Craig P. McGowan

The fastest runner on artificial legs: different limbs, similar function?

The recent competitive successes of a bilateral, transtibial amputee sprint runner who races with modern running prostheses has triggered an international controversy regarding the relative function provided by his artificial limbs. Here, we conducted three tests of functional similarity between this amputee sprinter and competitive male runners with intact limbs: the metabolic cost of running, sprinting endurance, and running mechanics. Metabolic and mechanical data, respectively, were acquired via indirect calorimetry and ground reaction force measurements during constant-speed, level treadmill running. First, we found that the mean gross metabolic cost of transport of our amputee sprint subject (174.9 ml O(2)*kg(-1)*km(-1); speeds: 2.5-4.1 m/s) was only 3.8% lower than mean values for intact-limb elite distance runners and 6.7% lower than for subelite distance runners but 17% lower than for intact-limb 400-m specialists [210.6 (SD 13.2) ml O(2)*kg(-1)*km(-1)]. Second, the speeds that our amputee sprinter maintained for six all-out, constant-speed trials to failure (speeds: 6.6-10.8 m/s; durations: 2-90 s) were within 2.2 (SD 0.6)% of those predicted for intact-limb sprinters. Third, at sprinting speeds of 8.0, 9.0, and 10.0 m/s, our amputee subject had longer foot-ground contact times [+14.7 (SD 4.2)%], shorter aerial [-26.4 (SD 9.9)%] and swing times [-15.2 (SD 6.9)%], and lower stance-averaged vertical forces [-19.3 (SD 3.1)%] than intact-limb sprinters [top speeds = 10.8 vs. 10.8 (SD 0.6) m/s]. We conclude that running on modern, lower-limb sprinting prostheses appears to be physiologically similar but mechanically different from running with intact limbs.

Independent effects of weight and mass on plantar flexor activity during walking: implications for their contributions to body support and forward propulsion.

The ankle plantar flexor muscles, gastrocnemius (Gas) and soleus (Sol), have been shown to play important roles in providing body support and forward propulsion during human walking. However, there has been disagreement about the relative contributions of Gas and Sol to these functional tasks. In this study, using independent manipulations of body weight and body mass, we examined the relative contribution of the individual plantar flexors to support and propulsion. We hypothesized that Gas and Sol contribute to body support, whereas Sol is the primary contributor to forward trunk propulsion. We tested this hypothesis by measuring muscle activity while experimentally manipulating body weight and mass by 1) decreasing body weight using a weight support system, 2) increasing body mass alone using a combination of equal added trunk load and weight support, and 3) increasing trunk loads (increasing body weight and mass). The rationale for this study was that muscles that provide body support would be sensitive to changes in body weight, whereas muscles that provide forward propulsion would be sensitive to changes in body mass. Gas activity increased with added loads and decreased with weight support but showed only a small increase relative to control trials when mass alone was increased. Sol activity showed a similar increase with added loads and with added mass alone and decreased in early stance with weight support. Therefore, we accepted the hypothesis that Sol and Gas contribute to body support, whereas Sol is the primary contributor to forward trunk propulsion.

Dynamics of leg muscle function in tammar wallabies (M. eugenii) during level versus incline hopping

SUMMARY The goal of our study was to examine whether the in vivo force-length behavior, work and elastic energy savings of distal muscle-tendon units in the legs of tammar wallabies (Macropus eugenii) change during level versus incline hopping. To address this question, we obtained measurements of muscle activation (via electromyography), fascicle strain (via sonomicrometry) and muscle-tendon force (via tendon buckles) from the lateral gastrocnemius (LG) and plantaris (PL) muscles of tammar wallabies trained to hop on a level and an inclined (10°, 17.4% grade) treadmill at two speeds (3.3 m s-1 and 4.2 m s-1). Similar patterns of muscle activation, force and fascicle strain were observed under both level and incline conditions. This also corresponded to similar patterns of limb timing and movement (duty factor, limb contact time and hopping frequency). During both level and incline hopping, the LG and PL exhibited patterns of fascicle stretch and shortening that yielded low levels of net fascicle strain [LG: level, -1.0±4.6% (mean ± s.e.m.) vs incline, 0.6±4.5%; PL: level, 0.1±1.0% vs incline, 0.4±1.6%] and muscle work (LG: level, -8.4±8.4 J kg-1 muscle vs incline, -6.8±7.5 J kg-1 muscle; PL: level, -2.0±0.6 J kg-1 muscle vs incline, -1.4±0.7 J kg-1 muscle). Consequently, neither muscle significantly altered its contractile dynamics to do more work during incline hopping. Whereas electromyographic (EMG) phase, duration and intensity did not differ for the LG, the PL exhibited shorter but more intense periods of activation, together with reduced EMG phase (P<0.01), during incline versus level hopping. Our results indicate that design for spring-like tendon energy savings and economical muscle force generation is key for these two distal muscle-tendon units of the tammar wallaby, and the need to accommodate changes in work associated with level versus incline locomotion is achieved by more proximal muscles of the limb.

