Pancheri FQ, Eng CM, Lieberman DE, Biewener AA, Dorfmann L.
A constitutive description of the anisotropic response of the fascia lata. J Mech Behav Biomed MaterJ Mech Behav Biomed MaterJ Mech Behav Biomed Mater. 2014;30 :306-23.
AbstractIn this paper we propose a constitutive model to analyze in-plane extension of goat fascia lata. We first perform a histological analysis of the fascia that shows a well-organized bi-layered arrangement of undulated collagen fascicles oriented along two well defined directions. To develop a model consistent with the tissue structure we identify the absolute and relative thickness of each layer and the orientation of the preferred directions. New data are presented showing the mechanical response in uniaxial and planar biaxial extension. The paper proposes a constitutive relation to describe the mechanical response. We provide a summary of the main ingredients of the nonlinear theory of elasticity and introduce a suitable strain-energy function to describe the anisotropic response of the fascia. We validate the model by showing good fit of the numerical results and the experimental data. Comments are included about differences and analogies between goat fascia lata and the human iliotibial band.
Lin HT, Ros IG, Biewener AA.
Through the eyes of a bird: modelling visually guided obstacle flight. J R Soc InterfaceJ R Soc InterfaceJ R Soc Interface. 2014;11 :20140239.
AbstractVarious flight navigation strategies for birds have been identified at the large spatial scales of migratory and homing behaviours. However, relatively little is known about close-range obstacle negotiation through cluttered environments. To examine obstacle flight guidance, we tracked pigeons (Columba livia) flying through an artificial forest of vertical poles. Interestingly, pigeons adjusted their flight path only approximately 1.5 m from the forest entry, suggesting a reactive mode of path planning. Combining flight trajectories with obstacle pole positions, we reconstructed the visual experience of the pigeons throughout obstacle flights. Assuming proportional-derivative control with a constant delay, we searched the relevant parameter space of steering gains and visuomotor delays that best explained the observed steering. We found that a pigeon's steering resembles proportional control driven by the error angle between the flight direction and the desired opening, or gap, between obstacles. Using this pigeon steering controller, we simulated obstacle flights and showed that pigeons do not simply steer to the nearest opening in the direction of flight or destination. Pigeons bias their flight direction towards larger visual gaps when making fast steering decisions. The proposed behavioural modelling method converts the obstacle avoidance behaviour into a (piecewise) target-aiming behaviour, which is better defined and understood. This study demonstrates how such an approach decomposes open-loop free-flight behaviours into components that can be independently evaluated.
Holt NC, Wakeling JM, Biewener AA.
The effect of fast and slow motor unit activation on whole-muscle mechanical performance: the size principle may not pose a mechanical paradox. Proc Biol SciProc Biol SciProc Biol Sci. 2014;281 :20140002.
AbstractThe output of skeletal muscle can be varied by selectively recruiting different motor units. However, our knowledge of muscle function is largely derived from muscle in which all motor units are activated. This discrepancy may limit our understanding of in vivo muscle function. Hence, this study aimed to characterize the mechanical properties of muscle with different motor unit activation. We determined the isometric properties and isotonic force-velocity relationship of rat plantaris muscles in situ with all of the muscle active, 30% of the muscle containing predominately slower motor units active or 20% of the muscle containing predominately faster motor units active. There was a significant effect of active motor unit type on isometric force rise time (p < 0.001) and the force-velocity relationship (p < 0.001). Surprisingly, force rise time was longer and maximum shortening velocity higher when all motor units were active than when either fast or slow motor units were selectively activated. We propose this is due to the greater relative effects of factors such as series compliance and muscle resistance to shortening during sub-maximal contractions. The findings presented here suggest that recruitment according to the size principle, where slow motor units are activated first and faster ones recruited as demand increases, may not pose a mechanical paradox, as has been previously suggested.
Lee DV, Isaacs MR, Higgins TE, Biewener AA, McGowan CP.
Scaling of the spring in the leg during bouncing gaits of mammals. Integr Comp BiolIntegr Comp BiolIntegr Comp Biol. 2014;54 :1099-108.
