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Biomedical subjects

A A Biewener

Publications and source records attributed to A A Biewener.

At least 19 recordsLinked to original sources
PubMed2006

Effects of load carrying on metabolic cost and hindlimb muscle dynamics in guinea fowl (Numida meleagris).

The goal of this study was to test whether the contractile patterns of two major hindlimb extensors of guinea fowl are altered by load-carrying exercise. We hypothesized that changes in contractile pattern, specifically a decrease in muscle shortening velocity or enhanced stretch activation, would result in a reduction in locomotor energy cost relative to the load carried. We also anticipated that changes in kinematics would reflect underlying changes in muscle strain. Oxygen consumption, muscle activation intensity, and fascicle strain rate were measured over a range of speeds while animals ran unloaded vs. when they carried a trunk load equal to 22% of their body mass. Our results showed that loading produced no significant (P > 0.05) changes in kinematic patterns at any speed. In vivo muscle contractile strain patterns in the iliotibialis lateralis pars postacetabularis and the medial head of the gastrocnemius showed a significant increase in active stretch early in stance (P < 0.01), but muscle fascicle shortening velocity was not significantly affected by load carrying. The rate of oxygen consumption increased by 17% (P < 0.01) during loaded conditions, equivalent to 77% of the relative increase in mass. Additionally, relative increases in EMG intensity (quantified as mean spike amplitude) indicated less than proportional recruitment, consistent with force enhancement via stretch activation, in the proximal iliotibialis lateralis pars postacetabularis; however, a greater than proportional increase in the medial gastrocnemius was observed. As a result, when averaged for the two muscles, EMG intensity increased in direct proportion to the fractional increase in load carried.

Animals↗
PubMed2006

Differential design for hopping in two species of wallabies.

Hindlimb musculoskeletal anatomy and steady speed over ground hopping mechanics were compared in two species of macropod marsupials, tammar wallabies and yellow-footed rock wallabies (YFRW). These two species are relatively closely related and are of similar size and general body plan, yet they inhabit different environments with presumably different musculoskeletal demands. Tammar wallabies live in relatively flat, open habitat whereas yellow-footed rock wallabies inhabit steep cliff faces. The goal of this study was to explore musculoskeletal differences between tammar wallabies and yellow-footed rock wallabies and determine how these differences influence each species' hopping mechanics. We found the cross-sectional area of the combined ankle extensor tendons of yellow-footed rock wallabies was 13% greater than that of tammar wallabies. Both species experienced similar ankle joint moments during steady-speed hopping, however due to a lower mechanical advantage at this joint, tammar wallabies produced 26% more muscle force. Thus, during moderate speed hopping, yellow-footed rock wallabies operated with 38% higher tendon safety factors, while tammar wallabies were able to store 73% more elastic strain energy (2.18 J per leg vs. 1.26 J in YFRW). This likely reflects the differing demands of the environments inhabited by these two species, where selection for non-steady locomotor performance in rocky terrain likely requires trade-offs in locomotor economy.

Animals↗
PubMed2005

Joint work and power associated with acceleration and deceleration in tammar wallabies (Macropus eugenii).

Measurements of joint work and power were determined using inverse dynamics analysis based on ground reaction force and high-speed video recordings of tammar wallabies as they decelerated and accelerated while hopping over a force platform on level ground. Measurements were obtained over a range of accelerations ranging from -6 m s(-2) to 8 m s(-2). The goal of our study was to determine which joints are used to modulate mechanical power when tammar wallabies change speed. From these measurements, we also sought to determine which hind limb muscle groups are the most important for producing changes in mechanical work. Because our previous in vivo analyses of wallaby distal muscle function indicated that these muscle-tendon units favor elastic energy savings and perform little work during steady level and incline hopping, we hypothesized that proximal muscle groups operating at the hip and knee joint are most important for the modulation of mechanical work and power. Of the four hind limb joints examined, the ankle joint had the greatest influence on the total limb work, accounting for 89% of the variation observed with changing speed. The hip and metatarsophalageal (MP) joints also contributed to modulating whole limb work, but to a lesser degree than the ankle, accounting for 28% (energy production) and -24% (energy absorption) of the change in whole limb work versus acceleration, respectively. In contrast, the work produced at the knee joint was independent of acceleration. Based on the results of our previous in vivo studies and given that the magnitude of power produced at the ankle exceeds that which these muscles alone could produce, we conclude that the majority of power produced at the ankle joint is likely transferred from the hip and knee joints via proximal bi-articular muscles, operating in tandem with bi-articular ankle extensors, to power changes in hopping speed of tammar wallabies. Additionally, over the observed range of performance, peak joint moments at the ankle (and resulting tendon strains) did not increase significantly with acceleration, indicating that having thin tendons favoring elastic energy storage does not necessarily limit a tammar wallaby's ability to accelerate or decelerate.

