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Biomedical subjects

John E A Bertram

Publications and source records attributed to John E A Bertram.

11 recordsLinked to original sources

Collateral ligaments of the distal sesamoid bone in the digit of Equus: re-evaluating midstance function.

The distal forelimb of the horse has a complex array of ligaments that play a critical role in determining function of the digit and are often associated with the initiation of foot pathologies. The collateral ligaments of the distal sesamoid bone (CLDS) play an important role in digit stabilization near the end of foot contact and there is also limited evidence to suggest that the CLDS stabilize the proximal interphalangeal joint (PIPJ) during weight bearing. By virtue of their anatomical attachments where the ligaments pass dorsal to the axis of rotation of the PIPJ, it is reasonable to assume that the CLDS prevent flexion of the PIPJ during weight bearing or midstance in a moving horse. To test this functional hypothesis, forelimb specimens from three mixed-breed horses were loaded in compression in a materials testing frame. Limb loading was applied with the CLDS intact and following transection. Average PIPJ angle and metacarpophalangeal joint (MCPJ) angle at maximum load (approximately 3000 N) were calculated from angular changes of proximal and middle phalanges and the third metacarpal, which were compared between intact and transected trials. PIPJ angles were found to be the same (175 degrees) at maximum load for intact and transected trials. The proximal and middle phalanges rotated together remaining aligned, regardless of the CLDS condition. Contrary to expectation, however, the combined proximal and middle phalanges unit rotates less relative to the third metacarpal under load after transection, indicating less digit extension at the metacarpophalangeal (fetlock) joint without the influence of CLDS. Since the mechanical properties of the fetlock joint are unchanged by CLDS transection, observed proximal and middle phalanx motion is dependent on increased rotation of the distal phalanx after transection. The original hypothesis was not supported and the results suggest that at midstance the CLDS function primarily to stabilize the articulation of the middle phalanx about the distal phalanx to limit distal interphalangeal joint extension during weight bearing. Establishing the functional role of the CLDS may help to better understand the biomechanical consequences of ligament injuries and diseases of the pastern.

Animals↗

Constrained optimization in human running.

Walking humans spontaneously select different speed, frequency and step length combinations, depending on which of these three parameters is specified. This behavior can be explained by constrained optimization of cost of transport (metabolic cost/distance) where cost of transport is seen as the main component of an underlying objective function that is minimized within the limitations of specified constraints. It is then of interest to ask whether or not such results are specific to walking only, or indicate a more general feature of locomotion control. The current study examines running gait selection within the framework of constrained optimization by comparing self-selected running gaits to the gaits predicted by constrained optimization of a cost surface constructed from cost data available in the literature. Normalizing speed and frequency values in the behavioral data by preferred speed and frequency reduced inter-subject variability and made group behavioral trends more visible. Although actual behavior did not coincide exactly with running cost optimization, self-selected gait and predictions from the general human cost surface did agree to within the 95% confidence interval and the region of minimal cost+0.005 ml O2 kg(-1) m(-1). This was similar to the level of agreement between actual and predicted behavior observed in walking. Thus, there seems to be substantial evidence to suggest that (i) selection of gait parameters in running can largely be predicted using constrained optimization, and (ii) general cost surfaces can be constructed using metabolic data from one group that will largely predict the behavior of other groups.

Adult↗

A collisional model of the energetic cost of support work qualitatively explains leg sequencing in walking and galloping, pseudo-elastic leg behavior in running and the walk-to-run transition.

Terrestrial legged locomotion requires repeated support forces to redirect the body's vertical velocity component from down to up. We assume that the redirection is accomplished by impulsive leg forces that cause small-angle glancing collisions of a point-mass model of the animal. We estimate the energetic costs of these collisions by assuming a metabolic cost proportional to positive muscle work involved in generating the impulses. The cost of bipedal running estimated from this collisional model becomes less than that of walking at a Froude number (v2/gl) of about 0.7. Two strategies to reduce locomotion costs associated with the motion redirection are: (1) having legs simulate purely elastic springs, as is observed in human running; and (2) sequencing the leg forces during the redirection phase; examples of this sequencing are the ba-da-dump pattern of a horse gallop and having push-off followed by heel-strike in human walking.

Animals↗

Constrained optimization in human walking: cost minimization and gait plasticity.

As walking speed increases, consistent relationships emerge between the three determinant parameters of walking, speed, step frequency and step length. However, when step length or step frequency are predetermined rather than speed, different relationships are spontaneously selected. This result is expected if walking parameters are selected to optimize to an underlying objective function, known as the constrained optimization hypothesis. The most likely candidate for the objective function is metabolic cost per distance traveled, where the hypothesis predicts that the subject will minimize the cost of travel under a given gait constraint even if this requires an unusual step length and frequency combination. In the current study this is tested directly by measuring the walking behavior of subjects constrained systematically to determined speeds, step frequencies or step lengths and comparing behavior to predictions derived directly from minimization of measured metabolic cost. A metabolic cost surface in speed-frequency space is derived from metabolic rate for 10 subjects walking at 49 speed-frequency conditions. Optimization is predicted from the iso-energetic cost contours derived from this surface. Substantial congruence is found between the predicted and observed behavior using the cost of walking per unit distance. Although minimization of cost per distance appears to dominate walking control, certain notable differences from predicted behavior suggest that other factors must also be considered. The results of these studies provide a new perspective on the integration of walking cost with neuromuscular control, and provide a novel approach to the investigation of the control features involved in gait parameter selection.