AbstractTrotting, bipedal running, and especially hopping have long been considered the principal bouncing gaits of legged animals. We use the radial-leg spring constant [Formula: see text] to quantify the stiffness of the physical leg during bouncing gaits. The radial-leg is modeled as an extensible strut between the hip and the ground and [Formula: see text] is determined from the force and deflection of this strut in each instance of stance. A Hookean spring is modeled in-series with a linear actuator and the stiffness of this spring [Formula: see text] is determined by minimizing the work of the actuator while reproducing the measured force-deflection dynamics of an individual leg during trotting or running, and of the paired legs during hopping. Prior studies have estimated leg stiffness using [Formula: see text], a metric that imagines a virtual-leg connected to the center of mass. While [Formula: see text] has been applied extensively in human and comparative biomechanics, we show that [Formula: see text] more accurately models the spring in the leg when actuation is allowed, as is the case in biological and robotic systems. Our allometric analysis of [Formula: see text] in the kangaroo rat, tammar wallaby, dog, goat, and human during hopping, trotting, or running show that [Formula: see text] scales as body mass to the two-third power, which is consistent with the predictions of dynamic similarity and with the scaling of [Formula: see text]. Hence, two-third scaling of locomotor spring constants among mammals is supported by both the radial-leg and virtual-leg models, yet the scaling of [Formula: see text] emerges from work-minimization in the radial-leg model instead of being a defacto result of the ratio of force to length used to compute [Formula: see text]. Another key distinction between the virtual-leg and radial-leg is that [Formula: see text] is substantially greater than [Formula: see text], as indicated by a 30-37% greater scaling coefficient for [Formula: see text]. We also show that the legs of goats are on average twice as stiff as those of dogs of the same mass and that goats increase the stiffness of their legs, in part, by more nearly aligning their distal limb-joints with the ground reaction force vector. This study is the first allometric analysis of leg spring constants in two decades. By means of an independent model, our findings reinforce the two-third scaling of spring constants with body mass, while showing that springs in-series with actuators are stiffer than those predicted by energy-conservative models of the leg.
Eng CM, Pancheri FQ, Lieberman DE, Biewener AA, Dorfmann L.
Directional differences in the biaxial material properties of fascia lata and the implications for fascia function. Ann Biomed EngAnn Biomed EngAnn Biomed Eng. 2014;42 :1224-37.
AbstractFascia is a highly organized collagenous tissue that is ubiquitous in the body, but whose function is not well understood. Because fascia has a sheet-like structure attaching to muscles and bones at multiple sites, it is exposed to different states of multi- or biaxial strain. In order to measure how biaxial strain affects fascia material behavior, planar biaxial tests with strain control were performed on longitudinal and transversely oriented samples of goat fascia lata (FL). Cruciform samples were cycled to multiple strain levels while the perpendicular direction was held at a constant strain. Structural differences among FL layers were examined using histology and SEM. Results show that FL stiffness, hysteresis, and strain energy density are greater in the longitudinal vs. transverse direction. Increased stiffness in the longitudinal layer is likely due to its greater thickness and greater average fibril diameter compared to the transverse layer(s). Perpendicular strain did not affect FL material behavior. Differential loading in the longitudinal vs. transverse directions may lead to structural changes, enhancing the ability of the longitudinal FL to transmit force, store energy, or stabilize the limb during locomotion. The relative compliance of the transverse fibers may allow expansion of underlying muscles when they contract.
Biewener AA, Wakeling JM, Lee SS, Arnold AS.
Validation of Hill-type muscle models in relation to neuromuscular recruitment and force-velocity properties: predicting patterns of in vivo muscle force. Integr Comp BiolIntegr Comp BiolIntegr Comp Biol. 2014;54 :1072-83.
AbstractWe review here the use and reliability of Hill-type muscle models to predict muscle performance under varying conditions, ranging from in situ production of isometric force to in vivo dynamics of muscle length change and force in response to activation. Muscle models are frequently used in musculoskeletal simulations of movement, particularly when applied to studies of human motor performance in which surgically implanted transducers have limited use. Musculoskeletal simulations of different animal species also are being developed to evaluate comparative and evolutionary aspects of locomotor performance. However, such models are rarely validated against direct measures of fascicle strain or recordings of muscle-tendon force. Historically, Hill-type models simplify properties of whole muscle by scaling salient properties of single fibers to whole muscles, typically accounting for a muscle's architecture and series elasticity. Activation of the model's single contractile element (assigned the properties of homogenous fibers) is also simplified and is often based on temporal features of myoelectric (EMG) activation recorded from the muscle. Comparison of standard one-element models with a novel two-element model and with in situ and in vivo measures of EMG, fascicle strain, and force recorded from the gastrocnemius muscles of goats shows that a two-element Hill-type model, which allows independent recruitment of slow and fast units, better predicts temporal patterns of in situ and in vivo force. Recruitment patterns of slow/fast units based on wavelet decomposition of EMG activity in frequency-time space are generally correlated with the intensity spectra of the EMG signals, the strain rates of the fascicles, and the muscle-tendon forces measured in vivo, with faster units linked to greater strain rates and to more rapid forces. Using direct measures of muscle performance to further test Hill-type models, whether traditional or more complex, remains critical for establishing their accuracy and essential for verifying their applicability to scientific and clinical studies of musculoskeletal function.