Acceleration↗
PubMed2005

The mechanics of jumping versus steady hopping in yellow-footed rock wallabies.

The goal of our study was to explore the mechanical power requirements associated with jumping in yellow-footed rock wallabies and to determine how these requirements are achieved relative to steady-speed hopping mechanics. Whole body power output and limb mechanics were measured in yellow-footed rock wallabies during steady-speed hopping and moving jumps up to a landing ledge 1.0 m high (approximately 3 times the animals' hip height). High-speed video recordings and ground reaction force measurements from a runway-mounted force platform were used to calculate whole body power output and to construct a limb stiffness model to determine whole limb mechanics. The combined mass of the hind limb extensor muscles was used to estimate muscle mass-specific power output. Previous work suggested that a musculoskeletal design that favors elastic energy recovery, like that found in tammar wallabies and kangaroos, may impose constraints on mechanical power generation. Yet rock wallabies regularly make large jumps while maneuvering through their environment. As jumping often requires high power, we hypothesized that yellow-footed rock wallabies would be able to generate substantial amounts of mechanical power. This was confirmed, as we found net extensor muscle power outputs averaged 155 W kg(-1) during steady hopping and 495 W kg(-1) during jumping. The highest net power measured reached nearly 640 W kg(-1). As these values exceed the maximum power-producing capability of vertebrate skeletal muscle, we suggest that back, trunk and tail musculature likely play a substantial role in contributing power during jumping. Inclusion of this musculature yields a maximum power output estimate of 452 W kg(-1) muscle. Similar to human high-jumpers, rock wallabies use a moderate approach speed and relatively shallow leg angle of attack (45-55 degrees) during jumps. Additionally, initial leg stiffness increases nearly twofold from steady hopping to jumping, facilitating the transfer of horizontal kinetic energy into vertical kinetic energy. Time of contact is maintained during jumping by a substantial extension of the leg, which keeps the foot in contact with the ground.

Analysis of Variance↗
PubMed2004

Walking and running in the red-legged running frog, Kassina maculata.

Although most frog species are specialized for jumping or swimming, Kassina maculata (red-legged running frog) primarily uses a third type of locomotion during which the hindlimbs alternate. In the present study, we examined Kassina's distinct locomotory mode to determine whether these frogs walk or run and how their gait may change with speed. We used multiple methods to distinguish between terrestrial gaits: the existence or absence of an aerial phase, duty factor, relative footfall patterns and the mechanics of the animal's center of mass (COM). To measure kinematic and kinetic variables, we recorded digital video as the animals moved over a miniature force platform (N=12 individuals). With respect to footfall patterns, the frogs used a single gait and walked at all speeds examined. Duty factor always exceeded 0.59. Based on COM mechanics, however, the frogs used both walking and running gaits. At slower speeds, the fluctuations in the horizontal kinetic energy (E(k)) and gravitational potential energy (E(p)) of the COM were largely out of phase, indicating a vaulting or walking gait. In most of the trials, Kassina used a combined gait at intermediate speeds, unlike cursorial animals with distinct gait transitions. This combined gait, much like a mammalian gallop, exhibited the mechanics of both vaulting and bouncing gaits. At faster speeds, the E(k) and E(p) of Kassina's COM were more in phase, indicating the use of a bouncing or running gait. Depending on the definition used to distinguish between walking and running, Kassina either only used a walking gait at all speeds or used a walking gait at slower speeds but then switched to a running gait as speed increased.

