Morphological variation of hypaxial musculature in salamanders (Lissamphibia: Caudata)

Despite the acknowledged importance of the locomotory and respiratory functions associated with hypaxial musculature in salamanders, variation in gross morphology of this musculature has not been documented or evaluated within a phylogenetic or ecological context. In this study, we characterize and quantify the morphological variation of lateral hypaxial muscles using phylogenetically and ecologically diverse salamander species from eight families: Ambystomatidae (Ambystoma tigrinum), Amphiumidae (Amphiuma tridactylum), Cryptobranchidae (Cryptobranchus alleganiensis), Dicamptodontidae (Dicamptodon sp.), Plethodontidae (Gyrinophilus porphyriticus), Proteidae (Necturus maculosus), Salamandridae (Pachytriton sp.), and Sirenidae (Siren lacertina). For the lateral hypaxial musculature, we document 1) the presence or absence of muscle layers, 2) the muscle fiber angles of layers at mid‐trunk, and 3) the relative dorsoventral positions and cross‐sectional areas of muscle layers. Combinations of two, three, or four layers are observed. However, all species retain at least two layers with opposing fiber angles. The number of layers and the presence or absence of layers vary within species (Necturus maculosus and Siren lacertina), within genera (e.g., Triturus), and within families. No phylogenetic pattern in the number of layers can be detected with a family‐level phylogeny. Fiber angle variation of hypaxial muscles is considerable: fiber angles of the M. obliquus externus range from 20–80°; M. obliquus internus, 14–34°; M. transversus abdominis, 58–80° (acute angles measured relative to the horizontal septum). Hypaxial musculature comprises 17–37% of total trunk cross‐sectional area. Aquatic salamanders show relatively larger total cross‐sectional hypaxial area than salamanders that are primarily terrestrial. J. Morphol. 241:153–164, 1999. © 1999 Wiley‐Liss, Inc.     

The Evolution of the Functional Role of Trunk Muscles During Locomotion in Adult Amphibians

SYNOPSIS. The axial musculature of all vertebrates consistsof two principal masses, the epaxial and hypaxial muscles. Theprimitive function of both axial muscle masses is to generatelateral bending of the trunk during swimming, as is seen inmost fishes. Within amphibians we see multiple functional andmorphological elaborations of the axial musculature. These elaborationsappear to be associated not only with movement into terrestrialhabits (salamanders), but also with subsequent locomotor specializationsof two of the three major extant amphibian clades (frogs andcaecilians). Salamanders use both epaxial and hypaxial musclesto produce lateral bending during swimming and terrestrial,quadrupedal locomotion. However during terrestrial locomotionthe hypaxial muscles are thought to perform an added function,resisting long-axis torsion of the trunk. Relative to salamanders,frogs have elaborate epaxial muscles, which function to bothstabilize and extend the iliosacral and coccygeosacral joints.These actions are important in the effective use of the hindlimbsduring terrestrial saltation and swimming. In contrast, caecilianshave relatively elaborate hypaxial musculature that is linkedto a helix of connective tissue embedded in the skin. The helixand associated hypaxial muscles form a hydrostatic skeletonaround the viscera that is continuously used to maintain bodyposture and also contributes to forward force production duringburrowing.     

Fiber‐type distribution of the perivertebral musculature in Ambystoma

Many salamanders locomote in aquatic and terrestrial environments. During swimming, body propulsion is solely produced by the axial musculature generating lateral undulations of the trunk and tail. During terrestrial locomotion, the trunk is oscillated laterally in a standing wave, and body propulsion is achieved by concerted trunk and limb muscle action. The goal of this study was to increase our knowledge of the functional morphology of the tetrapod trunk. We investigated the muscle‐fiber‐type distribution and the anatomical cross‐sectional area of all perivertebral muscles in Ambystoma tigrinum and A. maculatum. Muscle‐fiber‐type composition was determined in serial cross‐sections based on m‐ATPase activity. Five different body segments were investigated to test for cranio‐caudal changes along the trunk. The overall fiber‐type distribution was very similar between the species, but A. tigrinum had relatively larger muscles than A. maculatum, which may be related to its digging behavior. None of the perivertebral muscles possessed a homogeneous fiber‐type composition. The M. interspinalis showed a distinct layered organization and may function to ensure the integrity of the spine (local stabilization). The M. dorsalis trunci exhibited the plesiomorphic pattern for notochordates in having a distinct superficial layer of red and intermediate fibers, which covered the central white fibers; therefore, it is suggested to function as a mobilizer and a stabilizer of the trunk, but, may also be involved in modulating body stiffness. Similarly, the M. subvertebralis showed clear regionalizations, implying functional subunits that can stabilize and mobilize the trunk as well as modulate of body stiffness. Cranio‐caudally, neither the fiber‐type composition nor the a‐csa changed dramatically, possibly reflecting the need to perform well in both aquatic and terrestrial habitats. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Morphology and Function of Lateral Hypaxial Musculature in Salamanders

