Functional differentiation of fore- and hindlimb muscles inMacaca fuscata determined on the basis of enzymatic activities

When the functional differentiation of 83 kinds of limb and trunk muscles ofMacaca fuscata was investigated on the basis of the activities of two glycolytic enzymes [lactate dehydrogenase (LDH) and aldolase] and one oxidative enzyme [succinate dehydrogenase (SDH)], the forelimb rather than the hindlimb muscles proved have higher oxidative activities. These results indicated that, inMacaca fuscata, the forelimb muscles have a higher resistance to fatigue, and that the hindlimb muscles have a higher tetanic tension on the basis of the relationships between enzymatic activities and functional properties of the muscle fiber types. These findings were interpreted in relation to the fact thatMacaca fuscata is a quadrupedal primate with arboreal habits, as compared with nonprimate terrestrial quadrupeds. The two-joint muscles and the superficial muscles contract more rapidly than do the other muscles in the hindlimb, thereby suggesting that both types of muscles readily adapt to quick movement.     

Comparison of hind limb muscle mass in neonate and adult prosimian primates

Little ontogenetic data exist to indicate whether muscular organization of neonates reflects adult locomotion (e.g., leaping) or infant activities like clinging or the initial quadrupedal phase of locomotion that typifies most infant primates. In the present study, five species of primates with contrasting modes of locomotion were examined. Twenty-eight preserved neonatal and adult cadavers were studied by careful dissection of the hip, thigh, and leg muscles. Wet weights were taken of limb muscles after removal, and the muscles were combined into major functional groups (e.g., flexors, extensors) of each limb segment. Results demonstrate that the distribution of muscle mass within the thigh and within the leg are similar between neonates and adults for all species, with major groups varying by 5% or less in all but two age comparisons. Crural indices of the neonates are nearly identical to those of the adults, but leg/thigh muscle mass ratios were higher in the neonates. Species vary greatly in the percentage of adult limb segment muscle mass present in neonates, with Tarsius syrichta having the greatest percentage for all segments and two lemurids showing the least. These results primarily track differences in relative body mass at birth rather than developmental differences. The adaptive distribution of muscle, as discussed previously for adult prosimians, appears to be established at birth. Neonates of leaping species already have much larger quadriceps muscles than quadrupeds. Differences between large- and small-bodied leapers (e.g., pronounced superficial plantarflexor masses in tarsiers and pronounced deep plantarflexor masses in sifakas) also are present in neonates. Ratios of muscle mass over body mass are smaller in all neonates than in their adult counterparts, suggesting that the neonates are relatively poorly muscled, and that muscle mass must increase with positive allometry during growth.     

Locomotion in ornithischian dinosaurs: an assessment using three‐dimensional computational modelling

Ornithischian dinosaurs were primitively bipedal with forelimbs modified for grasping, but quadrupedalism evolved in the clade on at least three occasions independently. Outside of Ornithischia, quadrupedality from bipedal ancestors has only evolved on two other occasions, making this one of the rarest locomotory transitions in tetrapod evolutionary history. The osteological and myological changes associated with these transitions have only recently been documented, and the biomechanical consequences of these changes remain to be examined. Here, we review previous approaches to understanding locomotion in extinct animals, which can be broadly split into form–function approaches using analogy based on extant animals, limb‐bone scaling, and computational approaches. We then carry out the first systematic attempt to quantify changes in locomotor muscle function in bipedal and quadrupedal ornithischian dinosaurs. Using three‐dimensional computational modelling of the major pelvic locomotor muscle moment arms, we examine similarities and differences among individual taxa, between quadrupedal and bipedal taxa, and among taxa representing the three major ornithischian lineages (Thyreophora, Ornithopoda, Marginocephalia). Our results suggest that the ceratopsid Chasmosaurus and the ornithopod Hypsilophodon have relatively low moment arms for most muscles and most functions, perhaps suggesting poor locomotor performance in these taxa. Quadrupeds have higher abductor moment arms than bipeds, which we suggest is due to the overall wider bodies of the quadrupeds modelled. A peak in extensor moment arms at more extended hip angles and lower medial rotator moment arms in quadrupeds than in bipeds may be due to a more columnar hindlimb and loss of medial rotation as a form of lateral limb support in quadrupeds. We are not able to identify trends in moment arm evolution across Ornithischia as a whole, suggesting that the bipedal ancestry of ornithischians did not constrain the development of quadrupedal locomotion via a limited number of functional pathways. Functional anatomy appears to have had a greater effect on moment arms than phylogeny, and the differences identified between individual taxa and individual clades may relate to differences in locomotor performance required for living in different environments or for clade‐specific behaviours.     