Animals↗
PubMed2004

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

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 degrees, 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.

Animals↗
PubMed2003

In vivo and in vitro heterogeneity of segment length changes in the semimembranosus muscle of the toad.

Many studies examine sarcomere dynamics in single fibres or length-tension dynamics in whole muscles in vivo or in vitro, but few studies link the various levels of organisation. To relate data addressing in vitro muscle segment behaviour with in vivo whole muscle behaviour during locomotion, we measured in vivo strain patterns of muscle segments using three sonomicrometry crystals implanted along a fascicle of the semimembranosus muscle in the American toad (Bufo americanus; n = 6) during hopping. The centre crystal emitted an ultrasonic signal, while the outer crystals received the signal allowing the instantaneous measurement of lengths from two adjacent muscle segments. On the first day, we recorded from the central and distal segments. On the second day of recordings, the most distal crystal was moved to a proximal position to record from a proximal segment and the same central segment. When the toads hopped a distance of two body lengths, the proximal and central segments strained -15.1 +/- 6.1 and -14.0 +/- 4.9 % (i.e. shortening), respectively. Strain of the distal segment, however, was significantly lower and more variable in pattern, often lengthening before shortening during a hop. From rest length, the distal segment initially lengthened by 2.6 +/- 2.0 % before shortening by 6.5 +/- 3.2 % at the same hop distance. Under in vitro conditions, the central segment always shortened more than the distal segment, except when passively cycled, during which the segments strained similarly. When the whole muscle was cycled sinusoidally and stimulated phasically in vitro, the two adjacent segments strained in opposite directions over much (up to 34 %) of the cycle. These differences in strain amplitude and direction imply that two adjacent segments can not only produce and/or absorb varying amounts of mechanical energy, but can also operate on different regions of their force-length and force-velocity relationships when activated by the same neural signal. Understanding regional differences in contractile dynamics within muscles is therefore important to linking our understanding of sarcomere behaviour with whole muscle behaviour during locomotion.

Animals↗
PubMed2003

Comparative power curves in bird flight.

The relationship between mechanical power output and forward velocity in bird flight is controversial, bearing on the comparative physiology and ecology of locomotion. Applied to flying birds, aerodynamic theory predicts that mechanical power should vary as a function of forward velocity in a U-shaped curve. The only empirical test of this theory, using the black-billed magpie (Pica pica), suggests that the mechanical power curve is relatively flat over intermediate velocities. Here, by integrating in vivo measurements of pectoralis force and length change with quasi-steady aerodynamic models developed using data on wing and body movement, we present mechanical power curves for cockatiels (Nymphicus hollandicus) and ringed turtle-doves (Streptopelia risoria). In contrast to the curve reported for magpies, the power curve for cockatiels is acutely concave, whereas that for doves is intermediate in shape and shows higher mass-specific power output at most speeds. We also find that wing-beat frequency and mechanical power output do not necessarily share minima in flying birds. Thus, aspects of morphology, wing kinematics and overall style of flight can greatly affect the magnitude and shape of a species' power curve.

Animals↗
PubMed2001

Dynamics of mallard (Anas platyrynchos) gastrocnemius function during swimming versus terrestrial locomotion.