SYNOPSIS. The lateral hypaxial musculature (LHM) of salamandersmay serve as a useful model for understanding the functionsof LHM in tetrapods more generally. Salamanders have betweentwo and four layers of LHM, arranged segmentally in myomeres.These layers produce three primary mechanical actions: theybend the body, pressurize the body, and produce or resist torsionabout the long axis of the body. The optimum muscle fiber anglefor forceful bending is 0° to the long axis, the optimumangle for pressurization is 90°, and the optimum angle fortorsion is 45°. For generating bending and torsional moments,lateral (superficial) muscle layers have greater mechanicaladvantage than medial (deep) layers. For increasing body pressure,by contrast, medial layers have greater mechanical advantage.A comparison of muscle fiber angles in aquatic and terrestrialsalamanders reveals that some aquatic salamanders have one musclelayer with a low fiber angle which may represent a specializationfor swimming. Overall, however, the fiber angles in the LHMof terrestrial and aquatic salamanders are surprisingly similar.In contrast, the pattern of fiber angles in caecilians is different,suggesting that these amphibians use their LHM differently.The fiber angle models and morphological observations presentedhere form a framework which may be useful in future studiesof lateral hypaxial musculature.     

Fiber‐type composition in the perivertebral musculature of lizards: Implications for the evolution of the diapsid trunk muscles

The perivertebral musculature of lizards is critical for the stabilization and the mobilization of the trunk during locomotion. Some trunk muscles are also involved in ventilation. This dual function of trunk muscles in locomotion and ventilation leads to a biomechanical conflict in many lizards and constrains their ability to breathe while running (“axial constraint”) which likely is reflected by their high anaerobic scope. Furthermore, different foraging and predator‐escape strategies were shown to correlate with the metabolic profile of locomotor muscles in lizards. Because knowledge of muscle's fiber‐type composition may help to reveal a muscle's functional properties, we investigated the distribution pattern of muscle fiber types in the perivertebral musculature in two small lizard species with a generalized body shape and subjected to the axial constraint (Dipsosaurus dorsalis, Acanthodactylus maculatus) and one species that circumvents the axial constraint by means of gular pumping (Varanus exanthematicus). Additionally, these species differ in their predator‐escape and foraging behaviors. Using refined enzyme‐histochemical protocols, muscle fiber types were differentiated in serial cross‐sections through the trunk, maintaining the anatomical relationships between the skeleton and the musculature. The fiber composition in Dipsosaurus and Acanthodactylus showed a highly glycolytic profile, consistent with their intermittent locomotor style and reliance on anaerobic metabolism during activity. Because early representatives of diapsids resemble these two species in several postcranial characters, we suggest that this glycolytic profile represents the plesiomorphic condition for diapsids. In Varanus, we found a high proportion of oxidative fibers in all muscles, which is in accordance with its high aerobic scope and capability of sustained locomotion. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

Biomechanics and functional preconditions for terrestrial lifestyle in basal tetrapods,with special consideration of Tiktaalik roseae