Musculoskeletal Geometry,Muscle Architecture and Functional Specialisations of the Mouse Hindlimb

Mice are one of the most commonly used laboratory animals, with an extensive array of disease models in existence, including for many neuromuscular diseases. The hindlimb is of particular interest due to several close muscle analogues/homologues to humans and other species. A detailed anatomical study describing the adult morphology is lacking, however. This study describes in detail the musculoskeletal geometry and skeletal muscle architecture of the mouse hindlimb and pelvis, determining the extent to which the muscles are adapted for their function, as inferred from their architecture. Using I₂KI enhanced microCT scanning and digital segmentation, it was possible to identify 39 distinct muscles of the hindlimb and pelvis belonging to nine functional groups. The architecture of each of these muscles was determined through microdissections, revealing strong architectural specialisations between the functional groups. The hip extensors and hip adductors showed significantly stronger adaptations towards high contraction velocities and joint control relative to the distal functional groups, which exhibited larger physiological cross sectional areas and longer tendons, adaptations for high force output and elastic energy savings. These results suggest that a proximo-distal gradient in muscle architecture exists in the mouse hindlimb. Such a gradient has been purported to function in aiding locomotor stability and efficiency. The data presented here will be especially valuable to any research with a focus on the architecture or gross anatomy of the mouse hindlimb and pelvis musculature, but also of use to anyone interested in the functional significance of muscle design in relation to quadrupedal locomotion.     

Computational modelling of muscle fibre operating ranges in the hindlimb of a small ground bird (Eudromia elegans), with implications for modelling locomotion in extinct species

The arrangement and physiology of muscle fibres can strongly influence musculoskeletal function and whole-organismal performance. However, experimental investigation of muscle function during in vivo activity is typically limited to relatively few muscles in a given system. Computational models and simulations of the musculoskeletal system can partly overcome these limitations, by exploring the dynamics of muscles, tendons and other tissues in a robust and quantitative fashion. Here, a high-fidelity, 26-degree-of-freedom musculoskeletal model was developed of the hindlimb of a small ground bird, the elegant-crested tinamou (Eudromia elegans, ~550 g), including all the major muscles of the limb (36 actuators per leg). The model was integrated with biplanar fluoroscopy (XROMM) and forceplate data for walking and running, where dynamic optimization was used to estimate muscle excitations and fibre length changes throughout both gaits. Following this, a series of static simulations over the total range of physiological limb postures were performed, to circumscribe the bounds of possible variation in fibre length. During gait, fibre lengths for all muscles remained between 0.5 to 1.21 times optimal fibre length, but operated mostly on the ascending limb and plateau of the active force-length curve, a result that parallels previous experimental findings for birds, humans and other species. However, the ranges of fibre length varied considerably among individual muscles, especially when considered across the total possible range of joint excursion. Net length change of muscle–tendon units was mostly less than optimal fibre length, sometimes markedly so, suggesting that approaches that use muscle–tendon length change to estimate optimal fibre length in extinct species are likely underestimating this important parameter for many muscles. The results of this study clarify and broaden understanding of muscle function in extant animals, and can help refine approaches used to study extinct species.     