This study investigates how the contractile function of a muscle may be modulated to accommodate changes in locomotor mode and differences in the physical environment. In vivo recordings of lateral gastrocnemius (LG) activation, force development (measured using tendon buckle transducers) and length change (measured using sonomicrometry) were obtained from mallard ducks (Anas platyrhynchos) as they swam at steady speeds in a water tank and walked or ran on land. LG force recordings were compared with combined lateral and medial gastrocnemius (MG) muscle-tendon force recordings obtained from the contralateral limb, allowing force development by the MG to be estimated relative to that of the LG. Although similar stresses were calculated to act in the LG and MG muscles during terrestrial locomotion (126 and 115 kPa, respectively), stresses were considerably greater in the LG compared with the MG during swimming (62 versus 34 kPa, respectively). During both steady swimming and terrestrial locomotion, the LG developed force while shortening over a considerable range of its length (swimming 23.6 % versus terrestrial 37.4 %). Activation of the muscle occurred near the end of passive lengthening during the recovery stroke, just prior to muscle shortening. As a result, the muscle generated broad positive work loops during both locomotor modes. LG work during swimming (4.8 J x kg(-1)) averaged 37 % of the work performed during terrestrial locomotion (13.1 J x kg(-1)), consistent with the twofold greater force and 58 % greater strain of the muscle during walking and running. Because limb cycle frequency was similar for the two locomotor modes (swimming 2.65 versus terrestrial 2.61 Hz), differences in power output (swimming 12.6 W x kg(-1 )versus terrestrial 32.4 W x kg(-1)) largely reflected difference in work per cycle. Tendon elastic energy savings was a small fraction (<5 %) of the work performed by the muscle, consistent with a fiber-tendon design of these two muscles that favors muscle work to produce limb movement with little tendon strain. These results are consistent with a higher cost of terrestrial locomotion in ducks compared with other, more cursorial birds that may operate their muscles more economically and achieve greater tendon elastic savings.

Animals↗
PubMed2001

Hindlimb muscle function in relation to speed and gait: in vivo patterns of strain and activation in a hip and knee extensor of the rat (Rattus norvegicus).

Understanding how animals actually use their muscles during locomotion is an important goal in the fields of locomotor physiology and biomechanics. Active muscles in vivo can shorten, lengthen or remain isometric, and their mechanical performance depends on the relative magnitude and timing of these patterns of fascicle strain and activation. It has recently been suggested that terrestrial animals may conserve metabolic energy during locomotion by minimizing limb extensor muscle strain during stance, when the muscle is active, facilitating more economical force generation and elastic energy recovery from limb muscle-tendon units. However, whereas the ankle extensors of running turkeys and hopping wallabies have been shown to generate force with little length change (<6% strain), similar muscles in cats appear to change length more substantially while active. Because previous work has tended to focus on the mechanical behavior of ankle extensors during animal movements, the actions of more proximal limb muscles are less well understood. To explore further the hypothesis of force economy and isometric behavior of limb muscles during terrestrial locomotion, we measured patterns of electromyographic (EMG) activity and fascicle strain (using sonomicrometry) in two of the largest muscles of the rat hindlimb, the biceps femoris (a hip extensor) and vastus lateralis (a knee extensor) during walking, trotting and galloping. Our results show that the biceps and vastus exhibit largely overlapping bursts of electrical activity during the stance phase of each step cycle in all gaits. During walking and trotting, this activity typically commences shortly before the hindlimb touches the ground, but during galloping the onset of activity depends on whether the limb is trailing (first limb down) or leading (second limb down), particularly in the vastus. In the trailing limb, the timing of the onset of vastus activity is slightly earlier than that observed during walking and trotting, but in the leading limb, this activity begins much later, well after the foot makes ground contact (mean 7% of the step cycle). In both muscles, EMG activity typically ceases approximately two-thirds of the way through the stance phase. While electrically active during stance, biceps fascicles shorten, although the extent of shortening differs significantly among gaits (P<0.01). Total average fascicle shortening strain in the biceps is greater during walking (23+/-3%) and trotting (27+/-5%) than during galloping (12+/-5% and 19+/-6% in the trailing and leading limbs, respectively). In contrast, vastus fascicles typically lengthen (by 8-16%, depending on gait) over the first half of stance, when the muscle is electrically active, before shortening slightly or remaining nearly isometric over much of the second half of stance. Interestingly, in the leading limb during galloping, vastus fascicles lengthen prior to muscle activation and exhibit substantial shortening (10+/-2%) during the period when EMG activity is recorded. Thus, patterns of muscle activation and/or muscle strain differ among gaits, between muscles and even within the same muscle of contralateral hindlimbs (as during galloping). In contrast to the minimal strain predicted by the force economy hypothesis, our results suggest that proximal limb muscles in rats operate over substantial length ranges during stance over various speeds and gaits and exhibit complex and changing activation and strain regimes, exemplifying the variable mechanical roles that muscles can play, even during level, steady-speed locomotion.