The fossil Tiktaalik roseae from the Late Devonian induces clear definition of the biomechanical and functional preconditions for a terrestrial lifestyle including quadrupedal standing and locomotion on limbs. Therefore, we determined the internal stresses in this model organism under the influence of gravity using the finite element method. Stress patterns during symmetrical two-forelimb support result from bending of trunk and neck. During asymmetrical one-forelimb support, as occurs during terrestrial locomotion, torsional stresses are higher than those caused by bending. The observed patterns of compressive stresses correspond well with the arrangement of compression-resistant materials: vertebral column, shoulder girdle and ribs. The tensile stresses are in accordance with the arrangement of longitudinal and oblique muscles forming the body wall. Torsional stresses concentrate along the periphery of the trunk, leaving its cavity free from mechanical stresses. Theoretical mechanics indicate that the flat skull and the mobility of the neck were advantageous for lateral snapping, similar to crocodiles. The same movement on land requires sprawling and flexed forelimbs. Our results can be interpreted as explanations for the tetrapod bauplan as well as confirmation and refinement of existing hypotheses about the lifestyle at the border between water and land of this early predecessor of terrestrial tetrapods.     

Morphological and histochemical characterization of the pectoral fin muscle of batoids

Batoids differ from other elasmobranch fishes in that they possess dorsoventrally flattened bodies with enlarged muscled pectoral fins. Most batoids also swim using either of two modes of locomotion: undulation or oscillation of the pectoral fins. In other elasmobranchs (e.g., sharks), the main locomotory muscle is located in the axial myotome; in contrast, the main locomotory muscle in batoids is found in the enlarged pectoral fins. The pectoral fin muscles of sharks have a simple structure, confined to the base of the fin; however, little to no data are available on the more complex musculature within the pectoral fins of batoids. Understanding the types of fibers and their arrangement within the pectoral fins may elucidate how batoid fishes are able to utilize such unique swimming modes. In the present study, histochemical methods including succinate dehydrogenase (SDH) and immunofluoresence were used to determine the different fiber types comprising these muscles in three batoid species: Atlantic stingray (Dasyatis sabina), ocellate river stingray (Potamotrygon motoro) and cownose ray (Rhinoptera bonasus). All three species had muscles comprised of two muscle fiber types (slow-red and fast-white). The undulatory species, D. sabina and P. motoro, had a larger proportion of fast-white muscle fibers compared to the oscillatory species, R. bonasus. The muscle fiber sizes were similar between each species, though generally smaller compared to the axial musculature in other elasmobranch fishes. These results suggest that batoid locomotion can be distinguished using muscle fiber type proportions. Undulatory species are more benthic with fast-white fibers allowing them to contract their muscles quickly, as a possible means of escape from potential predators. Oscillatory species are pelagic and are known to migrate long distances with muscles using slow-red fibers to aid in sustained swimming.     

Patterns of regional variation in the vertebral column of terrestrial salamanders

Regional variation in the vertebral column of several species of salamanders (families Ambystomatidae, Salamandridae and Plethodontidae) is analyzed. Measurements of three dimensions, centrum length, prezygapophyseal width, and transverse process length, provide the data. Ontogenetic, interspecific, intergeneric and interfamilial patterns of positional variation are diagrammed and discussed. Distinctive patterns of variation characterize the families, genera, and to a lesser extent, the species. The patterns of ambystomatid salamanders are the most generalized, and probably reflect derivation from a primitive ancestral stock. The most specialized conditions occur in the fully terrestrial plethodontids, a group generally considered to be highly derived. Data such as those presented here will aid in the identification of fossils. The patterns described have functional significance. For example, species which have an aquatic larval stage and which return to aquatic breeding sites have vertebrae which taper in length and width behind the pelvis. This is a feature associated with production of a traveling wave in the tail which is necessary for propulsion in water. Fully terrestrial species do not have a tapering column. In them, standing waves, such as occur in the trunk region of all species, typically occur in the tail. The caudal vertebrae of terrestrial species are rather uniform in dimensions for some distance, and the tail is cylindrical in form. Other functionally important features include the narrowing and shortening of some anterior vertebrae, associated with the development of a neck in some species with tongue feeding mechanisms. In contrast, species which use their heads as wedges during locomotion have broadened anterior vertebrae which serve as sites of origin for hypertrophied neck muscles.     