Fiber architecture of the intrinsic muscles of the shoulder and arm in semiterrestrial and arboreal guenons

The internal organization of myofibers and connective tissues has important physiologic implications for muscle function and for naturalistic behavior. In this study of forelimb muscle morphology and primate locomotion, fiber architecture is examined in the intrinsic muscles of the shoulder (musculi deltoideus, infraspinatus, supraspinatus, subscapularis, teres major, and t. minor) and arm (m. coracobrachialis, biceps brachii, brachialis, and triceps brachii) in the semiterrestrial vervets (Chlorocebus aethiops) and arboreal red-tailed guenons (Cercopithecus ascanius). Wet weights and lengths of whole muscles, lengths of fasciculi and their associated proximal and distal tendons, and angles of pinnation were measured to estimate morphologic correlates of physiologic properties of individual muscles: force, velocity/excursion, energy expense, and relative isometric or isotonic contraction. Neither mean total-shoulder:total-arm ratios for muscle mass nor total reduced physiological cross-sectional area exhibited significant (P < 0.05) interspecific differences, thus emphasizing the importance of fine-tuning musculoskeletal analyses by the data collected here. The results generally support those previously published for quadriceps femoris and triceps surae of the hind limb in these species (Anapol and Barry [1996] Am. J. Phys. Anthropol. 99:429-447). The fiber architecture of the semiterrestrial vervets is largely suited for higher velocity while running on the ground. By contrast, the architectural configuration of red-tailed monkeys implies relatively isometric muscle contraction and passive storage of elastic strain energy for exploitation of the compliant canopy, where substrate components are situated beneath the sagittal plane of the animal. With respect to relative distribution of maximum potential force output among muscles of either shoulder or arm groups in these otherwise hind limb-dominated quadrupedal primates, statistically significant interspecific differences are best interpreted in light of braking, climbing, and, for vervets, the transition between ground and canopy.The interspecific differences shown here for the intrinsic muscles of the shoulder and arm underscore the significance of intramuscular morphology in reconciling structure and function with regard to locomotor behavior. Its analysis and interpretation lend support to consideration of "semiterrestrial" as a bona fide locomotor category uniquely different from what is practiced by dedicated arboreal and terrestrial quadrupeds that occasionally visit the habitat of one another. Data from a more committed terrestrial species would clarify this enigma.     

Coordinated network functioning in the spinal cord: An evolutionary perspective

The successful achievement of harmonious locomotor movement results from the integrated operation of all body segments. Here, we will review current knowledge on the functional organization of spinal networks involved in mammalian locomotion. Attention will not simply be restricted to hindlimb muscle control, but by also considering the necessarily coordinated activation of trunk and forelimb muscles, we will try to demonstrate that while there has been a progressive increase in locomotor system complexity during evolution, many basic organizational features have been preserved across the spectrum from lower vertebrates through to humans. Concerning the organization of axial neuronal networks that control trunk muscles, it has been found across the vertebrate range that during locomotor movement a motor wave travels longitudinally in the spinal cord via the coupling of rhythmic segmental networks. For hindlimb activation it has been found in all species studied that the rostral lumbar segments contain the key elements for pattern generation. We also showed that rhythmic arm movements are under the control of cervical forelimb generators in quadrupeds as well as in human. Finally, it is highlighted that the coordination of quadrupedal movements during locomotion derives principally from an asymmetrical coordinating influence occurring in the caudo-rostral direction from the lumbar hindlimb networks.     

External forces on the limbs of jumping lemurs at takeoff and landing

Ground reaction forces were recorded for jumps of three individuals each of Lemur catta and Eulemur fulvus. Animals jumped back and forth between a ground-mounted force plate and a 0.5-m elevated platform, covering horizontal distances of 0.5-2 m. In total, 190 takeoffs and 263 landings were collected. Animals typically jumped from a run up and into a run out, during which they gained or into which they carried horizontal impulse. Correspondingly, vertical impulses dominated takeoffs and landings. Peak forces were moderate in magnitude and not much higher than forces reported for quadrupedal gaits. This is in contrast to the forces for standing jumps of specialized leapers that considerably exceed forces associated with quadrupedal gaits. Force magnitudes for the lemur jumps are more comparable to peak forces reported for other quadrupeds performing running jumps. Takeoffs are characterized by higher hindlimb than forelimb peak forces and impulses. L. catta typically landed with the hindlimbs making first contact, and the hindlimb forces and impulses were higher than the forelimb forces and impulses at landing. E. fulvus typically landed with the forelimbs striking first and also bearing the higher forces. This pattern does not fully conform to the paradigm of primate limb force distribution, with higher hindlimb than forelimb forces. However, the absolute highest forces in E. fulvus also occur at the hindlimbs, during acceleration for takeoff.     