Running-specific prostheses limit ground-force during sprinting

Running-specific prostheses (RSP) emulate the spring-like behaviour of biological limbs during human running, but little research has examined the mechanical means by which amputees achieve top speeds. To better understand the biomechanical effects of RSP during sprinting, we measured ground reaction forces (GRF) and stride kinematics of elite unilateral trans-tibial amputee sprinters across a range of speeds including top speed. Unilateral amputees are ideal subjects because each amputees affected leg (AL) can be compared with their unaffected leg (UL). We found that stance average vertical GRF were approximately 9 per cent less for the AL compared with the UL across a range of speeds including top speed (p < 0.0001). In contrast, leg swing times were not significantly different between legs at any speed (p = 0.32). Additionally, AL and UL leg swing times were similar to those reported for non-amputee sprinters. We infer that RSP impair force generation and thus probably limit top speed. Some elite unilateral trans-tibial amputee sprinters appear to have learned or trained to compensate for AL force impairment by swinging both legs rapidly.

Modulation of leg muscle function in response to altered demand for body support and forward propulsion during walking

A number of studies have examined the functional roles of individual muscles during normal walking, but few studies have examined which are the primary muscles that respond to changes in external mechanical demand. Here we use a novel combination of experimental perturbations and forward dynamics simulations to determine how muscle mechanical output and contributions to body support and forward propulsion are modulated in response to independent manipulations of body weight and body mass during walking. Experimentally altered weight and/or mass were produced by combinations of added trunk loads and body weight support. Simulations of the same experimental conditions were used to determine muscle contributions to the vertical ground reaction force impulse (body support) and positive horizontal trunk work (forward propulsion). Contributions to the vertical impulse by the soleus, vastii and gluteus maximus increased (decreased) in response to increases (decreases) in body weight; whereas only the soleus increased horizontal work output in response to increased body mass. In addition, soleus had the greatest absolute contribution to both vertical impulse and horizontal trunk work, indicating that it not only provides the largest contribution to both body support and forward propulsion, but the soleus is also the primary mechanism to modulate the mechanical output of the leg in response to increased (decreased) need for body support and forward propulsion. The data also showed that a muscles contribution to a specific task is likely not independent of its contribution to other tasks (e.g., body support vs. forward propulsion).

Muscle Contributions to Whole-Body Sagittal Plane Angular Momentum during Walking

Walking is a complex dynamic task that requires the regulation of whole-body angular momentum to maintain dynamic balance while performing walking subtasks such as propelling the body forward and accelerating the leg into swing. In human walking, the primary mechanism to regulate angular momentum is muscle force generation. Muscles accelerate body segments and generate ground reaction forces that alter angular momentum about the bodys center-of-mass to restore and maintain dynamic stability. In addition, gravity contributes to whole-body angular momentum through its contribution to the ground reaction forces. The purpose of this study was to generate a muscle-actuated forward dynamics simulation of normal walking to quantify how individual muscles and gravity contribute to whole-body angular momentum in the sagittal plane. In early stance, the uniarticular hip and knee extensors (GMAX and VAS), biarticular hamstrings (HAM) and ankle dorsiflexors (TA) generated backward angular momentum while the ankle plantar flexors (SOL and GAS) generated forward momentum. In late stance, SOL and GAS were the primary contributors and generated angular momentum in opposite directions. SOL generated primarily forward angular momentum while GAS generated backward angular momentum. The difference between muscles was due to their relative contributions to the horizontal and vertical ground reaction forces. Gravity contributed to the bodys angular momentum in early stance and to a lesser extent in late stance, which was counteracted primarily by the plantar flexors. These results may provide insight into balance and movement disorders and provide a basis for developing locomotor therapies that target specific muscle groups.