Lung ventilation in salamanders and the evolution of vertebrate air-breathing mechanisms

Functional analysis of lung ventilation in salamanders combined with historical analysis of respiratory pumps provides new perspectives on the evolution of breathing mechanisms in vertebrates. Lung ventilation in the aquatic salamander Necturus maculosus was examined by means of cineradiography, measurement of buccal and pleuroperitoneal cavity pressures, and electromyography of hypaxial musculature. In deoxygenated water Necturus periodically rises to the surface, opens its mouth, expands its buccal cavity to draw in fresh air, exhales air from the lungs, closes its mouth, and then compresses its buccal cavity and pumps air into the lungs. Thus Necturus produces only two buccal movements per breath: one expansion and one compression. Necturus shares the use of this two-stroke buccal pump with lungfishes, frogs and other salamanders. The ubiquitous use of this system by basal sarcopterygians is evidence that a two-stroke buccal pump is the primitive lung ventilation mechanism for sarcopterygian vertebrates. In contrast, basal actinopterygian fishes use a four-stroke buccal pump. In these fishes the buccal cavity expands to fill with expired air, compresses to expel the pulmonary air, expands to fill with fresh air, and then compresses for a second time to pump air into the lungs. Whether the sarcopterygian two-stroke buccal pump and the actinopterygian four-stroke buccal pump arose independently, whether both are derived from a single, primitive osteichthyian breathing mechanism, or whether one might be the primitive pattern and the other derived, cannot be determined. Although Necturus and lungfishes both use a two-stroke buccal pump, they differ in their expiration mechanics. Unlike a lungfish (Protopterus), Necturus exhales by contracting a portion of its hypaxial trunk musculature (the m. Iransversus abdominis) to increase pleuroperitoneal pressure. The occurrence of this same expiratory mechanism in amniotes is evidence that the use of hypaxial musculature for expiration, but not for inspiration, is a primitive tetrapod feature. From this observation we hypothesize that aspiration breathing may have evolved in two stages: initially, from pure buccal pumping to the use of trunk musculature for exhalation but not for inspiration (as in Necturus); and secondarily, to the use of trunk musculature for both exhalation and inhalation by costal aspiration (as in amniotes).     

Breathing with your belly: Abdominal exhalation,loco-ventilatory integration and size constraints on locomotion in small mammals

In mammals, diaphragmatic contractions control inhalation while contraction of some thoracic hypaxial muscles and the transversus abdominis muscle contribute to exhalation. Additional thoracic hypaxial muscles are recruited as accessory ventilatory muscles to improve inhalation and exhalation during locomotion. However, the contribution of abdominal hypaxial muscles to resting and locomotor ventilation is little understood in mammals and loco-ventilatory integration has not been studied in small basal mammals. We show for the first time that all of the abdominal hypaxial muscles actively contribute to both resting and locomotory ventilation in mammals but in a size-dependent manner. In large opossums (Didelphis), hypaxial muscles exhibit uniform mild tonus during resting ventilation (pressurizing the gut to aid in exhalation) and shift to phasic bursts of activity during each exhalation during locomotion. Smaller opossums (Monodelphis) actively exhale by firing the abdominal hypaxial muscles at ～10 Hz at both rest and at preferred locomotor speeds. Furthermore, the large opossums entrained ventilation to limb cycling as speed increased while the small opossums entrained limb cycling to the resting ventilation rate during locomotion. Differences in these species are related to size effects on the natural frequency of the ventilatory system and increasing resting ventilation rates at small size. Large mammals, with lower resting ventilation rates, can increase ventilatory rates during locomotion, while the high resting ventilation rates of small mammals limits their ability to increase ventilation rates during locomotion. We propose that increase in mammalian body size during the Cenozoic may have been an adaptation or exaptation to overcome size effects on ventilation frequency.     