Adult↗

Testing the hindlimb-strength hypothesis: non-aerial locomotion by Chiroptera is not constrained by the dimensions of the femur or tibia.

In the evolution of flight bats appear to have suffered a trade-off; they have become poor crawlers relative to terrestrial mammals. Capable walking does occur in a few disparate taxa, including the vampire bats, but the vast majority of bats are able only to shuffle awkwardly along the ground, and the morphological bases of differences in crawling ability are not currently understood. One widely cited hypothesis suggests that the femora of most bats are too weak to withstand the compressive forces that occur during terrestrial locomotion, and that the vampire bats can walk because they possess more robust hindlimb skeletons. We tested a prediction of the hindlimb-strength hypothesis: that during locomotion, the forces produced by the hindlimbs of vampire bats should be larger than those produced by the legs of poorly crawling bats. Using force plates we compared the hindlimb forces produced by two species of vampire bats that walk well, Desmodus rotundus (N=8) and Diaemus youngi (N=2), to the hindlimb forces produced during over-ground shuffling by a similarly sized bat that is a poor walker (Pteronotus parnellii; N=6). Peak hindlimb forces produced by P. parnellii were larger (ANOVA; P<0.05; N=65) and more variable (93.5+/-36.6% body weight, mean +/- s.d.) than those of D. rotundus (69.3+/-8.1%) or D. youngi (75.0+/-6.2%). Interestingly, the vertical components of peak force were equivalent among species (P>0.6), indicating similar roles for support of body weight by the hindlimbs in the three species. We also used a simple engineering model of bending stress to evaluate the support capabilities of the hindlimb skeleton from the dimensions of 113 museum specimens in 50 species. We found that the hindlimb bones of vampires are not built to withstand larger forces than those of species that crawl poorly. Our results show that the legs of poorly crawling bats should be able to withstand the forces produced during coordinated crawling of the type used by the agile vampires, and this indicates that some mechanism other than hindlimb bone thickness, such as myology of the pectoral girdle, limits the ability of most bats to crawl.

Analysis of Variance↗

New perspectives on brachiation mechanics.

This review is designed to evaluate and interpret studies relevant to the locomotory mode known as brachiation, particularly as performed by the Hylobatid apes: the gibbon and siamang species. The older literature and its conclusions are evaluated against recent work performed by the author and other research groups working on brachiation models, either computer simulations or physical robots. The gibbon displays two types of brachiation: continuous contact, analogous to walking, and ricochetal, analogous to running. Both brachiation gaits display substantial pendular exchange between kinetic and potential energy. However, the fundamental feature of either of these gaits is the minimization of collisional energy loss. Collisional energy loss due to discontinuities in the trajectory of the center of mass is emerging as key in understanding locomotion using limbs in any terrestrial environment. The insight gained from this perspective applied to gibbon locomotion demonstrates that this is a critical factor in understanding many of the maneuvers employed by these animals, and can provide novel new interpretations of the morphological specializations that characterize the group. It is observed that these animals could brachiate using either totally active (muscle powered) or totally passive (nonmuscular) mechanisms. The active option would be metabolically costly, but provides substantial motion plasticity, while the passive option has the potential for profound economy, but does not allow a means to effectively contend with the inconsistencies present in the animal's natural environment. The conclusion is that the body form of brachiators and the locomotion behaviors they exhibit are a compromise between these two extremes, and these features of the gibbon's biology can only be understood by recognizing the role of collisional energy loss and evaluating both passive and active motion options together.

Animals↗

Gait transition cost in humans.

The energetics of locomotion depend largely on speed, gait and body size. Gait selection for a given speed appears partly, but perhaps not wholly, related to metabolic cost. One cost normally omitted from considerations of locomotion efficiency is the metabolic cost of the transition between gaits. We present the first direct assessment of the metabolic cost for the walk-run/run-walk transition in humans. The average increase in metabolic cost for a step involving a transition is 1.75 times that of a mean non-transition step at a speed where metabolic power requirements are identical for walking and running. Despite this substantial increase in cost for the transition step, the metabolic cost of gait transition is unlikely to have a strong bearing on the process of gait selection as the cost of using a metabolically inappropriate gait, even for only a few steps, will dominate.

Adult↗

Mechanisms of force and power production in unsteady ricochetal brachiation.