Animals↗
PubMed2001

Pectoralis muscle performance during ascending and slow level flight in mallards (Anas platyrhynchos).

In vivo measurements of pectoralis muscle length change and force production were obtained using sonomicrometry and delto-pectoral bone strain recordings during ascending and slow level flight in mallards (Anas platyrhynchos). These measurements provide a description of the force/length properties of the pectoralis under dynamic conditions during two discrete flight behaviors and allow an examination of the effects of differences in body size and morphology on pectoralis performance by comparing the results with those of a recent similar study of slow level flight in pigeons (Columbia livia). In the present study, the mallard pectoralis showed a distinct pattern of active lengthening during the upstroke. This probably enhances the rate of force generation and the magnitude of the force generated and, thus, the amount of work and power produced during the downstroke. The power output of the pectoralis averaged 17.0 W kg(-)(1 )body mass (131 W kg(-)(1 )muscle mass) during slow level flight (3 m s(-)(1)) and 23.3 W kg(-)(1 )body mass (174 W kg(-)(1 )muscle mass) during ascending flight. This increase in power was achieved principally via an increase in muscle strain (29 % versus 36 %), rather than an increase in peak force (107 N versus 113 N) or cycle frequency (8.4 Hz versus 8.9 Hz). Body-mass-specific power output of mallards during slow level flight (17.0 W kg(-)(1)), measured in terms of pectoralis mechanical power, was similar to that measured recently in pigeons (16.1 W kg(-)(1)). Mallards compensate for their greater body mass and proportionately smaller wing area and pectoralis muscle volume by operating with a high myofibrillar stress to elevate mechanical power output.

Animals↗
PubMed2001

Mechanics of limb bone loading during terrestrial locomotion in the green iguana (Iguana iguana) and American alligator (Alligator mississippiensis).

In vivo measurements of strain in the femur and tibia of Iguana iguana (Linnaeus) and Alligator mississippiensis (Daudin) have indicated three ways in which limb bone loading in these species differs from patterns observed in most birds and mammals: (i) the limb bones of I. iguana and A. mississippiensis experience substantial torsion, (ii) the limb bones of I. iguana and A. mississippiensis have higher safety factors than those of birds or mammals, and (iii) load magnitudes in the limb bones of A. mississippiensis do not decrease uniformly with the use of a more upright posture. To verify these patterns, and to evaluate the ground and muscle forces that produce them, we collected three-dimensional kinematic and ground reaction force data from subadult I. iguana and A. mississippiensis using a force platform and high-speed video. The results of these force/kinematic studies generally confirm the loading regimes inferred from in vivo strain measurements. The ground reaction force applies a torsional moment to the femur and tibia in both species; for the femur, this moment augments the moment applied by the caudofemoralis muscle, suggesting large torsional stresses. In most cases, safety factors in bending calculated from force/video data are lower than those determined from strain data, but are as high or higher than the safety factors of bird and mammal limb bones in bending. Finally, correlations between limb posture and calculated stress magnitudes in the femur of I. iguana confirm patterns observed during direct bone strain recordings from A. mississippiensis: in more upright steps, tensile stresses on the anterior cortex decrease, but peak compressive stresses on the dorsal cortex increase. Equilibrium analyses indicate that bone stress increases as posture becomes more upright in saurians because the ankle and knee extensor muscles exert greater forces during upright locomotion. If this pattern of increased bone stress with the use of a more upright posture is typical of taxa using non-parasagittal kinematics, then similar increases in load magnitudes were probably experienced by lineages that underwent evolutionary shifts to a non-sprawling posture. High limb bone safety factors and small body size in these lineages could have helped to accommodate such increases in limb bone stress.