Muscle fibers in the central nervous system of nemerteans: Spatial organization and functional role

The system of muscle fibers associated with the brain and lateral nerve cords is present in all major groups of enoplan nemerteans. Unfortunately, very little is known about the functional role and spatial arrangement of these muscles of the central nervous system. This article examines the architecture of the musculature of the central nervous system in two species of monostiliferous nemerteans (Emplectonema gracile and Tetrastemma cf. candidum) using phalloidin staining and confocal microscopy. The article also briefly discusses the body‐wall musculature and the muscles of the cephalic region. In both species, the lateral nerve cords possess two pairs of cardinal muscles that run the length of the nerve cords and pass through the ventral cerebral ganglia. A system of peripheral muscles forms a meshwork around the lateral nerve cords in E. gracile. The actin‐rich processes that ramify within the nerve cords in E. gracile (transverse fibers) might represent a separate population of glia‐like cells or sarcoplasmic projections of the peripheral muscles of the central nervous system. The lateral nerve cords in T. cf. candidum lack peripheral muscles but have muscles similar in their position and orientation to the transverse fibers. The musculature of the central nervous system is hypothesized to function as a support system for the lateral nerve cords and brain, preventing rupturing and herniation of the nervous tissue during locomotion. The occurrence of muscles of the central nervous system in nemerteans and other groups and their possible relevance in taxonomy are discussed. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Amphibious adaptations in a newly recognized amphibious fish: Terrestrial locomotion and the influences of body size and temperature

Amphibious animals are adapted for both aquatic and terrestrial habitats. The conflicting requirements for dual habitats are perhaps most pronounced in the air‐breathing fishes, which represent an intermediate stage between the totally aquatic habitat and terrestrial colonization. A key requirement for amphibious fishes is terrestrial locomotion. The different densities and compositions of air and water impose constraints for efficient terrestrial locomotion that differ from those required for aquatic locomotion. I investigated terrestrial locomotion in a small South African fish, Galaxias ‘nebula’, by exposing 60 individual fish to air in specially designed raceways and quantifying movement type and occurrence as a function of availability of water, fish size and environmental temperature. Nebula showed a sustained undulating form of terrestrial locomotion characteristic of amphibious fishes and also a transient ballistic locomotion (jumps) typical of fully aquatic species. Terrestrial movement was influenced by fish size, with medium‐sized fish undertaking more jumps towards water, and fewer jumps away from water, than their smaller or larger conspecifics. In contrast, axial undulation was mainly influenced by temperature. However, there was no consistent pattern in temperature effects presumably because temperature is just one of a suit of environmental factors that may affect terrestrial locomotion. Nebula's amphibious adaptations allow it to cope with the unpredictability inherent in its natural environment.     

Abdominal muscle and epipubic bone function during locomotion in Australian possums: Insights to basal mammalian conditions and eutherian‐like tendencies in Trichosurus

Mammals have four hypaxial muscle layers that wrap around the abdomen between the pelvis, ribcage, and spine. However, the marsupials have epipubic bones extending anteriorly into the ventral hypaxial layers with two additional muscles extending to the ventral midline and femur. Comparisons of South American marsupials to basal eutherians have shown that all of the abdominal hypaxials are active bilaterally in resting ventilation. However, during locomotion marsupials employ an asymmetrical pattern of activity as the hypaxial muscles form a crosscouplet linkage that uses the epipubic bone as a lever to provide long‐axis support of the body between diagonal limb couplets during each step. In basal eutherians, this system shifts off the femur and epipubic bones (which are lost) resulting in a shoulder to pelvis linkage associated with shifts in both the positions and activity patterns of the pectineus and rectus abdominis muscles during locomotion. In this study, we present data on hypaxial function in two species (Pseudocheirus peregrinus and Trichosurus vulpecula) representing the two major radiations of possums in Australia: the Pseudocheiridae (within the Petauroidea) and the Phalangeridae. Patterns of gait, motor activity, and morphology in these two Australian species were compared with previous work to examine the generality of 1) the crosscouplet lever system as the basal condition for the Marsupialia and 2) several traits hypothesized to be common to all mammals (hypaxial tonus during resting ventilation, ventilation to step synchrony during locomotion, and bilateral transversus abdominis activity during locomotor expiration). Our results validate the presence of the crosscouplet pattern and basic epipubic bone lever system in Australian possums and confirm the generality of basal mammalian patterns. However, several novelties discovered in Trichosurus, reveal that it exhibits an evolutionary transition to intermediate eutherian‐like morphological and motor patterns paralleling many other unique features of this species. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Osteoderm microstructure indicates the presence of a crocodylian‐like trunk bracing system in a group of armoured basal tetrapods