Brachiators travel by swinging beneath handholds, and it is not obvious how these animals manage to accelerate and decelerate in a horizontal direction, especially when moving rapidly. Most previous analyses focused on brachiation in highly constrained laboratory conditions that induced steady-state locomotion. Emerging understanding of brachiation suggests that much of gibbon locomotory behavior and morphology must be considered within the context of the complexities of the natural environment: the forest canopy is three-dimensional, with high variation in handhold availability and properties. The goal of this paper is to quantify the active mechanisms by which gibbons can dynamically control their velocity. Force production and kinematics were analyzed from a white-handed gibbon Hylabates lar during ricochetal brachiation. Both the mechanisms of force production and power input may be inferred for accelerating and decelerating brachiation by combining force data with kinematics. Examples of steady-state, accelerating, and decelerating ricochetal brachiation are highlighted. Gibbons are able to produce net horizontal impulses by releasing early (resulting in a loss of potential energy, but an accelerating horizontal impulse) or delaying release (associated with an increase in potential energy, and a decelerating horizontal impulse). Torque about the shoulder, leg-lifting (or dropping), and elbow flexing (or straightening) are discussed as potential mechanisms for controlling energy within the brachiating system. Of these possibilities, leg-lifting and arm-flexing were observed as mechanisms of adding mechanical energy. Net energy loss, and substantial torques about the shoulder, were not observed.

Acceleration↗

Understanding brachiation: insight from a collisional perspective.

Gibbons are able to brachiate effectively through the forest canopy with a suspended swinging motion via contact with handholds. The swing phase is unlikely to be a cause of significant energy loss as pendulums are able to oscillate with only gradual mechanical energy dissipation. We consider the energetics associated with the transition of either a swing (during continuous-contact brachiation) or a ballistic flight (ricochetal brachiation) to a subsequent swing. In both styles of brachiation, kinematic data suggest that a gibbon overshoots the path that would allow a smooth transition into the swing phase. The sudden change in velocity due to such an overshoot is associated with a collision. Assuming neither the handhold nor the gibbon stores elastic strain energy, the energetic consequences of such overshoots can be calculated. We suggest two reasons for overshooting smooth transition trajectories: in the case of continuous contact brachiation, excess mechanical energy can be maintained with a high amplitude swing, and an overshoot during ricochetal brachiation produces a safety margin. The degree of energy loss with the transition to the swing phase is dependent both on the alignment of the pre- and post-transition paths, and on the kinetic energy at that instant. Active mechanisms reduce the effects of overshoots in both brachiation gaits. During continuous-contact brachiation, the path of the centre of mass can be controlled actively by flexion both of the trailing arm and the legs. During ricochetal brachiation, the length between the hand and the centre of mass (determining the subsequent swing path) can be controlled throughout the flight phase with leg flexion/extension. In addition, the elongated arms characteristic of gibbons improves the geometry of a collision for a given overshoot, and so may be viewed as a morphological adaptation reducing the energetic losses caused by overshooting for safety.

Adaptation, Physiological↗

Insulin-like growth factor-I improves cellular and molecular aspects of healing in a collagenase-induced model of flexor tendinitis.

Flexor tendinitis is a common and debilitating injury of elite and recreational athletes. Healing may be improved through intratendinous injection of insulin-like growth factor-I (IGF-I), which has been shown in vitro to stimulate mitogenesis and enhance tendon matrix production. This study investigated the effects of intratendinous injection of IGF-I on tendon healing in an equine model of flexor tendinitis. Collagenase-induced lesions were created in the tensile region of theflexor digitorum superficialis tendon of both forelimbs of eight horses. Treated tendons were injected with 2 microg rhlGF-I intralesionally every other day for 10 injections, while controls received 0.9% NaCl. Tendon fiber deposition and organization were evaluated serially using ultrasonography throughout the 8 week trial period. Following euthanasia, the tendons were harvested and DNA, hydroxyproline, and glycosaminoglycan content determined, mechanical strength and stiffness evaluated, gene expression and spatial arrangement of collagen types I and III assessed by northern blot and in situ hybridization, and tendon fiber architecture assessed by polarized light microscopy. Local soft tissue swelling was reduced in the IGF-I treated limbs. Similarly, lesion size in IGF-I treated tendons was smaller 3 and 4 weeks after initiation of treatment. Cell proliferation and collagen content of the IGF-I treated tendons were increased compared to controls. Mechanically, IGF-I treated tendons showed a trend toward increased stiffness compared to saline treated controls. Considered together with the decreased soft tissue swelling and improved sonographic healing, these data support the potential use of intralesional IGF-I for treatment of debilitating tendon injuries.

Animals↗

Hypothesis testing as a laboratory exercise: a simple analysis of human walking, with a physiological surprise.

This paper describes a laboratory exercise designed to provide students with experience testing a hypothesis by systematically isolating and controlling determinant variables. The study involves an analysis of walking and is performed by the students on a subject from within their lab group. The study requires use of a motorized treadmill, tape measure, stop watch, metronome, personal cassette player, and calculator. The exercise is designed to include factors that the students are familiar with, so they can focus on the isolation of variables without being confused about the process they are investigating. However, the exercise will not turn out as the students anticipate, meaning they will be forced to reevaluate the assumptions that formed the basis of their original hypothesis. This exercise is designed for a college-level course in exercise science, physiology, or biology but could easily be managed by a high school honors class with appropriate guidance.

Adult↗