. The results indicate that this complex dynamic bioreactor may be appropriate for extended preclinical testing of vertebral-mounted spinal devices and therapies.

Studies on intervertebral disk (IVD) response to various loads and postures are essential to understand disk's mechanical functions and to suggest preventive and corrective actions in the workplace. The experimental and finite-element (FE) approaches are well-suited for these studies, but validating their findings is difficult, partly due to the lack of alternative methods. Analytical modeling could allow methodological triangulation and help validation of FE models. This paper presents an analytical method based on thin-shell, beam-on-elastic-foundation and composite materials theories to evaluate the stresses in the anulus fibrosus (AF) of an axisymmetric disk composed of multiple thin lamellae. Large deformations of the soft tissues are accounted for using an iterative method and the anisotropic material properties are derived from a published biaxial experiment. The results are compared to those obtained by FE modeling. The results demonstrate the capability of the analytical model to evaluate the stresses at any location of the simplified AF. It also demonstrates that anisotropy reduces stresses in the lamellae. This novel model is a preliminary step in developing valuable analytical models of IVDs, and represents a distinctive groundwork that is able to sustain future refinements. This paper suggests important features that may be included to improve model realism.

We developed a balance measurement tool (the balanced reach test (BRT)) to assess standing balance while reaching and pointing to a target moving in three-dimensional space according to a sum-of-sines function. We also developed a three-dimensional, 13-segment biomechanical model to analyze performance in this task. Using kinematic and ground reaction force (GRF) data from the BRT, we performed an inverse dynamics analysis to compute the forces and torques applied at each of the joints during the course of a 90 s test. We also performed spectral analyses of each joint's force activations. We found that the joints act in a different but highly coordinated manner to accomplish the tracking task—with individual joints responding congruently to different portions of the target disk's frequency spectrum. The test and the model also identified clear differences between a young healthy subject (YHS), an older high fall risk (HFR) subject before participating in a balance training intervention; and in the older subject's performance after training (which improved to the point that his performance approached that of the young subject). This is the first phase of an effort to model the balance control system with sufficient physiological detail and complexity to accurately simulate the multisegmental control of balance during functional reach across the spectra of aging, medical, and neurological conditions that affect performance. Such a model would provide insight into the function and interaction of the biomechanical and neurophysiological elements making up this system; and system adaptations to changes in these elements' performance and capabilities.