Buchwitz, M., Witzmann, F., Voigt, S. and Golubev, V. 2012. Osteoderm microstructure indicates the presence of a crocodylian‐like trunk bracing system in a group of armoured basal tetrapods. —Acta Zoologica (Stockholm) 93 : 260–280. The microstructure of dorsal osteoderms referred to the chroniosuchid taxa Chroniosuchus, Chroniosaurus, Madygenerpeton and cf. Uralerpeton is compared to existing data on the bystrowianid chroniosuchian Bystrowiella and further tetrapods. Chroniosuchid osteoderms are marked by thin internal and relatively thick external cortices that consist of lowly vascularised parallel‐fibred bone. They are structured by growth marks and, in case of Madygenerpeton, by lines of arrested growth. The cancellous middle region is marked by a high degree of remodelling and a primary bone matrix of parallel‐fibred bone that may include domains of interwoven structural fibres. Whereas the convergence of Bystrowiella and chroniosuchid osteoderms is not confirmed by our observations, the internal cortex of the latter displays a significant peculiarity: It contains distinct bundles of shallowly dipping Sharpey’s fibres with a cranio‐ or caudoventral orientation. We interpret this feature as indicative for the attachment of epaxial muscles which spanned several vertebral segments between the medioventral surface of the osteoderms and the transversal processes of the thoracic vertebrae. This finding endorses the hypothesis that the chroniosuchid osteoderm series was part of a crocodylian‐like trunk bracing system that supported terrestrial locomotion. According to the measured range of osteoderm bone compactness, some chroniosuchian species may have had a more aquatic lifestyle than others.     

Bird terrestrial locomotion as revealed by 3D kinematics

Most birds use at least two modes of locomotion: flying and walking (terrestrial locomotion). Whereas the wings and tail are used for flying, the legs are mainly used for walking. The role of other body segments remains, however, poorly understood. In this study, we examine the kinematics of the head, the trunk, and the legs during terrestrial locomotion in the quail (Coturnix coturnix). Despite the trunk representing about 70% of the total body mass, its function in locomotion has received little scientific interest to date. This prompted us to focus on its role in terrestrial locomotion. We used high-speed video fluoroscopic recordings of quails walking at voluntary speeds on a trackway. Dorso-ventral and lateral views of the motion of the skeletal elements were recorded successively and reconstructed in three dimensions using a novel method based on the temporal synchronisation of both views. An analysis of the trajectories of the body parts and their coordination showed that the trunk plays an important role during walking. Moreover, two sub-systems participate in the gait kinematics: (i) the integrated 3D motion of the trunk and thighs allows for the adjustment of the path of the centre of mass; (ii) the motion of distal limbs transforms the alternating forward motion of the feet into a continuous forward motion at the knee and thus assures propulsion. Finally, head bobbing appears qualitatively synchronised to the movements of the trunk. An important role for the thigh muscles in generating the 3D motion of the trunk is suggested by an analysis of the pelvic anatomy.     

Description and scaling of pectoral muscles in ictalurid catfishes

The pectoral spine of catfishes is an antipredator adaptation that can be bound, locked, and rubbed against the cleithrum to produce stridulation sounds. We describe muscle morphology of the pectoral spines and rays in six species in four genera of North American ictalurid catfishes. Since homologies of catfish pectoral muscles have not been universally accepted, we designate them functionally as the spine abductor and adductor and the arrector dorsalis and ventralis. The four muscles of the remaining pectoral rays are the superficial and deep (profundal) abductors and adductors. The large spine abductor and spine adductor are responsible for large amplitude movements, and the smaller arrector dorsalis and arrector ventralis have more specialized functions, that is, spine elevation and depression, respectively, although they also contribute to spine abduction. Three of the four spine muscles were pennate (the abductor and two arrectors), the spine adductor can be pennate or parallel, and ray muscles have parallel fibers. Insertions of pectoral muscles are similar across species, but there is a shift of origins in some muscles, particularly of the superficial abductor of the pectoral rays, which assumes a midline position in Ictalurus and increasingly more lateral placement in Ameiurus (one quarter way out from the midline), and Pylodictis and Noturus (half way out). Coincident with this lateral shift, the attachments of the hypaxial muscle to the ventral girdle become more robust. Comparison with its sister group supports the midline position as basal and lateral migration as derived. The muscles of the pectoral spine are heavier than muscles of the remaining rays in all species but the flathead, supporting the importance of specialized spine functions above typical movement. Further, spine muscles were larger than ray muscles in all species but the flathead catfish, which lives in water with the fastest currents. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