Are fixed limb inertial models valid for dynamic simulations of human movement?

During human movement, muscle activation and limb movement result in subtle changes in muscle mass distribution. Muscle mass redistribution can affect limb inertial properties and limb dynamics, but it is not currently known to what extent. The objectives of this study were to investigate: (1) how physiological alterations of muscle and tendon length affect limb inertial characteristics, and (2) how such changes affect dynamic simulations of human movement. To achieve these objectives, a digital model of a human leg, custom software, and Software for interactive musculoskeletal modeling were used to simulate mass redistribution of muscle–tendon structures within a limb segment during muscle activation and joint movement. Thigh and shank center of mass and moments of inertia for different muscle activation and joint configurations were determined and compared. Limb inertial parameters representing relaxed muscles and fully active muscles were input into a simulated straight-leg movement to evaluate the effect inertial parameter variations could have on movement simulation results. Muscle activation and limb movement altered limb segment center of mass and moments of inertia by less than 0.04 cm and 1.2%, respectively. These variations in limb inertial properties resulted in less than 0.01% change in maximum angular velocity for a simulated straight-leg hip flexion task. These data demonstrate that, for the digital human leg model considered, assuming static quantities for segment center of masses and moments of inertia in movement simulations appear reasonable and induce minimal errors in simulated movement dynamics.     

Unifying principles in terrestrial locomotion: do hopping Australian marsupials fit in?

Mammalian terrestrial locomotion has many unifying principles. However, the Macropodoidea are a particularly interesting group that exhibit a number of significant deviations from the principles that seem to apply to other mammals. While the properties of materials that comprise the musculoskeletal system of mammals are similar, evidence suggests that tendon properties in macropodoid marsupials may be size or function dependent, in contrast to the situation in placental mammals. Postural differences related to hopping versus running have a dramatic effect on the scaling of the pelvic limb musculoskeletal system. Ratios of muscle fibre to tendon cross-sectional areas for ankle extensors and digital flexors scale with positive allometry in all mammals, but exponents are significantly higher in macropods. Tendon safety factors decline with increasing body mass in mammals, with eutherians at risk of ankle extensor tendon rupture at a body mass of about 150 kg, whereas kangaroos encounter similar problems at a body mass of approximately 35 kg. Tendon strength appears to limit locomotor performance in these animals. Elastic strain energy storage in tendons is mass dependent in all mammals, but exponents are significantly larger in macropodid. Tibial stresses may scale with positive allometry in kangaroos, which result in lower bone safety factors in macropods compared to eutherian mammals.     

Pennation angles of the intrinsic muscles of the foot

As mathematical models of the musculoskeletal system become increasingly detailed and precise, they require more accurate information about the architectural parameters of the individual muscles. These muscles are typically represented as Hill-type models, which require data on fiber length, physiological cross-sectional area (PCSA) and pennation angle. Most of this information for lower limb muscles has been published, except for data on the pennation angle of the intrinsic muscles of the foot. Each (n=20) intrinsic muscle of three human feet was dissected free. The dorsal and plantar surfaces were photographed and a digitized color image was imported into Abobe Photoshop. The muscles were divided into "anatomical units". For each anatomical unit (n=26), a line was drawn along the tendon axis and a number of other lines were drawn along individual muscle fibers. The angle between the tendon line and each fiber line was defined as the pennation angle of that fiber. By visual inspection, an effort was made to take measurements such that they represented the distribution of fibers in various parts of the muscle. Although some individual muscles had higher or lower pennation angles, when averaged for all specimens, the second dorsal interosseous had the smallest pennation angle (6.7+/-6.81 degrees) while the abductor digiti minimi had the largest (19.1+/-11.19 degrees). Since the cosines of the angles range from 0.9932 to 0.9449, the effect of the pennation angle on the force generated by the muscle was not great.     