Alligators and Crocodiles↗
PubMed2000

Hindlimb extensor muscle function during jumping and swimming in the toad (Bufo marinus).

Many anurans use their hindlimbs to generate propulsive forces during both jumping and swimming. To investigate the musculoskeletal dynamics and motor output underlying locomotion in such physically different environments, we examined patterns of muscle strain and activity using sonomicrometry and electromyography, respectively, during jumping and swimming in the toad Bufo marinus. We measured strain and electromyographic (EMG) activity in four hindlimb muscles: the semimembranosus, a hip extensor; the plantaris, an ankle extensor; and the gluteus and cruralis, two knee extensors. During jumping, these four muscles are activated approximately simultaneously; however, joint extension appears to be temporally staggered, with the hip beginning to extend prior to or initially faster than the more distal knee and ankle joints. Mirroring this pattern, the gluteus and plantaris shorten quite slowly and over a small distance during the first half of limb extension during take-off, before beginning to shorten rapidly. The hip and knee extensors finish shortening near the point of take-off (when the feet leave the ground), while the ankle-extending plantaris, which exhibits the longest-duration EMG burst, on average, always completes its shortening after take-off (mean 26 ms). During swimming, activation of the four muscles is also nearly synchronous at the start of a propulsive stroke. The onset of fascicle shortening is temporally staggered, with the knee extensors beginning to shorten first, prior to the hip and ankle extensors. In addition, the knee extensors also often exhibit some degree of slow passive shortening prior to the onset of EMG activity. The offset of muscle shortening during swimming is also staggered, and to a much greater extent than during jumping. During swimming, the cruralis and gluteus finish shortening first, the semimembranosus finishes 30-60 ms later, and the plantaris, which again exhibits the longest EMG burst, finishes shortening last (mean 150 ms after the cruralis). Interestingly, much of this extended shortening in the plantaris occurs at a relatively slow velocity and may reflect passive ankle extension caused by fluid forces, associated with previously generated unsteady (accelerative) limb movements, acting on the foot. Average EMG burst intensity tends to be greater during jumping than during swimming in all muscles but the gluteus. However, EMG burst duration only changes between jumping and swimming in the cruralis (duration during jumping is nearly twice as long as during swimming). The cruralis is also the only muscle to exhibit substantially greater fractional shortening during jumping (mean 0.28) than during swimming (mean 0.20 active strain, 0.22 total strain). On the basis of these results, it appears that toad hindlimb function is altered between jumping and swimming. Moreover, these functional differences are influenced by passive effects associated with physical differences between the external environments, but are also actively mediated by shifts in the motor output and mechanical behavior of several muscles.

Animals↗
PubMed1999

Experimental alteration of limb posture in the chicken (Gallus gallus) and its bearing on the use of birds as analogs for dinosaur locomotion.

Extant birds represent the only diverse living bipeds, and can be informative for investigations into the life-history parameters of their extinct dinosaurian relatives. However, morphological changes that occurred during early avian evolution, including the unique adoption of a nearly horizontal femoral orientation associated with a shift in center of mass (CM), suggest that caution is warranted in the use of birds as analogs for nonavian dinosaur locomotion. In this study, we fitted a group of white leghorn chickens (Gallus gallus) with a weight suspended posterior to the hip in order to examine the effects on loading and morphology. This caused a CM shift that necessitated a change in femoral posture (by 35 degrees towards the horizontal, P < 0.001), and resulted in reorientation of the ground reaction force (GRF) vector relative to the femur (from 41 degrees to 82 degrees, P < 0.001). Despite similar strain magnitudes, an overall increase in torsion relative to bending (from 1.70 to 1.95 times bending, P < 0.001) was observed, which was weakly associated with a tendency for increased femoral cross-sectional dimensions (P = 0.1). We suggest that a relative increase in torsion is consistent with a change in femoral posture towards the horizontal, since this change increases the degree to which the bone axis and the GRF vector produce mediolateral long-axis rotation of the bone. These results support the hypothesis that a postural change during early avian evolution could underlie the allometric differences seen between bird and nonavian dinosaur femora by requiring more robust femoral dimensions in birds due to an increase in torsion.