The interplay between speed,kinetics, and hand postures during primate terrestrial locomotion

Nonprimate terrestrial mammals may use digitigrade postures to help moderate distal limb joint moments and metapodial stresses that may arise during high‐speed locomotion with high‐ground reaction forces (GRF). This study evaluates the relationships between speed, GRFs, and distal forelimb kinematics in order to evaluate if primates also adopt digitigrade hand postures during terrestrial locomotion for these same reasons. Three cercopithecine monkey species (Papio anubis, Macaca mulatta, Erythrocebus patas) were videotaped moving unrestrained along a horizontal runway instrumented with a force platform. Three‐dimensional forelimb kinematics and GRFs were measured when the vertical force component reached its peak. Hand posture was measured as the angle between the metacarpal segment and the ground (MGA). As predicted, digitigrade hand postures (larger MGA) are associated with shorter GRF moment arms and lower wrist joint moments. Contrary to expectations, individuals used more palmigrade‐like (i.e. less digitigrade) hand postures (smaller MGA) when the forelimb was subjected to higher forces (at faster speeds) resulting in potentially larger wrist joint moments. Accordingly, these primates may not use their ability to alter their hand postures to reduce rising joint moments at faster speeds. Digitigrady at slow speeds may improve the mechanical advantage of antigravity muscles crossing the wrist joint. At faster speeds, greater palmigrady is likely caused by joint collapse, but this posture may be suited to distribute higher GRFs over a larger surface area to lower stresses throughout the hand. Thus, a digitigrade hand posture is not a cursorial (i.e. high speed) adaptation in primates and differs from that of other mammals. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Regulation of hypaxial muscle development

The majority of skeletal muscles in higher vertebrates are hypaxial and stem from the lateral lip of the dermomyotomes. Various external signals converge on the dorsolateral quadrant of the somite to specify the hypaxial muscle precursors, to discriminate between migratory and non-migratory cells and to allow delamination of precursors destined for long-range migration. Within the somite, Pax3 acts as upstream regulator of hypaxial muscle development. Downstream targets are cMet and Lbx1, which may independently control identity, differentiation and motility of migratory muscle precursors. Received: 31 August 1998 / Accepted: 20 October 1998     

Axial allometry in a neutrally buoyant environment: effects of the terrestrial‐aquatic transition on vertebral scaling

Ecological diversification into new environments presents new mechanical challenges for locomotion. An extreme example of this is the transition from a terrestrial to an aquatic lifestyle. Here, we examine the implications of life in a neutrally buoyant environment on adaptations of the axial skeleton to evolutionary increases in body size. On land, mammals must use their thoracolumbar vertebral column for body support against gravity and thus exhibit increasing stabilization of the trunk as body size increases. Conversely, in water, the role of the axial skeleton in body support is reduced, and, in aquatic mammals, the vertebral column functions primarily in locomotion. Therefore, we hypothesize that the allometric stabilization associated with increasing body size in terrestrial mammals will be minimized in secondarily aquatic mammals. We test this by comparing the scaling exponent (slope) of vertebral measures from 57 terrestrial species (23 felids, 34 bovids) to 23 semi‐aquatic species (pinnipeds), using phylogenetically corrected regressions. Terrestrial taxa meet predictions of allometric stabilization, with posterior vertebral column (lumbar region) shortening, increased vertebral height compared to width, and shorter, more disc‐shaped centra. In contrast, pinniped vertebral proportions (e.g. length, width, height) scale with isometry, and in some cases, centra even become more spool‐shaped with increasing size, suggesting increased flexibility. Our results demonstrate that evolution of a secondarily aquatic lifestyle has modified the mechanical constraints associated with evolutionary increases in body size, relative to terrestrial taxa.