Dimensions and moment arms of the hind- and forelimb muscles of common chimpanzees (Pan troglodytes).

This paper supplies quantitative data on the hind- and forelimb musculature of common chimpanzees (Pan troglodytes) and calculates maximum joint moments of force as a contribution to a better understanding of the differences between chimpanzee and human locomotion. We dissected three chimpanzees, and recorded muscle mass, fascicle length, and physiological cross-sectional area (PCSA). We also obtained flexion/extension moment arms of the major muscles about the limb joints. We find that in the hindlimb, chimpanzees possess longer fascicles in most muscles but smaller PCSAs than are predicted for humans of equal body mass, suggesting that the adaptive emphasis in chimpanzees is on joint mobility at the expense of tension production. In common chimpanzee bipedalism, both hips and knees are significantly more flexed than in humans, necessitating muscles capable of exerting larger moments at the joints for the same ground force. However, we find that when subject to the same stresses, chimpanzee hindlimb muscles provide far smaller moments at the joints than humans, particularly the quadriceps and plantar flexors. In contrast, all forelimb muscle masses, fascicle lengths, and PCSAs are smaller in humans than in chimpanzees, reflecting the use of the forelimbs in chimpanzee, but not human, locomotion. When subject to the same stresses, chimpanzee forelimb muscles provide larger moments at the joints than humans, presumably because of the demands on the forelimbs during locomotion. These differences in muscle architecture and function help to explain why chimpanzees are restricted in their ability to walk, and particularly to run bipedally.     

Modeling short-range stiffness of feline lower hindlimb muscles

The short-range stiffness (SRS) of skeletal muscles is a critical property for understanding muscle contributions to limb stability, since it represents a muscle's capacity to resist external perturbations before reflexes or voluntary actions can intervene. A number of studies have demonstrated that a simple model, consisting of a force-dependent active stiffness connected in series with a constant passive stiffness, is sufficient to characterize the SRS of individual muscles over the entire range of obtainable forces. The purpose of this study was to determine if such a model could be used to characterize the SRS-force relationship in a number of architecturally distinct muscles. Specifically, we hypothesized that the active and passive stiffness components for a specific muscle can be estimated from anatomical measurements, assuming uniform active and passive stiffness properties across all muscles. This hypothesis was evaluated in six feline lower hindlimb muscle types with different motor unit compositions and architectures. The SRS-force relationships for each muscle type were predicted based on anatomical measurements and compared to experimental data. The model predictions were accurate to within 30%, when uniform scaling properties were assumed across all muscles. Errors were the greatest for the extensor digitorum longus (EDL). When this muscle was removed from the analysis, prediction errors dropped to less than 8%. Subsequent analyses suggested that these errors might have resulted from differences in the tendon elastic modulus, as compared to the other muscles tested.     

Finding the neck–trunk boundary in snakes: Anteroposterior dissociation of myological characteristics in snakes and its implications for their neck and trunk body regionalization

The neck and trunk regionalization of the presacral musculoskeletal system in snakes and other limb‐reduced squamates was assessed based on observations on craniovertebral and body wall muscles. It was confirmed that myological features characterizing the neck in quadrupedal squamates (i.e., squamates with well‐developed limbs) are retained in all examined snakes, contradicting the complete lack of the neck in snakes hypothesized in previous studies. However, the posterior‐most origins of the craniovertebral muscles and the anterior‐most bony attachments of the body wall muscles that are located at around the neck–trunk boundary in quadrupedal squamates were found to be dissociated anteroposteriorly in snakes. Together with results of a recent study that the anterior expression boundaries of Hox genes coinciding with the neck–trunk boundary in quadrupedal amniotes were dissociated anteroposteriorly in a colubrid snake, these observations support the hypothesis that structures usually associated with the neck–trunk boundary in quadrupedal squamates are displaced relative to one another in snakes. Whereas certain craniovertebral muscles are elongated in some snakes, results of optimization on an ophidian cladogram show that the most recent common ancestor of extant snakes would have had the longest craniovertebral muscle, M. rectus capitis anterior, that is elongated only by several segments compared with that of quadrupedal squamates. Therefore, even such a posteriorly displaced “cervical” characteristic plesiomorphically lies fairly anteriorly in the greatly elongated precloacal region of snakes, suggesting that the trunk, not the neck, would have contributed most to the elongation of the snake precloacal region. A similar dissociation of structures usually associated with the neck–trunk boundary in quadrupedal squamates is observed in limb‐reduced squamates, suggesting that these forms and snakes may share a developmental mechanism producing modifications in the anterior–posterior patterning associated with body elongation. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Three-dimensional model of the feline hindlimb