Dynamic pressure maps for wings and tails of pigeons in slow, flapping flight, and their energetic implications

SUMMARY Differential pressure measurements offer a new approach for studying the aerodynamics of bird flight. Measurements from differential pressure sensors are combined to form a dynamic pressure map for eight sites along and across the wings, and for two sites across the tail, of pigeons flying between two perches. The confounding influence of acceleration on the pressure signals is shown to be small for both wings and tail. The mean differential pressure for the tail during steady, level flight was 25.6 Pa, which, given an angle of attack for the tail of 47.6°, suggests the tail contributes 7.91% of the force required for weight support, and requires a muscle-mass specific power of 19.3 W kg-1 for flight to overcome its drag at 4.46 m s-1. Differential pressures during downstroke increase along the wing length, to 300-400 Pa during take-off and landing for distal sites. Taking the signals obtained from five sensors sited along the wing at feather bases as representative of the mean pressure for five spanwise elements at each point in time, and assuming aerodynamic forces act within the x-z plane (i.e. no forces in the direction of travel) and perpendicular to the wing during downstroke, we calculate that 74.5% of the force required to support weight was provided by the wings, and that the aerodynamic muscle-mass specific power required to flap the wings was 272.7 W kg-1.

Obesity does not increase external mechanical work per kilogram body mass during walking

Walking is the most common type of physical activity prescribed for the treatment of obesity. The net metabolic rate during level walking (W/kg) is approximately 10% greater in obese vs. normal weight adults. External mechanical work (W(ext)) is one of the primary determinants of the metabolic cost of walking, but the effects of obesity on W(ext) have not been clearly established. The purpose of this study was to compare W(ext) between obese and normal weight adults across a range of walking speeds. We hypothesized that W(ext) (J/step) would be greater in obese adults but W(ext) normalized to body mass would be similar in obese and normal weight adults. We collected right leg three-dimensional ground reaction forces (GRF) while twenty adults (10 obese, BMI=35.6 kg/m(2) and 10 normal weight, BMI=22.1 kg/m(2)) walked on a level, dual-belt force measuring treadmill at six speeds (0.50-1.75 m/s). We used the individual limb method (ILM) to calculate external work done on the center of mass. Absolute W(ext) (J/step) was greater in obese vs. normal weight adults at each walking speed, but relative W(ext) (J/step/kg) was similar between the groups. Step frequencies were not different. These results suggest that W(ext) is not responsible for the greater metabolic cost of walking (W/kg) in moderately obese adults.

Leg stiffness of sprinters using running-specific prostheses

Running-specific prostheses (RSF) are designed to replicate the spring-like nature of biological legs (bioL) during running. However, it is not clear how these devices affect whole leg stiffness characteristics or running dynamics over a range of speeds. We used a simple spring–mass model to examine running mechanics across a range of speeds, in unilateral and bilateral transtibial amputees and performance-matched controls. We found significant differences between the affected leg (AL) of unilateral amputees and both ALs of bilateral amputees compared with the bioL of non-amputees for nearly every variable measured. Leg stiffness remained constant or increased with speed in bioL, but decreased with speed in legs with RSPs. The decrease in leg stiffness in legs with RSPs was mainly owing to a combination of lower peak ground reaction forces and increased leg compression with increasing speeds. Leg stiffness is an important parameter affecting contact time and the force exerted on the ground. It is likely that the fixed stiffness of the prosthesis coupled with differences in the limb posture required to run with the prosthesis limits the ability to modulate whole leg stiffness and the ability to apply high vertical ground reaction forces during sprinting.

Forward dynamics simulations provide insight into muscle mechanical work during human locomotion.

Complex musculoskeletal models and computer simulations can provide critical insight into muscle mechanical work output during locomotion. Simulations provide both a consistent mechanical solution that can be interrogated at multiple levels (muscle fiber, musculotendon, net joint moment, and whole-body work) and an ideal framework to identify limitations with different estimates of muscle work and the resulting implications for metabolic cost and efficiency.