Animals↗
PubMed1999

Dynamics of muscle function during locomotion: accommodating variable conditions.

Much of what we know about animal locomotion is derived from studies examining animals moving within a single, homogeneous environment, at a steady speed and along a flat grade. As a result, the issue of how musculoskeletal function might shift to accommodate variability within the external environment has remained relatively unexplored. One possibility is that locomotor muscles are differentially recruited depending upon the environment in which the animal is moving. A second possibility is that the same muscles are recruited, but that they are activated in a different manner so that their contractile function differs according to environment. Finally, it is also possible that, in some cases, animals may not need to alter their musculoskeletal function to move under different external conditions. In this case, however, the mechanical behavior appropriate for one environmental condition may constrain locomotor performance in another. To begin to explore the means by which animals accommodate variable conditions in their environment, we present three case studies examining how musculoskeletal systems function to allow locomotion under variable conditions: (1) eels undulating through water and across land, (2) turkeys running on level and inclined surfaces, and (3) ducks using their limbs to walk and to paddle. In all three of these examples, the mechanical behavior of some muscle(s) involved in locomotion are altered, although to different degrees and in different ways. In the running turkeys, the mechanical function of a major ankle extensor muscle shifts from contracting isometrically on a flat surface (producing little work and power), to shortening actively during contraction on an uphill gradient (increasing the amount of work and power generated). In the ducks, the major ankle extensor undergoes the same general pattern of activation and shortening in water and on land, except that the absolute levels of muscle stress and strain and work output are greater during terrestrial locomotion. In eels, a transition to land elicits changes in electromyographic duty cycles and the relative timing of muscle activation, suggesting some alteration in the functional mechanics of the underlying musculature. Only by studying muscle function in animals moving under more variable conditions can we begin to characterize the functional breadth of the vertebrate musculoskeletal system and understand more fully its evolutionary design.

Animals↗
PubMed1998

Evaluation of a bone's in vivo 24-hour loading history for physical exercise compared with background loading.

The present study sought to answer two research questions. First, how distinctive, as a potential osteogenic stimulus, are short-duration bouts of treadmill exercise relative to sedentary background activity? Second, how well does daily effective strain stimulus relate the loading history for one such exercise program, in comparison with other experimental loading programs, to bone formation? In vivo cortical strains were measured in the tibiotarsus of White Leghorn chickens at a late stage of skeletal growth (14-34 weeks old) under the conditions of a previous investigation of bone formation in response to an exercise program (15 min/day, treadmill gait at 60% maximum speed while carrying 20% body mass) that included sedentary background activity. These strain data were compiled into 24-hour loading histories of peak cyclic strain, demonstrating that strains were statistically different for exercise and background activities (p < 0.0001), with both the magnitude and number of cyclic strain events being greater during exercise (generally greater than 500 microstrain, 2,500 cycles/day) than during background activity (generally less than 500 microstrain, mean: 775 cycles/day). Strains during exercise accounted for more than 97% of the daily effective strain stimulus for bone adaptation, despite the fact that exercise comprised only 1% of the daily period (15 min/day). The levels of the daily effective strain stimulus were similar to those calculated for strains engendered by artificial loading of functionally isolated avian ulnae, which either maintained bone mass or resulted in a 15% increase of cortical cross-sectional area in both sets of studies. These results indicate that short-duration bouts of treadmill exercise and sedentary background activity can represent distinct osteogenic stimuli for adaptive bone modeling. They also provide experimental support for the use of a daily effective strain stimulus to quantify skeletal loading histories for differing programs of physical exercise, although the relative importance of other mechanical and nonmechanical factors requires further investigation.

Animals↗