This article describes a three-dimensional musculoskeletal model of the feline hindlimb based on digitized musculoskeletal anatomy. The model consists of seven degrees of freedom: three at the hip and two each at the knee and ankle. Lines of action and via points for 32 major muscles of the limb are described. Interspecimen variability of muscle paths was surprisingly low; most via points displayed a scatter of only a few millimeters. Joint axes identified by mechanical techniques as noncoincident and nonorthogonal were further honed to yield moment arms consistent with previous reports. Interspecimen variability in joint axes was greater than that of muscle paths and highlights the importance of joint axes in kinematic models. The contribution of specific muscles to the direction of endpoint force generation is discussed.     

Accelerated contractile function and improved fatigue resistance of calf muscles in newborn piglets with IUGR

Asymmetrical intrauterine growth restriction is denoted by disproportional reduction of muscle mass compared with body weight reduction. However, effects on contractile function or tissue development of skeletal muscles were not studied until now. Therefore, isometric force output of serial-stimulated hindlimb plantar flexors was measured in thiopental-anesthetized normal weight (NW) and intrauterine growth-restricted (IUGR) 1-day-old piglets under conditions of normal, reduced (aortic cross clamping), and reestablished (clamp release) blood supply (measured by colored microspheres technique). Furthermore, muscle fiber type distribution was determined after histochemical staining, specific muscle force of the plantar flexors [quotient from absolute force divided by muscle mass (N/g)] was calculated, and glycogen content and morphometric data of the investigated muscles were estimated. Regional blood flow of hindlimb muscles was similar in NW (6 +/- 2 ml. min(-1). 100 g(-1)) and IUGR piglets (8 +/- 1 ml. min(-1). 100 g(-1)). Isometric muscle contractions induced a marked increase in regional blood flow of 4.1-fold in NW and 5-fold in stimulated hindlimb muscles of IUGR piglets (baseline blood flow). Specific force of NW piglet muscles (5.2 +/- 0.2 N/g) was significantly lower than IUGR piglet muscles (6.1 +/- 0.6 N/g; P < 0.05). Isometric muscle contractions (NW: 32.7 +/- 4.7 N; IUGR: 21.7 +/- 4.0 N) resulted in a higher rate of force decrease in the calf muscles of NW animals compared with IUGR piglets (8 +/- 2 vs. 3 +/- 1%; P < 0. 01). Functional restoration of contractile performance after hindlimb recirculation was nearly complete in IUGR piglets (98 +/- 1%), whereas in NW piglets a deficit of 9 +/- 3% was found (P < 0. 01). Muscle fiber type estimation revealed an increased proportion of type I fibers in flexor digitalis superficialis and gastrocnemius medialis in IUGR piglets (P < 0.05). These data clearly indicate that contractile function is accelerated in newborn IUGR piglets.     

A musculoskeletal model of the hand and wrist: model definition and evaluation

Abstract

To improve our understanding on the neuromechanics of finger movements, a comprehensive musculoskeletal model is needed. The aim of this study was to build a musculoskeletal model of the hand and wrist, based on one consistent data set of the relevant anatomical parameters. We built and tested a model including the hand and wrist segments, as well as the muscles of the forearm and hand in OpenSim. In total, the model comprises 19 segments (with the carpal bones modeled as one segment) with 23 degrees of freedom and 43 muscles. All required anatomical input data, including bone masses and inertias, joint axis positions and orientations as well as muscle morphological parameters (i.e. PCSA, mass, optimal fiber length and tendon length) were obtained from one cadaver of which the data set was recently published. Model validity was investigated by first comparing computed muscle moment arms at the index finger metacarpophalangeal (MCP) joint and wrist joint to published reference values. Secondly, the muscle forces during pinching were computed using static optimization and compared to previously measured intraoperative reference values. Computed and measured moment arms of muscles at both index MCP and wrist showed high correlation coefficients (r?=?0.88 averaged across all muscles) and modest root mean square deviation (RMSD?=?23% averaged across all muscles). Computed extrinsic flexor forces of the index finger during index pinch task were within one standard deviation of previously measured in-vivo tendon forces. These results provide an indication of model validity for use in estimating muscle forces during static tasks.     

Muscle weights and succinic dehydrogenase distribution in the hind limb musculature of two rodents (Thryonomys gregorianus and Pedetes capensis) with different locomotory habits

The hind limb muscles of the spring hare (Pedetes capensis) were found to be relatively heavier than the hind limb muscles of the cane rat (Thryonomys gregorianus). The distribution of succine dehydrogenase activity was investigated in four of these muscles (m. gluteus superficialis, m.semimembranosus, m. biceps femoris and m.rectus femoris) from both animals. It was found that the spring hare had a higher proportion of low-activity fibres in all four muscles than the cane rat. All muscle fibre types were also smaller in diameter in the spring hare than the cane rat. These results are discussed in relation to the different locomotry habits of the two animals.     

A three-dimensional model of the rat hindlimb: musculoskeletal geometry and muscle moment arms

As a first step towards developing a dynamic model of the rat hindlimb, we measured muscle attachment and joint center coordinates relative to bony landmarks using stereophotogrammetry. Using these measurements, we analyzed muscle moment arms as functions of joint angle for most hindlimb muscles, and tested the hypothesis that postural change alone is sufficient to alter the function of selected muscles of the leg. We described muscle attachment sites as second-order curves. The length of the fit parabola and residual errors in the orthogonal directions give an estimate of muscle attachment sizes, which are consistent with observations made during dissection. We modeled each joint as a moving point dependent on joint angle; relative endpoint errors less than 7% indicate this method as accurate. Most muscles have moment arms with a large range across the physiological domain of joint angles, but their moment arms peak and vary little within the locomotion domain. The small variation in moment arms during locomotion potentially simplifies the neural control requirements during this phase. The moment arms of a number of muscles cross zero as angle varies within the quadrupedal locomotion domain, indicating they are intrinsically stabilizing. However, in the bipedal locomotion domain, the moment arms of these muscles do not cross zero and thus are no longer intrinsically stabilizing. We found that muscle function is largely determined by the change in moment arm with joint angle, particularly the transition from quadrupedal to bipedal posture, which may alter an intrinsically stabilizing arrangement or change the control burden.     

The role of hind limb flexor muscles during swimming in the toad, Bufo marinus

Most work examining muscle function during anuran locomotion has focused largely on the roles of major hind limb extensors during jumping and swimming. Nevertheless, the recovery phase of anuran locomotion likely plays a critical role in locomotor performance, especially in the aquatic environment, where flexing limbs can increase drag on the swimming animal. In this study, I use kinematic and electromyographic analyses to explore the roles of four anatomical flexor muscles in the hind limb of Bufo marinus during swimming: m. iliacus externus, a hip flexor; mm. iliofibularis and semitendinosus, knee flexors; and m. tibialis anticus longus, an ankle flexor. Two general questions are addressed: (1) What role, if any, do these flexors play during limb extension? and (2) How do limb flexors control limb flexion? Musculus iliacus externus exhibits a large burst of EMG activity early in limb extension and shows low levels of activity during recovery. Both m. iliofibularis and m. semitendinosus are biphasically active, with relatively short but intense bursts during limb extension followed by longer and typically weaker secondary bursts during recovery. Musculus tibialis anticus longus becomes active mid way through recovery and remains active through the start of extension in the next stroke. In conclusion, flexors at all three joints exhibit some activity during limb extension, indicating that they play a role in mediating limb movements during propulsion. Further, recovery is controlled by a complex pattern of flexor activation timing, but muscle intensities are generally lower, suggesting relatively low force requirements during this phase of swimming.